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UNIVERSITY of CALIFORNIA, SAN DIEGO Consumers UNIVERSITY OF CALIFORNIA, SAN DIEGO Consumers on coral reefs: variation in the structure and function of herbivorous echinoid communities and their consequent effects on benthic dynamics A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Oceanography by Levi Steele Lewis Committee in charge: Professor Jennifer Smith, Chair Professor Andreas Andersson Professor James Leichter Professor Lisa Levin Professor Stuart Sandin Professor Jonathan Shurin 2016 Copyright Levi Steele Lewis, 2016 All rights reserved. The Dissertation of Levi Steele Lewis is approved, and it is acceptable in quality and form for publication on microfilm and electronically: ______________________________________________________________________ ______________________________________________________________________ ______________________________________________________________________ ______________________________________________________________________ ______________________________________________________________________ ______________________________________________________________________ Chair University of California, San Diego 2016 iii DEDICATION To my ridiculous and wonderful parents & family, my personal Wonder Woman, the warm and conscientious people of Maui Makai Watch, the dedicated brilliant minds that must and will overcome, the socially aware and responsible, the sponge-crab lovers, yellow-cake makers, tank chuckers, the gracious tigermonkacuda, the inspiring viperfish, broad spectrum antibiotics, minivans, and the legion of volunteers who gift their time and talents to the understanding and preservation of nature. iv EPIGRAPH “The ideal scientist thinks like a poet and works like a bookkeeper,” - E.O. Wilson "Look at me now..." - a coral v TABLE OF CONTENTS DEDICATION .............................................................................................................. iv EPIGRAPH .....................................................................................................................v TABLE OF CONTENTS ............................................................................................... vi LIST OF FIGURES ...................................................................................................... vii LIST OF TABLES ..........................................................................................................x ACKNOWLEDGEMENTS .......................................................................................... xii VITA ............................................................................................................................xiv ABSTRACT OF THE DISSERTATION ....................................................................... xv CHAPTER 1 Introduction: Herbivory, functional diversity, and the structure and function of coral reef ecosystems .....................................................................................1 CHAPTER 2 Interactions between environmental characteristics, consumer effects, and the development of benthic communities on coral reefs: a comparative experimental approach ........................................................................................................................ 24 CHAPTER 3 Metabolic ecology of tropical herbivorous echinoids: allometric scaling of mass and metabolism in relation to the Metabolic Theory of Ecology ............................ 97 CHAPTER 4 Diversity in the ecological functions of sympatric herbivorous echinoids: metabolism, consumption rate, and dietary preference ................................................. 127 CHAPTER 5 Identity, diversity, and the ecological effects of herbivorous echinoid communities on a coral reef ......................................................................................... 168 CHAPTER 6 Conclusion: Advancing our understanding of spatial variation in consumer effects on benthic succession, interactions with the environmental setting, and the functional diversity and ecology of herbivorous echinoids on Hawaiian coral reefs ..... 208 vi LIST OF FIGURES Figure 1.1. Prominent ecological dynamics of coral reef systems examined in the dissertation ......................................................................................................................7 Figure 2.1. Study sites and design of settlement tiles and cages ..................................... 84 Figure 2.2. Environmental characteristics measured at each of the 8 coral reef study sites ...................................................................................................................................... 85 Figure 2.3. Biomass of (a) macroalgae, (b) total non-calcified and (c) calcified material on CAU tiles .................................................................................................................. 86 Figure 2.4. Scatterplot matrix of relationships between environmental characteristics (herbivore biomass: HRB, dissolved inorganic nutrients: DIN, and sediment accumulation rate: SAR) and dry mass of macroalgae, non-calcified and calcified material on CAU tiles. ................................................................................................... 87 Figure 2.5. Relative magnitude of consumer effects (log response ratio of open:caged tiles) on total biomass of (a) macroalgae, (b) total non-calcified, and (c) calcified biomass on CAU tiles .................................................................................................................. 88 Figure 2.6. Scatterplot matrix of relationships between environmental characteristics (herbivore biomass: HRB, dissolved inorganic nutrients: DIN, and sediment accumulation rate: SAR) and the magnitude of the effects of consumers on macroalgae, non-calcified, and calcified biomass on CAU tiles. ........................................................ 89 Figure 2.7. Community structure of sessile organisms and substrates on exposed surfaces of CAU tiles. ................................................................................................................. 90 Figure 2.8. Effect size (open-cage, raw %) of consumers on the abundance (% cover) of 3 dominant benthic functional groups on CAU tiles ....................................................... 91 Figure 2.9. Community structure of sessile organisms on CAU surfaces ....................... 92 Figure 2.10. Coral recruitment among sites and in relation to sediment exposure .......... 93 Figure 2.11. Composition of benthic communities at study sites along leeward Maui. ... 94 Figure 2.12. Examination of CAU tile cage artifacts. ..................................................... 95 Figure 2.13. Relationships between non-calcified and calcified biomass among sites. ... 96 vii Figure 3.1. Taxonomy and morphology of 5 tropical echinoids examined in the this study .................................................................................................................................... 124 Figure 3.2. Allometric relationships of echinoid mass and metabolism for 5 tropical herbivorous echinoids .................................................................................................. 125 Figure 3.3. Allometric scaling coefficients for mass and metabolism of 5 tropical echinoids ..................................................................................................................... 126 Figure 4.1. Location of echinoid collection and grazing assay site on Maui, Hawaii; and the Maui Ocean Center where metabolic analyses were conducted. ............................. 160 Figure 4.2. Taxonomy and mophology of echinoids (top) and algae (bottom) used in algal grazing assays. ............................................................................................................. 161 Figure 4.3. Design of field-based grazing assays (A-C) and metabolic assays (D-F). ... 162 Figure 4.4. Control-corrected total and mass-specific grazing (A-B) and metabolic (C-D) rates, and relationships between total (E) and mass-specific (F) grazing vs. metabolic rates. ............................................................................................................................ 163 Figure 4.5. Corrected percent consumption (A) and preference (B) derived from results of field-based grazing assays............................................................................................ 164 Figure 4.6. Dissimilarity in algal preference profiles and mass-specific metabolic rates .................................................................................................................................... 165 Figure 4.7. Method for estimating consumption of turf and CCA ................................. 166 Figure 4.8. Echinoid sizes and control values for grazing and metabolic assays .......... 167 Figure 5.1. Location of study site on Maui, Hawaii (a), and examples of the four general diversity treatments: open (b), full exclosure (c), single -sp. (d), multi-sp. (e) .............. 199 Figure 5.2. Schematic of the 8 experimental treatments established within each of the 5 spatial blocks ............................................................................................................... 200 Figure 5.3. Mean (± SE) (a) densities of algal thalli, (b) turf heights, and (c) percent community composition (by density) of erect macroalgal functional groups and juvenile P. meandrina corals; each at initial, 1-yr and 2-yr time points (year and month provided)
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