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UNIVERSITY of CALIFORNIA, SAN DIEGO Historical Change in Coral UNIVERSITY OF CALIFORNIA, SAN DIEGO Historical change in coral reef communities in Caribbean Panama A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Oceanography by Katie Lynn Cramer Committee in charge: Professor Jeremy Jackson, Chair Professor Christopher Charles Professor James Leichter Professor Richard Norris Professor Naomi Oreskes 2011 Copyright Katie Lynn Cramer, 2011 All rights reserved. The Dissertation of Katie Lynn Cramer is approved, and it is acceptable in quality and form for publication on microfilm and electronically: _____________________________________________ _____________________________________________ _____________________________________________ _____________________________________________ _____________________________________________ Chair University of California, San Diego 2011 iii DEDICATION For my parents, Harry and Connie Cramer iv TABLE OF CONTENTS Signature page……………………………………………………………………………iii Dedication………………………………………………………………………………...iv Table of contents…………………………………………………………………………..v List of Figures…………………………………………………………………………….ix List of Tables………………………………………………………………………….....xv Acknowledgements……………………………………………………………………...xvi Vita……………………………………………………………………………………….xx Abstract of the Dissertation……………………………………………………..............xxi CHAPTER 1: Geography, human occupation, and environmental change in western and central Caribbean Panama…………………………………………………………1 ABSTRACT…………………………………………………………………………...1 INTRODUCTION…………………………………………………………………….2 GEOLOGICAL SETTING……………………………………………………………4 ENVIRONMENTAL SETTING……………………………………………………...6 Panama…………………………………………………………………………….6 Bocas del Toro…………………………………………………………………….9 Costa Arriba……………………………………………………………………...10 HUMAN HISTORY…………………………………………………………………12 Pre-contact period………………………………………………………………..12 Bocas del Toro…………………………………………………...………15 Costa Arriba……………………………………………………………...19 Colonial period…………………………………………………………………...22 v Bocas del Toro……………………………………………………….…..24 Costa Arriba……………………………………………………………...26 Post-colonial period……………………………………………………………...28 Bocas del Toro…………………………………………………….……..30 Costa Arriba……………………………………………………...………35 CONCLUSIONS…………………………………………………………………….39 REFERENCES……………………………………………………………...……….41 CHAPTER 2: Historical change in coral communities in Caribbean Panama…….57 ABSTRACT………………………………………………………………………….57 INTRODUCTION…………………………………………………………………...58 METHODS…………………………………………………………………………..61 Study sites………………………………………………………………………..61 Sampling…………………………………………………………………………62 Taxa………………………………………………………………………………65 Statistical analysis………………………………………………………………..66 Radiocarbon dating………………………………………………………………68 Potential taphonomic biases……………………………………………………...69 RESULTS……………….…………………………………………………………...71 Coral data………………………………………………………………………...71 Bocas del Toro…………………………………………………………...72 Radiocarbon dates………………………………………………..72 Coral community composition…………………………………...74 Costa Arriba……………………………………………………………...78 vi Radiocarbon dates………………………………………………..78 Coral community composition…………………………………...79 Comparison of regions…………………………………………...81 DISCUSSION………………………………………………………………………..82 Community change………………………………………………………………82 Community change and human disturbance……………………………………..85 Comparison with previous studies……………………………………………….87 Potential taphonomic biases……………………………………………………...88 CONCLUSIONS…………………………………………………………………….88 REFERENCES………………………………………………………………………90 CHAPTER 3: Historical change in coral reef bivalve and gastropod communities in Caribbean Panama…………………………………………………………………118 ABSTRACT………………………………………………………………………...118 INTRODUCTION………………………………………………………………….119 METHODS…………………………………………………………………………123 Data collection………………………………………………………………….123 Mollusk data…………….………………………………………………………125 Statistical analyses……………………………………………………………...126 Radiocarbon dating……………………………………………………………..127 Potential taphonomic biases…………………………………………………….128 RESULTS…………………………………………………………………………..129 Radiocarbon dates………………………………………………………………129 Gastropods……………………………………………………………………...129 vii Gastropod data………………………………………………………….129 Bocas del Toro………………………………………………………….130 Costa Arriba…………………………………………………………….134 Comparison of regions………………………………………………….136 Bivalves…………………………………………………………………………137 Bivalve data………………………………………………………...