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RESILIENCE AND ACCLIMATIZATION POTENTIAL OF REEF CORALS UNDER PREDICTED CLIMATE CHANGE STRESSORS A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI‘I MĀNOA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY DECEMBER 2012 By Hollie M. Putnam Dissertation Committee: Ruth D. Gates, Chairperson H Gert de Couet Guylaine Poisson Robert J. Toonen Eric H. DeCarlo ACKNOWLEDGEMENTS My sincere thanks go out to my amazing advisor, Dr. Ruth Gates. You were always there to challenge, encourage, and support me. No matter how large the problems seemed, after a meeting with you, I felt I could conquer the world. Thank you for all the time you have taken to teach and mentor me, and I look forward to continuing our work together! To my committee, Dr. Eric DeCarlo, Dr. Gert de Couet, Dr. Guylaine Poisson, and Dr. Rob Toonen, thank you for your support, your flexibility, your feedback, your time, and the open doors whenever I had questions. Last but certainly not least, I am thankful for my unofficial committee member from afar, Dr. Peter Edmunds. Thank you for everything over the past 7 years! You are an outstanding scientist and role model. I cannot thank you and Ruth enough for pushing me and supporting me every step of the way! I want to say a special thanks to the Gates lab members past and present! Dr. Michael Stat, Dr. Xavier Pochon and Dr. Michelle Phillips, you have been great teachers and friends! I appreciate all the time you gave me and the many exciting discussions along the way. Dr. Denise Yost and Marisa Guarinello, thank you for your friendship and the late nights on the kayak and in the lab during spawning. You made a difficult job so much easier with your help! To all the Gates lab members and Leong lab members (Laetitia, Anderson, Derek, Rebecca, Maggie S, Nicole, John, Jen, Mindy, Madison, Keisha, Kirsten, Carly, Kelsey and Dean), thank you all for an intellectually stimulating and fun time for the last 4 years! I want to say a huge thanks to Madison Kosma, Keisha Rodriguez, Kirsten Fujitani, and Carly Richer. Thank you so much for the many hours and fun times in the wetlab and at the titrator! You ladies are amazing! To all my friends ii (especially Maggie J, Viv, Michelle, Megan, Nyssa, Sherril & Amy), and the HIMB faculty and staff, thank you! To my family in Minnesota and Vermont, I want to thank you for your support in my extended time in school. Mom and Dad, Autumn and Chuck, Bill and Donna, Melanie and Bryan, you have all been patient, accepting, and encouraging. Thank you all! Finally, I am extremely grateful for the love and encouragement of my husband Travis. We got married my first month in grad school 7 years ago and you have been a constant support since then. I feel like this degree is as much yours as it is mine, and I thank you so much for everything you have done for me and that you continue to do for us while you are gone. I love you! To anyone I may have missed, the list could go on and on, so thank you all! This research was funded or supported by grants from UH EPSCoR (EPS- 0903833), NSF to RDG (OCE-0752604), NSF to PJ Edmunds (BIO-OCE 08-44785), the National Marine Sanctuary Program and HIMB Reserve Partnership (memorandum of agreement 2005-008/66882) and funding to HMP from the International Society for Reef Studies, the Ocean Conservancy, American Fisheries Society, Lerner Gray Fund for Marine Research, the Achievement Rewards for College Scientists (ARCS) Foundation, and the UH Edmondson Research Fund. In addition, this research was developed under STAR Fellowship Assistance Agreement no. FP917199 awarded by the U.S. Environmental Protection Agency (EPA). This manuscript has not been formally reviewed by the EPA and the views expressed are solely those of the authors. The EPA does not endorse any products or commercial services mentioned in this document. iii ABSTRACT Coral reef ecosystems are among the most diverse and productive in the world. The basis for this productivity is the symbiosis between cnidarian hosts and single-celled dinoflagellates (Symbiodinium) that together structure the reef. Coral reefs are currently under threat locally from a variety of stressors, as well as globally from increasing temperature and CO2-induced ocean acidification. While rates of adaptation are anticipated to be slower than the rate of climate change, rapid acclimatory processes, such as trans-generational acclimatization and other epigenetic mechanisms may contribute to the maintenance of coral reefs in the future. The goal of this dissertation was to advance our understanding of the processes involved in coral response to climate change. In particular, I focused on identifying the variability in physical setting (temperature, pH, pCO2) for coastal reefs in Kaneohe Bay, Hawaii, and developing the experimental infrastructure with which to test the effects of increasing temperature and ocean acidification on the reef building coral Pocillopora damicornis. This brooding coral provides the ideal model to test life-stage specific response and the connection between adults and brooded larvae in a trans-generational context. In a 9-day factorial experimental exposure to either ambient (25°C) and high temperature (29°C) and ambient (~415 µatm) and high CO2 (~635 µatm), P. damicornis larvae displayed strong metabolic suppression and decline in Rubiso protein expression (ribulose-1,5-bisphosphate carboxylase/oxygenase, a rate-limiting enzyme in the Calvin cycle) at high temperatures regardless of CO2 concentration, likely resulting in energetic debt with negative fitness implications. When adult corals were preconditioned to ambient (26.5°C and ~ 415 µatm) or high temperature and CO2 (29°C and ~800 µatm) for 1.5 months prior to larval iv release, adults in high conditions displayed significant declines in productivity while maintaining metabolic rate and calcification, with reproductive consequences. Coral larvae from adults exposed to high conditions (29°C and ~800µatm) were significantly affected by adult environment resulting in smaller larvae with lower metabolic rates. However, positive trans-generational acclimatization was documented in a secondary reciprocal exposure in larvae from adults with a high history displaying higher size- normalized metabolic rates, with implications for protein turnover, energetics, and fitness. This work highlights the necessity of considering rapid acclimatization, or epigenetic processes (i.e., trans-generational acclimatization) in our examination of the response of coral to climate change in order to best inform our predictions for the future of coral reefs. v TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................................................................................... ii ABSTRACT ...................................................................................................................... iii LIST OF TABLES AND APPENDICES ......................................................................... vii LIST OF FIGURES ......................................................................................................... viii CHAPTER ONE .................................................................................................................. 1 CHAPTER TWO ............................................................................................................... 34 Abstract ......................................................................................................................... 35 Introduction ................................................................................................................... 36 Materials and Methods ................................................................................................... 38 Results ........................................................................................................................... 45 Discussion ...................................................................................................................... 47 Acknowledgements ....................................................................................................... 52 References ..................................................................................................................... 54 CHAPTER THREE ........................................................................................................... 66 Abstract ......................................................................................................................... 67 Introduction ................................................................................................................... 68 Materials and Methods ................................................................................................... 72 Results ........................................................................................................................... 80 Discussion ...................................................................................................................... 83 Acknowledgements ....................................................................................................... 93 References ..................................................................................................................... 94 vi TABLE OF CONTENTS (continued) CHAPTER FOUR ........................................................................................................... 114 Abstract ......................................................................................................................