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California State University, Northridge an Ecological CALIFORNIA STATE UNIVERSITY, NORTHRIDGE AN ECOLOGICAL AND PHYSIOLOGICAL ASSESSMENT OF TROPICAL CORAL REEF RESPONSES TO PAST AND PROJECTED DISTURBANCES A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biology By Elizabeth Ann Lenz May 2014 The thesis of Elizabeth A. Lenz is approved by: Robert C. Carpenter, Ph.D. Date: Eric D. Sanford, Ph.D. Date: Mark A. Steele, Ph.D. Date: Peter J. Edmunds, Ph.D., Chair Date: California State University, Northridge ii ACKNOWLEDGEMENTS I would like to thank Dr. Peter J. Edmunds first and foremost for being my fearless leader and advisor - for the incredible opportunities and invaluable mentorship he has provided to me as a graduate student in the Polyp Lab. I am ever so grateful for his guidance, endless caffeinated energy, constructive critiques, and dry British humor. I would also like to thank my loyal committee members Drs. Robert Carpenter and Mark Steele at CSUN for their availability and expert advise during this process. Their suggestions have greatly contributed to my thesis. I would not only like to acknowledge Dr. Eric Sanford from UC Davis for serving on my committee, but thank him for his incessant support throughout my career over the last 7 years. I will always admire his contagious enthusiasm for invertebrates, passion for scientific research, and unlimited knowledge about marine ecology. My research would not have been possible without the technical support and assistance from my colleagues in Moorea, French Polynesia and St. John, USVI. I am grateful to Dr. Lorenzo Bramanti, Dr. Steeve Comeau, Vince Moriarty, Nate Spindel, Emily Rivest, Christopher Wall, Darren Brown, Alexandre Yarid, Nicolas Evensen, Craig Didden, the VIERS staff, and undergraduate assistants: Kristin Privitera-Johnson and Amanda Arnold. I would like to thank the former and current members of the CSUN Biology Department. Particularly, I am thankful for the MBGSA community, WiS group, Hollie Putnam, Anya Brown, Lianne Jacobson, Stella Swanson, Heather Hillard, Amy Briggs, Camdilla Wirth, Lauren Valentino, Mia Adreani, Justin Hackitt, and of course my fellow female scholars in the Polyp Lab: Sylvia Zamudio, Jennifer Smolenski, and Ananda Ellis. I would like to thank my constant sources of encouragement - Dr. Thomas Lenz and Debbra Lenz for their unconditional love and patience, my five siblings especially Mary Lenz for her generous time and reliability, and for my dear friends for enhancing my graduate experience: Sarah McVay, Emily Tung, Lauren Burgunder, Megan Boyd, Theresa Dineen, Malia Collins, Melanie Levy, Jackie Sones, DSO Jason Heron, Kristin Aquilino, Christopher Kwan, Bart Bagdasaryan, Casey Scott, Hamilton W. Fennie, Alexander G. Snyder, Kerry Nichols, Evelyne Kuo, Brian Lee, Kirk Sato, Tren Kauzer, Paul Logston, Maggie Sogin, Sean McCann, and Lauren Mickool. Finally, I am grateful to Drs. Annaliese Hettinger and John J. Stachowicz for recommending I pursue my master’s degree at CSUN in the Polyp Lab. My graduate thesis was made possibly by the financial support of CSUN’s College of Science and Mathematics Graduate Research Promise Fellowship, the National Science Foundation research support for Moorea Coral Reef Long-Term Ecological Research (OCE 10-26852), Division of Ocean Science (OCE 10-41270), St. John USVI Long- Term Research in Environmental Biology (DEB 03-43570, DEB 08-41441, and OCE 13- 32915), and the Gordon and Betty Moore Foundation. iii Signature Page ii Acknowledgements iii Abstract v Chapter 1 General Introduction 1 Chapter 2 Decadal-scale changes in the density of Caribbean gorgonians Introduction 14 Methods 17 Results 22 Discussion 27 Tables 36 Figures 48 Chapter 3 Contrasting morphologies of the Pacific coral Porites rus are not affected by ocean acidification Introduction 56 Methods 60 Results 70 Discussion 75 Tables 83 Figures 96 Chapter 4 Concluding remarks 105 Literature Cited 109 iv Abstract An Ecological and Physiological Assessment of Tropical Coral Reef Responses to Past and Projected Disturbances By Elizabeth A. Lenz Masters of Science in Biology Tropical reef structure and topographical complexity is a product of coral morphology, which greatly enhances biodiversity, ecological function, and ecosystem services. However, reef-building corals have declined in cover by 50-80% in response to the accumulated effects of natural and anthropogenic disturbances over the last 3 to 4 decades. This thesis is comprised of two different studies that address potential changes in shallow reef communities and benthic structure in response to natural and anthropogenic disturbances. The first objective was to examine changes in Caribbean benthic communities as scleractinians have declined and the second objective was to