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California State University, Northridge the Effects Of CALIFORNIA STATE UNIVERSITY, NORTHRIDGE THE EFFECTS OF OCEAN ACIDIFICATION ON BIOEROSION IN THE BACK REEF OF MOOREA, FRENCH POLYNESIA A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biology By Lauren Michele Valentino August 2014 The thesis of Lauren M. Valentino is approved: ___________________________________________ __________________ Peter J. Edmunds, Ph.D. Date ___________________________________________ __________________ Mark A. Steele, Ph.D. Date ___________________________________________ __________________ Robert C. Carpenter, Ph.D., Chair Date California State University, Northridge ii ACKNOWLEDGEMENTS I would like to thank my advisor Dr. Robert Carpenter for believing in me and providing opportunities that will continue to foster my abilities as a student, writer, and scientist. Dr. Carpenter’s guidance and unwavering patience helped me become a more confident and independent researcher. I will never forget all of the delicious dinners, boat lunches, and round-island adventures surveying the reefs of Moorea. I truly can’t thank him enough for making me an additional member of the Carpenter lab family, something that I will carry proudly as I continue my career in marine science. I am grateful to Dr. Peter Edmunds who met with me countless times to help me work through many different questions I had regarding my thesis. His attention to detail and ability to streamline the most complicated processes always left me with a deeper understanding and enthusiasm for my research. Any self-doubt was immediately met with the mantra, “You’re good enough. You’re smart enough. And doggone it, people like you.” This and many other Pete-isms will help remind me of the importance of my confidence as a scientist (and a good coffee buzz) throughout my research and life. I would also like to thank Dr. Mark Steele for being an incredible teacher and committee member. He always made me feel comfortable being myself, while still taking my research and me seriously. I used to think that stats were abstract and arduous, but Dr. Steele has a way of explaining it in a simple, straightforward way that makes it seem so practical and useful. His paper discussions and pragmatic approach to marine research helped me look at science with a more critical eye. I feel very lucky to have learned from the diverse perspectives of Bob, Pete, and Mark who have all made me the scientist I am today, and for that I am forever thankful. Also, thanks to my Mom, Lindsay, and Johnnie for being patient and understanding throughout my grad school experience. Going home always helped put things in perspective and reminded me that where I am from is as important as where I am going. Thanks to Heather Hillard, the ‘eather in Leather, who was my lab partner in crime beginning to end. I have her to thank for getting me through what I once thought was not possible. Merci bien to the rest of the Carpenter lab, especially Amy, Carolina, Vinny, Anya, Coulson, Steeve, and Stella who were always positive, helpful, and improved my quality of life both on campus and in the field. For all their love and support, thanks to my adoptive LA family, Sylvia (Stout), Cecilia (Harley), and Lauren M. for giving me a home away from home. I am so grateful to my friends Camdilla, Sara, Lareen, and Mark. The countless dinners, adventures, and work parties I shared with them helped me though the most challenging times, kept me sane, and most importantly made me laugh. This research was made possible with funding from the US National Science Foundation to the Moorea Coral Reef Long Term Ecological Research MCR-LTER (1026851 and 1236905), R. Carpenter and P. Edmunds (1041270), and from generous donations made by the Gordon and Betty Moore Foundation. Additionally, this research was supported by California State University Northridge Graduate Studies, Associated Students, California State University’s Thesis Support Grant, and California State University’s Gradate Equity Fellowship. iii TABLE OF CONTENTS Signature Page ii Acknowledgements iii List of Tables v List of Figures vi Abstract vii Chapter 1 General Introduction 1 Chapter 2 Distribution and Abundance of Internal and External Bioeroders Across the Back Reef of Moorea, French Polynesia Introduction 9 Methods 15 Results 19 Discussion 21 Tables 26 Figures 30 Chapter 3 Effects of Ocean Acidification on Calcification, Bioerosion, and Respiration Rates of Lithophaga laevigata within Massive Porites in Moorea, French Polynesia Introduction 35 Methods 40 Results 47 Discussion 50 Tables 56 Figures 59 Chapter 4 Summary 64 Literature Cited 69 iv LIST OF TABLES PAGE 2.1 - Results from a two-factor analysis of variance (ANOVA) testing for 26 differences in external bioeroder densities upstream and downstream. 2.2 - Results from a two-factor analysis of variance (ANOVA) testing for 27 differences in internal bioerosion on bommie and rubble substrate at upstream and downstream locations. 