The Biology and Ecology of Tintinnid Ciliates – Models for Marine Plankton

John Dolan has worked on plankton ecology for over 25 years in systems ranging from eutrophic estuaries to the oligotrophic South Pacific. His research subjects are plankton protists, especially ciliates and dinoflagellates and research topics are biodiversity, species packing, and the relationships of form and function. David Montagnes is an aquatic ecologist. He combines modelling, laboratory, and field studies investigations into the ecophysiology and behaviour of protists- aquatic taxa forming a rich assemblage and occupying environmentally important niches. His research themes include the flow of energy and biomass in ecosystems and how abiotic and biotic properties and anthropogenic perturbations alter individuals, populations, and ecosystems. Sabine Agatha is a ciliate taxonomist. She has spent over 20 years describing and classifying ciliates mostly from marine habitats by combining morphologic and genetic approaches. The biodiversity, evolution, and biogeography of ciliates are her main research topics. Wayne Coats is a recently retired protistologist. His research encompasses ecology, morphology, life history, and taxonomy of dinoflagellates and ciliates. Parasitism and mixotrophy are featured themes of his work. Diane Stoecker is a biological oceanographer. Her laboratory and field projects focus on microzooplankton and their grazing on phytoplankton and role as prey for copepods and other mesozooplankton. Plastid retention and mixotrophy in ciliates and dinoflagellates and their effects on population dynamics and trophic transfer are other favorite research topics.

CONTRIBUTORS vii CHAPTER 1 INTRODUCTION TO TINTINNIDS 1 1.1 Why a book on tintinnid ciliates? 2 1.2 What is a tintinnid ciliate? 3 1.3 The lorica as the defining characteristic of tintinnid ciliates 3 1.4 History of tintinnid studies 7 1.5 Tintinnids as model organisms for marine plankton 13 1.6 Key points 16 Acknowledgments 16 CHAPTER 2 THE TINTINNID LORICA 17 2.1 Introduction 18 2.2 Diversity formation and variability of loricae 19 2.3 Ultrastructure of loricae 32 2.4 Chemical composition of loricae 35 2.5 Lorica sedimentation 40 2.6 Key points 41 Acknowledgments 41 CHAPTER 3 SYSTEMATICS AND EVOLUTION OF TINTINNID CILIATES 42 3.1 Introduction 43 3.2 History of tintinnid taxonomy and systematics 44 3.3 Evolution of tintinnids 49 3.4 How to read the tintinnid cladogram 64 3.5 Molecular analysis and comparison with morphologic data 69 3.6 Systematics 75 3.7 Comparison with the evolution of related planktonic ciliates: the aloricate choreotrichids oligotrichids and halteriids 81 3.8 Key points 83 Acknowledgments 84 CHAPTER 4 ECOPHYSIOLOGY AND BEHAVIOR OF TINTINNIDS 85 4.1 Introduction 86 4.2 Feeding 86 4.3 Growth 99 4.4 Swimming behavior 106 4.5 Response to abiotic factors and interactions with biotic factors 111 4.6 Tintinnids as models and in models 114 4.7 Key points 121 Acknowledgments 121 CHAPTER 5 PREDATORS OF TINTINNIDS 122 5.1 Introduction 123 5.2 Predators of tintinnids 133 5.3 Anti-predator defenses of tintinnids 141 5.4 Top-down control of tintinnids 142 5.5 Importance of tintinnids as prey for the predator 143 5.6 Tintinnids as vectors for algal toxins 143 5.7 Key points 144 Acknowledgments 144 CHAPTER 6 PARASITES OF TINTINNIDS 145 6.1 Background 146 6.2 Dinoflagellate parasites: morphology and infection cycle 149 6.3 Molecular evolution of dinoflagellates parasitizing tintinnids 160 6.4 Ecology of tintinnid parasites 165 6.5 Summary and future directions 169 6.6 Key points 169 Acknowledgments 170 CHAPTER 7 COMPARATIVE BIOLOGY OF TINTINNID CYSTS 171 7.1 Introduction 172 7.2 Morphology 173 7.3 Encystment and excystment 176 7.4 Ecological function of cysts 182 7.5 Key points 184 Acknowledgments 185 CHAPTER 8 FOSSIL TINTINNIDS 186 8.1 Introduction: a paucity of data 187 8.2 Proterozoic tintinnids: no evidence of a beginning 187 8.3 Paleozoic tintinnid reports: too thin to know 191 8.4 Mesozoic tintinnids and calpionellids: same shape different composition 191 8.5 Cenozoic tintinnids: sparse 192 8.6 Organic and agglutinated phanerozoic fossils: the most likely candidates 193 8.7 Conclusions and perspectives 195 8.8 Key points 197 Acknowledgments 197 CHAPTER 9 TINTINNIDS IN MICROZOOPLANKTON COMMUNITIES 198 9.1 Ecology of microzooplankton 199 9.2 Quantitative importance of tintinnids in microzooplankton assemblages 205 9.3 Characteristics that set tintinnids apart from other microzooplankton 211 9.4 Key points 212 Acknowledgments 213 CHAPTER 10 DIVERSITY AND DISTRIBUTIONS OF TINTINNIDS 214 10.1 Introduction 215 10.2 Biogeography 215 10.3 Assemblages of coastal systems 233 10.4 Assemblages of open waters 239 10.5 Key points 241 Acknowledgments 243 REFERENCES 244 ORGANISM INDEX 288 SUBJECT INDEX 293