determine differential responses of contrasting morphotypes of corals to acidified conditions in Moorea, French Polynesia. In Chapter 2, I assessed the abundance of arborescent gorgonians (AGs) at local and regional scales to test the hypothesis that AGs have increased in abundance on Caribbean reefs over multiple decades. In St. John, mean abundance of AGs increased 42% from 1992-2012, with each of the dominant genera (Gorgonia, Antillogorgia, and pooled genera: Sea Rods) increasing 11-221% over the same period. Regionally, the compiled data show that AGs have increased in abundance, with mean densities rising from 7.0 to 15.1 colonies m-2 over the last 45 years. This study highlights the apparent success and importance of AGs on contemporary Caribbean v reefs, where scleractinian percent cover has remained at ~10%. In Chapter 3, I evaluated the calcification rates of branches and plates of Porites rus in response to ocean acidification (OA) (~800 µatm and 1000 µatm PCO2) with temperature, light and water flow. Skeletal morphology of reef-building hermatypic corals is influenced by abiotic factors, but calcification of coral skeleton is expected to decline as ocean acidification (OA) increases. Theory predicts that hermatypic corals with contrasting morphologies will differ in calcification rates due to differences in: 1) light absorption by Symbiodinium spp. for photosynthesis to enhance coral calcification, and 2) differential mass transfer characteristics (e.g., modulating the flux of dissolved inorganic carbon and hydrogen), which may mitigate the negative effects on calcification in high PCO2. In the second study, I determined that branches and plates were both tolerant of elevated temperature and PCO2. However, branches had higher variability in calcification rates than plates in response to light, and flow. Together these results of this research demonstrate two successful taxa (gorgonians and a weedy scleractinian) in tropical reef communities that have relatively high tolerances against natural and anthropogenic disturbances. Although, gorgonians and P. rus may be alternatives to formerly dominant, slow growing, and diverse communities of hermatypic corals, the functional roles and the ecosystem services they provide will likely differ. vi Chapter 1 General Introduction Disturbances on Tropical Coral Reefs Tropical coral reefs facilitate the highest biodiversity of any ecosystems in the world (Connell 1978) and provide economic value through coastal protection, fisheries, and ecotourism (Moberg and Folke 1999; Cesar et al. 2003). Hermatypic corals (P: Cnidaria, C: Anthozoa, SC: Hexacorallia, O: Scleractinia) are the ecosystem engineers responsible for the topographically complex framework that many organisms rely on for food and shelter (Idjadi and Edmunds 2006; Wild et al. 2011). The skeletal framework of scleractinians is created from the deposition of calcium carbonate (CaCO3) in the form of aragonite. However, natural and anthropogenic stressors are dramatically changing corals, from the molecular (e.g., gene regulation) to ecological scale (e.g., community structure) (Nyström et al. 2000; Bellwood et al. 2004; Hoegh-Guldberg et al. 2007). Tropical hermatypic coral communities are restricted geographically and constantly exposed to multiple stressors acting on reef systems. These stressors include both pulse (e.g., hurricanes, disease outbreaks, and thermal bleaching events) and press (e.g., ocean acidification and increased sea surface temperature [SST]) disturbances (Connell 1997). These assaults can have several grave consequences, with the effects on calcification and maintenance of reef complexity (Kleypas et al. 1999; Langdon and Atkinson 2005; Kleypas et al. 2001) at the forefront of concerns over the future of tropical coral reef communities (Knowlton 2001; Hough-Guldberg et al. 2007). As anthropogenic and natural disturbances continue, it is crucial to understand the physiological and ecological 1 responses of tropical corals to projected environmental conditions in order to determine reef resilience and shifts in coral reef communities. Changes in Coral Reef Communities Field surveys, photoquadrats, video recordings, and aerial photographs have been common tools used for long-term monitoring of scleractinian communities with percent cover the most widely used metric of coral abundance (Gardner et al. 2003; De’ath et al. 2013; Edmunds 2013; Ruzicka et al. 2013). In the last 3-5 decades, monitoring efforts have determined coral loss to be ~50% on the Great Barrier Reef from 1965-2012 (De’ath et al. 2013), ~80% throughout the Caribbean from 1977 to 2001 (Gardner et al. 2003), and ~22% in the Florida Keys from 1999-2009 (Ruzicka et al. 2013). In the Caribbean, the massive decline in scleractinian
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