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Influence of Environmental Features on Tubifex Tubifex And INFLUENCE OF ENVIRONMENTAL FEATURES ON TUBIFEX TUBIFEX AND MYXOBOLUS CEREBRALIS INFECTED TUBIFEX TUBIFEX IN YELLOWSTONE NATIONAL PARK: IMPLICATIONS FOR WHIRLING DISEASE RISK by Julie Diane Alexander A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biological Sciences MONTANA STATE UNIVERSITY Bozeman, Montana November 2010 ©COPYRIGHT by Julie Diane Alexander 2010 All Rights Reserved ii APPROVAL of a dissertation submitted by Julie Diane Alexander This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citation, bibliographic style, and consistency and is ready for submission to the Division of Graduate Education. Dr. Billie L. Kerans, Committee Chair Approved for the Department of Ecology Dr. David W. Roberts, Department Head Approved for the Division of Graduate Education Dr. Carl A. Fox iii STATEMENT OF PERMISSION TO USE In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying of this dissertation is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to ProQuest Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted “the exclusive right to reproduce and distribute my dissertation in and from microform along with the non- exclusive right to reproduce and distribute my abstract in any format in whole or in part.” Julie Diane Alexander November 2010 iv ACKNOWLEDGEMENTS Funding was provided by the Whirling Disease Initiative of the National Partnership on Management of Wild and Native Coldwater Fisheries and the Center for Resources, Yellowstone National Park and an ESPSCOR grant to BLK. Additional support was provided by the Western Fisheries Research Center (WFRC), USGS. I thank employees of the Bozeman Fish Technology Center for histology and Yellowstone‟s Center for Resources for field assistance. I thank Charlotte Rasmussen at the WFRC for instruction in molecular techniques used in my research. She helped turn an endeavor that was fraught with frustration into a great learning experience. Dr. Matt Lavin generously provided advice regarding phylogenetic analyses. I thank my committee members Dr. Todd Koel, Dr. Jim Winton, Dr. Al Zale, and Dr. Steven Kalinowski for generously providing much needed critical input and their time. Dr. Sandra Halonen, as graduate representative, also provided time, input, and gender balance in the room. I especially thank Dr. Billie Kerans for all of her time and input throughout this process. Her ideas, frustrations, humor, and above all, tenacity, have shaped my work and fostered enjoyment in trying to understand how the pieces fit together. Fellow graduate students and members of Billie‟s lab (past and present) contributed ideas, listened to my grumblings, and provided moral support. I thank all of my technicians, friends, and family members who walked many miles with me in grizzly country to retrieve sentinel fish and T. tubifex. Finally, I acknowledge and thank my family, Bill, Margie, and Michael Alexander, and Frank Tazzara, for their unwavering support over the years. I could not have finished without their encouragement and help. v TABLE OF CONTENTS 1. INTRODUCTION TO DISSERTATION .............................................................. 1 Overview of Dissertation ........................................................................................ 4 Literature Cited ....................................................................................................... 6 2. BACKGROUND .................................................................................................. 14 The Parasite ........................................................................................................... 15 Factors Affecting Myxobolus cerebralis Spores ................................................... 17 Whirling Disease Risk .......................................................................................... 18 The Salmonid Host ............................................................................................... 21 Factors Affecting Myxobolus cerebralis infections in the Salmonid Host .... 22 The Oligochaete Host ........................................................................................... 23 Myxobolus cerebralis Infections in the Oligochaete Host ............................. 24 Host Factors ................................................................................................... 24 Community Factors ....................................................................................... 25 Environmental Factors ................................................................................... 26 Whirling Disease and Yellowstone Cutthroat Trout ............................................. 28 Literature Cited ..................................................................................................... 30 3. CONTEXT SPECIFIC PARASITISM IN TUBIFEX TUBIFEX IN GEOTHERMALLY INFLUENCED STREAM REACHES IN YELLOWSTONE NATIONAL PARK ............................................................... 40 Contribution of Authors and Co-Authors ............................................................. 40 Manuscript Information Page ............................................................................... 41 Abstract ................................................................................................................. 42 Introduction ........................................................................................................... 44 Methods ................................................................................................................ 48 Study Site and Environmental Features ......................................................... 48 Host Ecology ................................................................................................. 50 Parasite Ecology ............................................................................................ 54 Whirling Disease Risk to Fish ....................................................................... 55 Analyses ......................................................................................................... 56 Results ................................................................................................................... 57 Environmental Features ................................................................................. 57 Host Ecology ................................................................................................. 57 Parasite Ecology ............................................................................................ 59 Whirling Disease Risk to Fish ....................................................................... 59 Discussion ............................................................................................................. 60 Acknowledgements ............................................................................................... 66 vi TABLE OF CONTENTS-CONTINUED Literature Cited ..................................................................................................... 67 4. HABITAT AND PARASITE SUCCESS: INFLUENCE OF HOST ENVIRONMENT ON MYXOBOLUS CEREBRALIS IN TRIBUTARIES IN YELLOWSTONE NATIONAL PARK ............................................................... 83 Abstract ................................................................................................................. 83 Introduction ........................................................................................................... 85 Methods ................................................................................................................ 87 Study Site and Environmental Features ......................................................... 88 Invertebrate Community ................................................................................ 91 Host Ecology ................................................................................................. 93 Parasite Ecology ............................................................................................ 95 Confinement and Invertebrates ...................................................................... 97 Confinement Types and Host Ecology .......................................................... 98 Confinement Types and Parasite Ecology ..................................................... 98 Results ................................................................................................................... 99 Environmental Features ................................................................................. 99 Invertebrate Ecology ...................................................................................... 99 Oligochaete Host Ecology ........................................................................... 100 Parasite Ecology .......................................................................................... 100 Discussion ........................................................................................................... 101
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