The Functions of Sound Production in the Lined Seahorse, Hippocampus Erectus, and Effects of Loud Ambient Noise on Its Behavior And
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THE FUNCTIONS OF SOUND PRODUCTION IN THE LINED SEAHORSE, HIPPOCAMPUS ERECTUS, AND EFFECTS OF LOUD AMBIENT NOISE ON ITS BEHAVIOR AND PHYSIOLOGY IN CAPTIVE ENVIRONMENTS By PAUL AUGUST ANDERSON A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2009 1 © 2009 Paul August Anderson 2 To my parents, Josephine J. McLaughlin and Paul F. Anderson, whose unconditional love and support have paved the way for me to be everything that I want to be 3 ACKNOWLEDGMENTS First and foremost, I owe a debt of gratitude to my Ph.D. advisor, W.J. Lindberg (University of Florida/UF), who accepted the daunting role of advising a student with complex, multidisciplinary research interests. Among the many contributions made to the project, Dr. Lindberg provided wisdom and teachings in the scientific process, helped guide complex negotiations among many stakeholders involved, offered funding via the D.M. Smith Fellowship, and assisted in the acquisition of additional funds along the way. Dr. Lindberg provided enlightened guidance all along the route of this circuitous adventure to its destination! Similarly, I offer many thanks to each of my Ph.D. committee members, I. Berzins (The Florida Aquarium) , H. Masonjones (University of Tampa), D. Murie (UF), D. Parkyn (UF), C. St. Mary (UF), each of whom played important roles in shaping the project into its final form. In particular, I thank I. Berzins for offering a fruitful and collaborative relationship with The Florida Aquarium Center for Conservation, and for generating ideas about the effects of noise on stress in fishes (that led to co-authorship in Chapter 4). I. Berzins also provided funding to conduct The Seahorse Sound Survey (Chapter 2) and has created a position for me at The Center for Conservation that has to date enabled me to fulfill mutual career goals; it is an opportunity that I truly cherish and enjoy. The concept of the Dissertation was developed in consult with H. Masonjones, who first suggested effects of noise on seahorse health and reproduction, having previously detected effects in her study system (H. zosterae) and suggesting further exploration of the phenomenon. H. Masonjones also captained the vessel (and furnished a crew of competent shipmates, S., G., and K. Masonjones) upon which ambient noise measurements were taken from the field. D. Murie provided thorough critique of study design and also suggested the study of acoustic roles in prey capture behavior, that led to Chapter 5 of this Dissertation. D. Parkyn offered wisdom and teachings in the complex world of acoustic neuroethology, and was 4 instrumental in securing stipend funding along the way. Finally, C. St. Mary also offered valuable critique of study design, assisted with fund acquisition, and advised statistical methods in Chapter 4. In the company of these mentors, I also wish to acknowledge R. Francis-Floyd, who opened the door for me at The University of Florida and helped me to get started on my path. Some Chapters in this Dissertation are the result of fruitful collaborations with co-authors, who provided knowledge and resources instrumental to the work. I am indebted to D. Mann (University of South Florida), who provided the analytical system necessary to conduct the study in Chapter 3, along with patient teachings in the methods of AEP audiometry and sound analysis in general. I also acknowledge his students, B. Casper, M. Hill-Cook, R. Hill, and J. Locascio, who provided guidance along the way. L. J. Guillette and H. Hamlin (UF) taught enzyme immunoassay methods and offered their laboratory to conduct cortisol assays included in Chapter 4. They also provided valuable advice in considering the stress phenomenon, advising the examination of variance among response measures, that led to a fruitful examination of results. I also acknowledge members of their lab, in particular T. Edwards and B. Moore, for guidance. E. Adams and F. Fogarty (UF) served as undergraduate research assistants along the way. Among the many (acknowledged below), these individuals invested tremendous effort and original thought over an extended period of time that contributed greatly to Chapters 6 and 4 (respectively). I am very thankful for their sustained hard work. The Dissertation was further molded by fruitful conversations with, and wisdom contributed by, professionals in diverse fields. A. Slater (The Florida Aquarium) molded my skills in syngnathid husbandry. I owe many thanks to J. Pattee (Pioneer Hill Software) and R. Shrivastav (UF), who taught concepts of acoustics and sound analysis. A. Noxon (Acoustic 5 Sciences Corporation) shared