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CALIFORNIA STATE UNIVERSITY, NORTHRIDGE Foundation Species CALIFORNIA STATE UNIVERSITY, NORTHRIDGE Foundation species loss alters ecosystem functioning within temperate tidepool communities A thesis submitted in partial fulfillment of the requirements For the degree of Master of Science in Biology By Jennifer Fields December 2020 The thesis of Jennifer B. Fields is approved: _______________________________________ _______________ Dr. Kerry J. Nickols, Ph.D. Date ______________________________________ _______________ Dr. Steve R. Dudgeon, Ph.D. Date _______________________________________ _______________ Dr. Nyssa J. Silbiger, Ph.D. (Chair) Date California State University, Northridge ii ACKNOWLEDGEMENTS Science never happens in isolation. This Master’s is the product of endless hard work from a team of amazing and passionate people. My thesis advisor, Dr. Nyssa Silbiger, has been the mentor I could only dream of being myself one day. Without your connections with Oregon Department of Fish and Wildlife (ODFW) Marine Reserve team, I would not be able to do the master’s project that I have been dreaming about since studying barnacles in undergrad. Thank you for your endless edits, letters of recommendation, and support as I pushed my computational abilities and gained confidence in R stats. You have encouraged all of us to be not only better scientists and science communicators, but also better humans that advocate to make this world a more inclusive and equitable space. The theme of #everythingisconnected and memories from the Silbiger Lab (e.g. impromptu dance parties) are something I will carry fondly forever. Thank you to my thesis committee members, Dr. Kerry Nickols and Dr. Steve Dudgeon for their expertise and feedback on study design and statistical analyses. I have learned so much from you both and I always felt supported by you with my project and my scientific abilities. Dr. Dudgeon, thank you for your insight into grazing experiments and for helping with the omnivore’s dilemma. Your knowledge of intertidal intricacies is always incredibly appreciated. Dr. Nickols, thank you for always bringing me back to the bigger oceanographic and physical picture. You have challenged me to dive deeper into my data to understand more of “how”, and “why” questions within tide pool systems. Thank you for being my inspiration to pursue coastal policy as my future career. I literally would not be a future CA Sea Grant State Fellow without your support. I am extremely grateful for my labmates, Amanda Chiachi, Danielle Barnas, Danielle Becker, Katie Hannibal, and Jamie Kerlin, for their advice, support, and, in non-COVID times, iii hugs. I am grateful for my shared (Silbiger/Nickols) labmates, Deme Panos, Brian Cohn, Taylorann Smith, and Emily Wilson, for always inspiring me, providing tea, and endless laughter. For all the undergraduate students in our lab present and past, Maria Martone, Emily Rukstales, Natalie Dahan, Miranda Gilhuys, and Julio Rosales, it has been amazing to see you grow as rockstar scientists and thank for inspiring me with your passion. Thank you to my broader California State University, Northridge (CSUN) cohort, especially the CSUN trail run club (Sarah Wenner, Robert Hogg, Kiran Reed, Jack Girard, and Chelsea Brisson) and my housemates Jennica Moffat (feat. Hoodini), Chris Polinski, and Danielle Becker, whom I would not have gotten through the last 2.5 years without. Thank you to the Biology Department and stock room staff for always providing generous support and assistance to my laundry list of tasks. Thanks to my incredible team of intertidal rockstar volunteers: A. Chiachi, M. Barrios, C. Polinski, J. Moffat, S. McClarence, C. Garvey, S. Vogel, D. Zuk, A. Marino, O. Martin, M. Hauer-Jensen, K. Bauer, C. Fields, D. Becker, M. Nguyen, A. Chin, M. Yamane, T. Miles, C. Weiler, and H. Fulton-Bennett, especially during night water sampling. Special thank you to my core assistants M. Gilhuys, J. Rosales, and D. Barnas, whom learned along with me how to be flexible and manage a multifaceted field experiment. I could have never accomplished this without your enthusiasm and hard work. This project would not have come to life without the research, lab, and housing support from ODFW Marine Reserve staff, Oregon State Parks, and Oregon State University’s Hatfield Marine Science Center. Funding for this project was provided by CSUN Research and Graduate Studies, CSUN Department of Biology, CSU Council on Ocean Affairs, Science & Technology Graduate Research Grant, Association of Retired Faculty, Sigma Xi Grants-in-Aid of Research, and Dr. iv Julie Gorchynski research grant. This project was conducted under ODFW Scientific Taking Permit #22930 and Oregon Parks and Recreation Department Scientific Take Permit #017-19 as well as California