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UNIVERSITY of CALIFORNIA SAN DIEGO the Trophic Ecology of Reef

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UNIVERSITY of CALIFORNIA SAN DIEGO the Trophic Ecology of Reef UNIVERSITY OF CALIFORNIA SAN DIEGO The trophic ecology of reef-building corals: the influence of resource availability on coral nutrition at multiple scales A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Marine Biology by Michael Douglas Fox Committee in charge: Professor Jennifer Smith, Chair Professor Andreas Andersson Professor Elsa Cleland Professor Stuart Sandin Professor Brice Semmens 2018 Copyright Michael Douglas Fox, 2018 All rights reserved. The Dissertation of Michael Douglas Fox is approved, and it is acceptable in quality and form for publication on microfilm and electronically: _____________________________________________________________________ _____________________________________________________________________ _____________________________________________________________________ _____________________________________________________________________ _____________________________________________________________________ Chair University of California San Diego 2018 iii DEDICATION To my parents, Mary Kay and Douglas Fox, who always encouraged me to follow my dreams and instilled in me a passion for natural history at an early age. Thank you for always believing in me and for your never-ending support. iv EPIGRAPH “From our present vantage point in time, when every square meter of earth has been surveyed by satellite and can be ‘goggled’ at will, our challenge is no longer to discover what is there, but to understand how the many complex parts of the Earth’s systems interact. The challenge is not to push outward but to connect together.” Charles F. Kennel, Director of Scripps Institution of Oceanography 1997-2006 From the Forward of the book, Ocean: Reflections on a Century of Exploration by Wolf. H Berger v TABLE OF CONTENTS Signature Page .............................................................................................................. iii Dedication ..................................................................................................................... iv Epigraph ......................................................................................................................... v Table of Contents .......................................................................................................... vi List of Tables ................................................................................................................ vii List of Figures ............................................................................................................... viii Acknowledgements ....................................................................................................... x Vita ................................................................................................................................. xiii Abstract of the Dissertation .......................................................................................... xv Introduction ................................................................................................................... 1 CHAPTER 1: Gradients in nearshore primary production predict the trophic strategies of mixotrophic corals at multiple spatial scales ......................................... 7 CHAPTER 2: Amino acid δ13C analysis reveals trophic plasticity in a common reef-building coral .................................................................................. 49 CHAPTER 3: Nutrient enrichment differentially affects calcification in two Hawaiian coral species .......................................................................................... 82 Conclusion .................................................................................................................... 111 vi LIST OF TABLES Chapter 1 Table 1.S1: Analysis of Covariance (ANCOVA) results .............................................. 45 Table 1.S2: Global data set of coral δ13C and δ15N and nearshore chl-a ....................... 46 Table 1.S3: Results of linear regression analyses designed to examine the influence of taxonomic resolution and location on the global relationship between Δ13C and chl-a ............................................................................................ 47 Table 1.S4: Results of linear regression analysis examining the relationship between coral δ13C and δ15N with absolute latitude, chl-a, and estimated thermocline depth ....................................................................................... 48 Chapter 2 Table 2.S1: All sites around Palmyra and the sample size for each isotopic metric analyzed ........................................................................................... 79 Table 1.S2: Analysis of Variance (ANOVA) results examining coral and endosymbiont stable isotope values as well as their relative difference (Δ) ...................... 80 13 Table 1.S3: Individual ANOVAs comparing differences in δ CAA across the three source groups .............................................................................................. 81 Chapter 3 Table 3.1: Statistical summary for the fixed effects of linear mixed effects models. .... 104 vii LIST OF FIGURES Chapter 1 Figure 1.1: Oceanographic climate of the Southern Line Islands .................................. 40 Figure 1.2: Pocillopora meandrina host and endosymbiont δ13C and ∆13C across depth in the SLI ............................................................................................ 41 Figure 1.3: Global patterns of coral and endosymbiont δ13C and ∆13C as a function of surfac chl-a .............................................................................................. 42 Figure 1.S1: Surface area normalized chl-a concentrations (µg chl-a cm-2) of Pocillopora meandrina across islands and depth ..................................... 43 Figure 1.S2: Linear relationships between mean coral and symbionts δ13C and ∆13C with latitude, mean chl-a, and estimated depth of the thermocline ........... 44 Chapter 2 13 Figure 2.1: A comparison between δ C analysis of six essential amino acids (AAESS) and bulk tissue δ13C and δ15N analysis of a mixotrophic coral ................... 72 Figure 2.2: Individual variation in coral nutrition across 19 colonies from four Locations ..................................................................................................... 73 Figure 2.S1: Map of Palmyra Atoll National Wildlife Refuge ...................................... 74 Figure 2.S2: Boxplot of δ13C values of seven non-essential and six essential amino acids ............................................................................................... 75 Figure 2.S3: A comparison of the modeled percent contribution of heterotrophy to P. meandrina diets using MixSIAR and the bootstrapped LDA approach .................................................................................................... 76 Figure 2.S4: Linear discriminant analysis of the seven analyzed non-essential amino acids ............................................................................................... 77 Figure 2.S5: Linear discriminant analysis of six essential amino acids ........................ 78 Chapter 3 Figure 3.1: Mean inorganic nutrient concentrations for each treatment level ............... 105 Figure 3.2: Growth and total protein content in Porites compressa and Pocillopora acuta ........................................................................................ 106 viii Figure 3.3: Photophysiological response of Porites compressa to nutrient enrichment ................................................................................................... 107 Figure 3.4: Photophysiological response of Pocillopora acuta to nutrient enrichment ................................................................................................... 108 Figure 3.5: Maximum quantum yield in both coral species over the course of the experiment ........................................................................................ 109 Figure 3.S1: Environmental conditions throughout the experiment .............................. 110 ix ACKNOWLEDGEMENTS Thank you to my advisor, Jennifer Smith, for her guidance and support over the past five years. I am incredibly grateful for the opportunities you have provided me to study and learn from some of the most remote and spectacular reefs on the planet. Jen’s love of natural history and passion for benthic ecology is contagious and has helped me remember to step back and appreciate the little things even while we’re working like crazy in the field. Thank you for the countless opportunities and for introducing me to an exceptional group of colleagues. I will never forget the experiences I have had and the lessons I’ve learned during the numerous expeditions that I have been a part of while in your lab. I would also like to thank my committee, who has provided insightful feedback and constant encouragement over the last few years. Stuart, thank you for always being willing to chat with me about ways to analyze and explore data, those conversations have been invaluable to my development as an ecologist. Andreas, thank you for your mentorship and support over the 18 months it took to get some positive reviews for my first manuscript. The scientific process can be daunting at times but your positive attitude and thoughtful words of encouragement really helped me keep my eye on the prize. Elsa, thank you for the opportunity to take my
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