2.3 - Results from the mixed model analysis of variance (ANOVA) testing for 28 differences in Lithophaga abundance at upstream, top, downstream locations on a Porites colony. 2.4 - Results from Tukey’s honestly significant difference tests of the mixed 29 model analysis of variance (ANOVA) testing for differences in Lithophaga abundance at upstream, top, downstream locations on a Porites colony. 3.1 - Results from the simple linear regression of the major axis of the borehole 56 and the length of the valve (n=38). 3.2 - Summary of carbonate chemistry measurements in 6 randomly assigned 57 tanks throughout a 28-d incubation period in elevated and ambient pCO2 treatments. Mean ± SE. N=28 sampling days for all parameters. 3.3 - Results from the mixed model analysis of variance (ANOVA) testing 58 effects of Core Type (Porites with and without Lithophaga) and CO2 (ambient and elevated treatments) on area-normalized net calcification rates. v LIST OF FIGURES PAGE 2.1 - Study site on the north shore of Moorea, French Polynesia. 30 Upstream and downstream locations shown for transects conducted on external bioeroders, internal erosion, and Lithophaga abundance. Arrows depict the unidirectional flow across the back reef. 2.2 - External bioeroder abundance on bommies and rubble in upstream 31 and downstream locations. Location P=0.214, Substrate type P=<0.001, L x S P=0.636. 2.3 - Percent bioerosion in bommie and rubble pieces collected at 32 upstream and downstream locations. Location P=0.337, Substrate type P=<0.001, L x S P=0.010. 2.4 - Lithophaga abundance in upstream and downstream locations. 33 Location P=0.912. 2.5 - Spatial Abundance of Lithophaga on Porites on upstream, top, and 34 downstream sides of a Porites colony. Zone P=0.002, Upstream P=0.001, Top P=0.008, Downstream P=0.002, (n=10). 3.1 - Correlation between the major axis of the borehole opening of 59 Lithophaga laevigata and valve length (n=38). 3.2 - Area-normalized net calcification rates of Porites with and without 60 Lithophaga in ambient and elevated treatments (n=28). 3.3 - Weight-normalized net calcification rates of Lithophaga in burrow 61 mimics in ambient and elevated treatments (n=20). 3.4 - Estimated bioerosion rates of Lithophaga in Porites in elevated and 62 ambient pCO treatments (n=28). 2 3.5 - Respiration rates of Lithophaga in burrow mimics after 28 days in 63 elevated and ambient pCO treatments (n=12). 2 vi ABSTRACT Effects of ocean acidification on bioerosion in the back reef of Moorea, French Polynesia. By Lauren M. Valentino Master of Science in Biology Coral reefs are among the most diverse ecosystems on the planet and have been compared to rainforests because of their complexity and high species diversity. Tropical reefs have relatively nutrient-poor waters, but they are one of the most productive ecosystems providing benefits and ecosystem services to society in the form of coastal protection, food, and economic resources such as tourism. Rising carbon dioxide emissions by humans will have serious environmental implications for the ocean environment. Coral reef ecosystems are particularly vulnerable to this unprecedented increase of CO2 due to their carbon chemistry and thermal sensitivity. Anthropogenic CO2 is predicted to decrease ocean surface pH by 0.14–0.35 units by 2100 causing ocean acidification (OA). Most studies have focused on how OA will affect rates of calcification of coral reef organisms. However, bioerosion also could be sensitive to rapid changes in ocean carbonate chemistry. I tested the effects of decreased pH on the distribution of bioeroders in the field and on the boring capacity of the mollusk Lithophaga laevigata living within corals, massive Porites spp. (a complex of three species: P. lobata, P. australiensis, and P. lutea) in the lab. Field studies showed higher vii external bioeroder abundance on coral bommies, and higher internal bioerosion in coral rubble, however, there was no differences in bioerosion between variable pH environments found at upstream and downstream transects. L. laevigata, a boring bivalve, is abundant within massive Porites sp. on the back reef of Moorea, French Polynesia. L. laevigata abundance in massive Porites across the back reef ranged from 3 to 95 ind/m2. Size analysis of L. laevigata showed a significant correlation of the borehole opening and the size of the bivalve, which allowed for a non-destructive method for collection of uniformly sized bivalves as a way to standardize bioerosion rates for analyses. I conducted a month-long mesocosm experiment where massive Porites cores with and without L. laevigata, were incubated in ambient (400 µatm) and elevated (850 µatm) pCO2 treatments held at a constant temperature. Net calcification rates of Porites cores significantly decreased in the elevated treatment. Presence of L. laevigata decreased net calcification rates of Porites regardless of CO2 treatment. I also compared the bioerosion rate of L.
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