DFW permit #13047 and CA State Parks permit #18- 820-50 for preliminary testing. Lastly, to my family and friends that responsible for the person I am today, thank you for your endless support of my dreams and for always inspiring me to pursue my passions. v DEDICATION I dedicate this thesis to literal metric ton of mussels I removed from the intertidal. I thank you for the endless wisdom I have gained from destroying thousands of critters’ home. I hope you made some hungry Pisaster very happy. vi Table of Contents Signature Page ii Acknowledgements iii Dedication vi List of Tables viii List of Figures ix Abstract x Introduction 1 Methods 5 Results 17 In-text tables and figures 19 Discussion 30 References 36 Appendix A: Tables 43 Appendix B: Figures 55 vii LIST OF TABLES TABLE 1 43 TABLE 2 45 TABLE 3 46 TABLE 4 47 TABLE 5 48 TABLE 6 49 TABLE 7 50 TABLE 8 51 TABLE 9 52 TABLE 10 53 viii LIST OF FIGURES FIGURE 1 19 FIGURE 2 20 FIGURE 3 21 FIGURE 4 23 FIGURE 5 25 FIGURE 6 26 FIGURE 7 29 Appendix B FIGURE 1 55 Appendix B FIGURE 2 56 Appendix B FIGURE 3 57 Appendix B FIGURE 4 58 ix ABSTRACT Foundation species loss alters ecosystem functioning within temperate tidepool communities By Jennifer Fields Master of Science in Biology Foundation species are organisms that create shelter, enhance biodiversity, and maintain ecosystem functioning within their environment. Within the rocky intertidal ecosystem, a coastal ecosystem that can be dominated by mussels and surfgrass, foundation species are expected to decline with climate change, extreme climatic events, and increased human impact. Although studies have investigated the impact of intertidal foundation species on community structure and some resource fluxes, there is a gap in knowledge to understand the magnitude of effects that intertidal foundation species loss will have on multiple facets of ecosystem functioning in concert using a mechanistic approach. I studied the direct and indirect effects of foundation species loss of mussels (Mytilus californianus) and surfgrass (Phyllospadix spp.) on community structure and resource fluxes via in situ tide pool manipulations. I measured multiple facets of ecosystem function including stocks (community structure), fluxes (light, temperature, dissolved oxygen, dissolved inorganic nutrients, pH), and ecosystem metabolism (net ecosystem calcification [NEC] and net ecosystem production [NEP]) in mussel- and surfgrass-dominated tide pools in central Oregon before and after removal of foundation species. I quantified the interdependency between changes in stocks, fluxes, and ecosystem metabolism along a gradient of foundation species loss using a structural equation model. Surfgrass and mussel loss caused significant changes in the sessile communities, increased the light and temperature environment, shifted biogeochemistry, and directly and indirectly increased pH and ecosystem metabolism. Shifts in both surfgrass and mussel sessile communities with foundation species loss were driven mostly by increasing diatom cover. Surfgrass loss increased daily maximum temperature by 7.7°C and percent light by 30%, whereas mussel loss increased temperature by 3.8°C, but not the light environment over the month-long study. Tide pools with foundation species present shifted in accordance with changing ocean biogeochemistry, while pools with foundation species loss had higher in dissolved oxygen, pH, and temperature variance values. Surfgrass loss had strong positive indirect effects of NEP, pH, and NEC mediated by the direct increase in light environment with percent surfgrass loss. Mussel loss indirectly increased NEP mediated by direct increases in micro/macroalgal cover with mussel loss and directly increased pH through the removal of the dominant consumer within the system. This study highlights the importance of measuring the ecosystem as a whole with a mechanistic approach to understand the magnitude of changes that will occur with intertidal foundation species loss with increasing anthropogenic impacts. x Introduction Foundation species are a major focus of ecological studies due to their importance to ecosystem processes and conservation management (Byers et al. 2006, Halpern et al. 2007, Angelini et al. 2011, Ellison 2019). As the dominant, habitat-building organism within biological communities, foundation species facilitate interactions that promote biodiversity, stabilize community structure, and maintain fluxes (e.g. nutrient cycling, carbon cycling, or production rates) of ecosystem functioning (Bruno and Bertness 2001, Kinzig et al. 2001, Angelini et al. 2011, Ellison 2019). Recent studies have emphasized the connection between foundation species and ecological services, such as shoreline protection, nutrient cycling, and food security, as the direct consequences
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