his knowledge in acoustic engineering that shaped soundproofing methods that were used in the methods of Chapters 4-6. The muting surgery that was employed in Chapters 5 and 6 was first developed and advised by D. Colson (Rhode Island ENT Physicians, Inc.), who first published a study using the muting method. K. Harr (UF) advised hematological methods used in Chapter 4. A. Dove (Georgia Aquarium) and E. Greiner (UF) assisted with parasite identification in Chapter 4. L. Farina (UF) guided histological interpretation. Statistical analyses employed in the Chapters were selected and shaped as a result of consultation with several people in addition to those previously mentioned, including M. Allen, M. Brennan, J. Colee, D. Dutterer, J. Hill, and R. Littell (UF). Several people provided technical guidance and offered access to technical resources that made elements of methodology possible. I am especially grateful to D. Petty (UF) and her laboratory (including J. Holloway and T. Crosby), who provided invaluable veterinary services all along the way to maintain the health of the research collection. They also participated in, and helped shape the methodology for, the physiological methods used in Chapter 4 and the muting surgery used in Chapters 5 and 6. D. Samuelson and P. Lewis (UF) taught histological techniques and provided access to their laboratory for histological processing and evaluation. H. Rutherford (The Pier Aquarium) loaned hydrophones that were used extensively for sound recording. I appreciate the generous contributions of people in the fishing and marine ornamental industries that provided animals to study as well as resources to care for them. I owe many thanks to M. Helmholtz (Above The Reef), R. Stevens (Mary Jane Shrimp Co.), and the crew of The Twin Rivers Marina, who believed in the project and donated many animals to build the research collection. I am grateful to A. and M. Maxwell and the crew of Sea Critters, Inc., who 6 provided a live food supply for the collection over a period of several years. R. Lewis (Aquatic Indicators) provided Mysidopsis bahia for the experiment described in Chapter 5. I enjoyed the opportunity to engage undergraduate students of the University of Florida in research experience. In return, this team provided a powerful workforce, ensuring proper care of the research collection as well as the collection of quality data that are at the base of the results reported here. These people are E. Adams, F. Bastos, J. Bound, F. Fogarty, S. Koka, J. Liu, B. Macke, A. Maness, J. Moriarty, K. Nuessly, S. Osborn, Z. Punjani, J. Rosenbaum, B. Slossberg, D. Snipelsky, and A. Weppelman. Nine public aquaria from throughout the United States participated in The Seahorse Sound Survey (Chapter 2); for their contributions I am grateful. Thanks to R. Doege (Dallas World Aquarium), R. Curttright (Kingdom of the Seas Aquarium), K. Dobson (Maritime Aquarium), J. Reynolds and S. Reiner (Moody Gardens), S. Spina (New England Aquarium), J. Moffatt (Pittsburgh Aquarium), J. Skoy (Ripley’s Aquarium of the Smokies), J. Rawlings (Riverbanks Zoo), and K. Alford (Tennessee Aquarium). I also acknowledge N. Dunham (Florida Fish and Wildlife Conservation Commission), who graciously offered data and maps on sites in Tampa Bay where seahorses have been collected, and R. Watkins (UF), who provided Figure 2-2. Many funding sources contributed to the success of this Dissertation. As previously mentioned, the D.M. Smith Fellowship funded laboratory construction and research operations, and The Florida Aquarium Center for Conservation funded The Seahorse Sound Survey. The Mulvihill Scholarship (granted by Aquaculture Research/Environmental Associates, Inc. and the United States Aquaculture Association), Project A.W.A.R.E., and the Twin Rivers Marina also funded operations. The American Association of Zoo Veterinarians’ Mazuri Fund fully sponsored Chapter 4. My scholarship and stipend was provided by the University of Florida’s 7 Alumni Fellowship, the Morris Animal Foundation, The Spurlino Foundation, and The Florida Aquarium Center for Conservation. Animal collection was authorized by the Florida Fish and Wildlife Conservation Commission Special Activities License #05SR-944. Husbandry and experimental protocols were authorized by the University of Florida IACUC Protocol #D-432, the University of South Florida IACUC Protocol #2118, and The Florida Aquarium Animal Care and Use Committee. 8 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES .........................................................................................................................13 LIST OF FIGURES .......................................................................................................................14