The Biology of Coral Reefs (Biology of Habitats Series)

## Abstract Coral reefs represent the most spectacular and diverse marine ecosystem on the planet as well as a critical source of protein and income for many millions of people. Ecologically they are as complex as they are diverse and colourful. However, the combined effects of human activities have led to a rapid decline worldwide in the health of reefs. This timely book provides an integrated overview of the function, physiology, ecology and behaviour of coral reef organisms. Each chapter is enriched with a selection of ‘boxes’ on specific aspects written by internationally recognized experts. As with other books in the Biology of Habitats Series, the emphasis in this book is on the organisms that dominate this marine environment, although pollution, conservation, climate change and experimental aspects are also included. Indeed, particular emphasis is placed on conservation and management due to the habitat’s critically endangered status. A global range of examples is employed which gives the book international relevance. This accessible text is intended for students, naturalists and professionals and assumes no previous knowledge of coral reef biology. It is particularly suitable for both senior undergraduate and graduate students (in departments of biology, geography, and environmental science) taking courses in coral reef ecology, marine biology, oceanography and conservation biology, as well as the many professional ecologists and conservation biologists requiring a concise overview of the topic. It will also be of relevance and use to reef managers, recreational divers and amateur naturalists. Cover 1 Contents 8 Abbreviations 12 List of boxes 14 Plates 16 1 Coral reefs: Biodiverse and productive tropical ecosystems 24 1.1 Introduction 24 1.2 Areas and distributions of reefs 33 1.3 Biodiversity on coral reefs 36 1.4 Coral communities and reef growth 40 1.4.1 The fate of coral limestone 42 1.5 Types of reefs 43 1.6 Profiles and zones of reefs 47 1.6.1 The reef flat 47 1.6.2 The reef crest 50 1.6.3 Reef slopes 52 1.7 Biodiversity on reef slopes 55 1.8 Values of reefs to people 57 2 The main reef builders and space occupiers 58 2.1 Corals 62 2.2 Soft corals and sea fans 71 2.3 Sponges 75 2.4 Other animal species 79 2.4.1 Molluscs, echinoderms, crustaceans, polychaetes 79 2.4.2 Cryptic and bioeroding species 80 2.5 Macroalgae 81 2.6 Seagrasses and mangroves 84 2.7 Rates of coral growth, rates of reef growth 87 2.8 Soft substrates 89 3 The abiotic environment 91 3.1 Controls on coral distribution 91 3.2 Salinity 91 3.3 Temperature 96 3.4 Light 101 3.5 Nutrients 104 3.6 Exposure and other hydrodynamic factors 109 3.7 Sediment 112 3.8 Seawater carbonate chemistry 116 4 Symbiotic interactions 123 4.1 What is symbiosis? 123 4.2 Algal–invertebrate symbioses in corals and soft corals 123 4.3 Diversity of zooxanthellae 126 4.4 Physiology of coral calcification 130 4.5 Photosynthesis and carbon fluxes 132 4.6 Nitrogen acquisition and fluxes 139 4.7 Phosphorus 142 4.8 Symbiosis establishment and stability 143 4.9 Coral–microbial associations 148 4.10 Sponge symbioses with non-photosynthetic bacteria, cyanobacteria and algae 152 4.11 ‘Macro’ or iconic symbioses (e.g. fish and sea anemones, shrimps and fish) 154 5 Microbial, microalgal and planktonic reef life 157 5.1 Microbial reef life 157 5.1.1 Bacteria and archaea 157 5.1.2 Viruses 165 5.1.3 Fungi 167 5.1.4 Protozoa 167 5.1.5 Microalgae 169 5.1.6 Microbial productivity and turnover 171 5.2 Trophic links 173 5.2.1 The microbial loop 173 5.2.2 Consumption of microorganism 175 5.3 Zooplankton behaviour and ecology 182 5.3.1 Planktonic durations of larvae and planktonic dispersal 182 5.3.2 Diurnal cycles of demersal plankton 184 5.3.3 Reef connectivity 187 6 Reef fish: Evolution, diversity and function 190 6.1 Evolution and biogeography 190 6.2 Age and growth 192 6.3 Larval fish ecology 194 6.4 Reef fish diversity 195 6.5 Reef fish: Colourful for a reason 199 6.5.1 Poisonous and venomous fish 199 6.5.2 Camouflage 201 6.5.3 Mimicry 201 6.6 Abundance, biomass and trophic structure 201 6.7 Feeding and ecosystem function 203 6.7.1 Detritivores 203 6.7.2 Herbivores 205 6.7.3 Planktivores 207 6.7.4 Corallivores 208 6.7.5 Invertebrate feeders 209 6.7.6 Piscivores 209 6.7.7 Ecosystem function 210 6.8 Disturbances and coral reef fish 211 7 Reef fisheries and reef aquaculture 220 7.1 Fisheries resources on reefs 221 7.1.1 Vertebrate resources 221 7.1.2 Invertebrate resources 224 7.1.3 Practical issues with reef fisheries 232 7.1.4 Fishing with dynamite and poisons 233 7.2 Live reef fish trade 235 7.3 Aquaculture on reefs 240 7.4 Impacts of fishing 243 7.5 Approaches to managing coral reef fisheries 246 7.6 Controlling coral reef fisheries 250 7.7 Future for reef fisheries 252 8 Coral reefs in the modern world 255 8.1 Damaging impacts on reefs 256 8.1.1 Nutrient enrichment 256 8.1.2 Industrial and physical impacts: Landfill, dredging, sedimentation 257 8.1.3 Chemical and oil pollution 262 8.2 Other physical impacts 263 8.2.1 Structural stress from construction 263 8.2.2 Boat anchoring on reefs 264 8.2.3 Nuclear testing 266 8.3 Coral diseases 267 8.4 Diseases of other reef organisms 270 8.4.1 Red algae 270 8.4.2 Caribbean Diadema 270 8.4.3 Caribbean gorgonians 271 8.5 Climate change 272 8.5.1 Temperature rise 272 8.5.2 The progression of a bleaching event 276 8.5.3 Sea surface temperature curves and forecasts 277 8.6 