…..137 Bocas del Toro………………………………………………………….138 Costa Arriba…………………………………………………………….141 Comparison of regions………………………………………………….144 DISCUSSION………………………………………………………………………145 CONCLUSIONS…………………………………………………………………...149 ACKNOWLEDGEMENTS………………………………………………………...150 REFERENCES……………………………………………………………………..151 CHAPTER 4: Ecological change on Caribbean coral reefs before coral disease and bleaching……………………………………………………………………………….196 ABSTRACT………………………………………………………………………...196 TEXT……………………………………………………………………………….196 REFERENCES……………………………………………………………………..204 viii LIST OF FIGURES Figure 1.1. Study regions and the two prehistoric human settlements discussed in text outside of study regions……………………………………………………...51 Figure 1.2. Map of Bocas del Toro with triangles indicating archaeological sites………52 Figure 1.3. Map of Costa Arriba with triangles indicating archaeological sites………...53 Figure 1.4. Population of Bocas del Toro and Costa Arriba, pre-contact period to present………………………………………………………………………..54 Figure 1.5. Human population of Bocas del Toro since 1832 and major events affecting the nearshore environment…………………………………………………...55 Figure 1.6. Human population of Costa Arriba since ~1550 and major events affecting the nearshore environment………………………………………………...…56 Figure 2.1. Sampling design. (A) regions and reef sites, (B) pits and transects………..103 Figure 2.2. Total mass of coral rubble analyzed for offshore and lagoonal sites.……...104 Figure 2.3. Bocas del Toro. Principal Components Analysis of species proportions, offshore and lagoonal sites.…………………………………………………105 Figure 2.4. Bocas del Toro. Species richness by time horizon ………………………...106 Figure 2.5. Offshore sites, Bocas del Toro. Mean relative abundance (proportion total weight) of species comprising at least 1% of community at any one time horizon ……………………………………………………………………..107 Figure 2.6. Lagoonal sites, Bocas del Toro. Mean relative abundance (proportion total weight) of species comprising at least 1% of community at any one time horizon.……………………………………………………………………..108 ix Figure 2.7. Offshore sites, Bocas del Toro. Relative abundance of dominant species over time.………………………………………………………………………...109 Figure 2.8. Lagoonal sites, Bocas del Toro. Relative abundance of dominant species over time ………………………………………………………………………...110 Figure 2.9. Costa Arriba. Principal Components Analysis of species proportions, offshore sites and lagoonal sites……………………………………………………...111 Figure 2.10. Costa Arriba. Species richness by time horizon ………………………….112 Figure 2.11. Offshore sites, Costa Arriba. Mean relative abundance (proportion total weight) of species comprising at least 1% of community at any one time horizon ……………………………………………………………………..113 Figure 2.12. Lagoonal sites, Costa Arriba. Mean relative abundance (proportion total weight) of species comprising at least 1% of community at any one time horizon…………………………………………...…………………………114 Figure 2.13. Offshore sites, Costa Arriba. Relative abundance of dominant species over time…………………………………………………………………………115 Figure 2.14. Lagoonal sites, Costa Arriba. Relative abundance of dominant species over time…………………………………...…………………………………….116 Figure 2.15. Bocas del Toro and Costa Arriba. A) Principal Components Analysis of species proportions, B) Scores for first principal component by region and environment……………………………………………………………...…117 Figure 3.1. Gastropods, Bocas del Toro. Richness, total weight, total abundance, and average individual weight of gastropod shells……………………………...160 x Figure 3.2. Gastropods, Bocas del Toro. Richness, total weight, number of individuals, and individual shell weight by time horizon ……………………………….161 Figure 3.3. Gastropods, Bocas del Toro. Principal Components Analysis of species proportion by abundance (number of individuals)………………………….162 Figure 3.4. Gastropods, offshore sites, Bocas del Toro. Mean relative abundance (proportion individuals) of species comprising ≥1% of community in at least one time horizon …………………………………………………………...163 Figure 3.5. Gastropods, lagoonal sites, Bocas del Toro. Mean relative abundance (proportion individuals) of species comprising ≥1% of community in at least one time horizon …………………………………………………………...164 Figure 3.6. Gastropods, offshore sites, Bocas del Toro. Proportion over time of common gastropod species (>5% of community on average)………………………..165 Figure 3.7. Gastropods, lagoonal sites, Bocas del Toro. Proportion over time of common gastropod species (>5% of community on average)…………......................166 Figure 3.8. Gastropods, Bocas del Toro. Proportion carnivorous individuals………….167 Figure 3.9. Gastropods, Costa Arriba. Total weight, total abundance, and average individual weight of shells …………………………………………………168 Figure 3.10. Gastropods, Costa Arriba. Richness, total weight, number of individuals, and individual shell weight by time horizon ……………………………………169 Figure 3.11. Gastropods, Costa Arriba. Principal Components Analysis of species proportion by abundance (number of individuals). ………………………...170 xi Figure 3.12. Gastropods, offshore sites, Costa Arriba. Mean relative abundance (proportion individuals) of species comprising ≥ 1% of gastropod community in one or more time horizons.………………………………………….…...171 Figure 3.13. Gastropods, lagoonal
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