Acidification 280 8.6.1 Slowing of reef calcification 280 8.7 Sea level rise 282 8.8 Cyclones, hurricanes, typhoons 283 8.8.1 Damage from storm energy 285 8.8.2 Tsunamis 286 8.9 Synergies, stasis and feedbacks 286 9 Consequences to reefs of changing environmental stress 289 9.1 Ecological consequences of environmental impacts 289 9.1.1 Alternative stable states, thresholds, phase shifts and hysteresis 289 9.2 Changes to the main architectural species 293 9.2.1 Acropora die-off in the Caribbean 293 9.2.2 Montastraea and Orbicella in the Caribbean 297 9.3 Changes on Indo-Pacific reefs 298 9.3.1 Indo-Pacific shallow assemblages 298 9.3.2 Changes to reef fish 301 9.3.3 Effects of bioeroding species 303 9.3.4 Role of reefs in wave energy reduction 304 9.4 Sizes of the coral ‘reservoir’ 308 9.5 Changing food chains and trophic balances 310 10 The future, human population and management 312 10.1 The timescale available 313 10.1.1 Possible adaptation to stressors 315 10.2 The elephant in the room 316 10.2.1 Shifting-baseline syndrome 318 10.3 Protected areas and coral reefs 319 10.3.1 Connectivity and selection of areas 321 10.4 Environmental assessments 323 10.4.1 Offset schemes 324 10.5 Costs, prices and values 325 10.5.1 Economic costs 326 10.6 The ‘slippery slope to slime’ 327 10.7 The future for reefs 328 Bibliography 338 Index 388 Coral reefs represent the most spectacular and diverse marine ecosystem on the planet as well as a critical source of income for millions of people. However, the combined effects of human activity have led to a rapid decline in the health of reefs worldwide, with many now facing complete destruction. Their world-wide deterioration and over-exploitation has continued and even accelerated in many areas since the publication of the first edition in 2009. At the same time, there has been a near doubling in the number of scientific papers that have been written in this short time about coral reef biology and the ability to acclimate to ocean warming and acidification. This new edition has been thoroughly revised and updated, incorporating the significant increase in knowledge gained over the last decade whilst retaining the book's focus as a concise and affordable overview of the field. The Biology of Coral Reefs provides an integrated overview of the function, physiology, ecology, and behaviour of coral reef organisms. Each chapter is enriched with a selection of 'boxes' on specific aspects written by internationally recognised experts. As with other books in the Biology of Habitats Series, the emphasis in this book is on the organisms that dominate this marine environment although pollution, conservation, climate change, and experimental aspects are also included. Indeed, particular emphasis is placed on conservation and management due to the habitat's critically endangered status. A global range of examples is employed which gives the book international relevance. "Coral reefs represent the most spectacular and diverse marine ecosystem on the planet as well a being a critical source of income for millions of people. However, the combined effects of human activity have led to a rapid decline in the health of the reefs worldwide, with a significant proportion having been completely destroyed. This process has continued and even accelerated in many areas since the publication of the first edition in 2009. At the same time, there has been a near doubling in the number of scientific papers that have been written in this short time about coral reef biology. This new edition has been thoroughly revised and updated, incorporating this significant increase in knowledge whilst retaining the book's focus as a concise and affordable overview of the field." "'The biology of coral reefs' provides an integrated overview of the function, physiology, ecology and behaviour of coral reef organisms. Each chapter is enriched with a selection of 'boxes' on specific aspects written by internationally recognised experts. As with other books in the Biology of Habitats series, the emphasis in this book is on the organisms that build, regulate and dominate this marine environment, and the factors that affect them including pollution, conservation, climate change and experimental aspects. Indeed, particular emphasis is placed on conservation and management due to the habitat's critically endangered status. A global range of examples is employed which gives the book international relevance." "This accessible text is intended for students, naturalists and professionals and assumes no previous knowledge of coral reef biology. It is particularly suitable for both senior undergraduate and graduate students (in departments of biology, geography and environmental sciences) taking courses in coral reef ecology, marine biology, oceanography and conservation biology, as well as the many professional ecologists and conservation biologists requiring a concise overview of the topic. It will also be of relevance and use to reef managers, recreational divers and amateur naturalists." -- rear cover A Concise But Comprehensive Introduction To The Biology Of Coral Reefs, Providing An Overview Of The Ecology Of Coral Reefs And Their Functioning, And The Biology Of Their Major Species Groups. The Responses To Modern Environmental Pressures, Climate Change, And Use Of Their Resources Is Also Described. Charles R.c. Sheppard, Simon K. Davy, Graham M. Pilling, Nicholas A.j. Graham. Includes Bibliographical References And Index.