Paleobiology of the Polycystine Radiolaria

About the Editors David Lazarus has studied the paleobiology and earth science applications of Cenozoic radiolaria for more than 40 years, formerly holding research positions at Columbia University/Lamont Earth Observatory, the Woods Hole Oceanographic Institution, and the Eidgenössische Technische Hochschule Zürich. He is currently Curator for Micropaleontology at the Museum für Naturkunde in Berlin. Noritoshi Suzuki has studied the taxonomy and species diversity of radiolarians thoughout the Phanerozoic. He started his career in field geology, switched to Devonian radiolarians for his Masters degree, and received his PhD degree for a study of Cenozoic radiolarians from Tohoku University, Japan. He has co-published a monograph on the radiolarians of the Ehrenberg Collection (Berlin), and has published integrative studies of radiolarian morphology and phylogenetics. He is currently Associate Professor at Tohoku University. Yoshiyuki Ishitani is a paleobiologist, focusing on the evolution of radiolarians. He is currently a researcher at the University of Tsukuba, and was formerly at Japan Agency for Marine-Earth Science and Technology, Glasgow University, and the University of Tokyo. Kozo Takahashi has studied the distribution and ecology of radiolarians and other siliceous plankton collected from ocean waters for several decades. Following an early career of staff scientist positions at the Woods Hole and Scripps oceanographic institutions he held multiple professorships in Japan, including universities in Sapporo and Kyushu University in Fukuoka.

Preface xi

Acknowledgements xv

Chapter 1 History 1

Introduction 1

Scientific Context 4

Early Studies (First Half of the Nineteenth Century) 8

C.G. Ehrenberg and J. Müller 8

Second Half of the Nineteenth Century to ca. 1920 13

E. Haeckel and his Disciples 13

Legacy of Early Studies 16

Early Twentieth Century (ca. 1920–1940) 17

The Early New Period (ca. 1940–1970) 20

The Origins of Radiolarian Biostratigraphy: 1940s to 1950s 20

Deep-Sea Drilling 21

Taxonomy 25

Biology 27

Mid New Period (1970–2000) 28

Current Period (2000-Present) 37

Chapter 2 Biology 41

General Characteristics of Planktonic Protist Biology 41

Physical Characteristics of the Pelagic Ocean 42

Plankton Taxa 46

Ecologic and Behavioral Constraints due to Small Body Size 46

Basic Radiolarian Cellular Structure 48

Skeleton 53

Skeleton Formation and Growth 55

Size 59

Colonial Forms 59

Life Cycle 60

Longevity 62

Motility 63

Feeding 63

Predators 65

Abundance and Role in Carbon Cycle 66

Symbiosis 67

Bioluminescence 68

Summary 69

Chapter 3 Ecology 71

Introduction 71

Biogeography 75

Vertical Distribution 83

Tropical Submergence 86

Longitudinal Gradients and Upwelling Assemblages 89

Latitudinal Gradients 90

Coastal Gradients 90

Seasonal Variability 91

Interannual Variability 93

Chapter 4 Genetics 95

Introduction 95

Molecular Phylogenetic Position of “Radiolarians” within Eukaryotes 96

Molecular Studies of Radiolarian’s Position within Eukaryotes 97

Relationships of Radiolarian Clades 98

Origination Times of Radiolarian Clades 102

Family-Level Phylogeny 102

Spumellaria (Shell-Bearing Radiolarians) 105

Collodaria (Colonial or Naked Radiolarians) 105

Nassellaria 106

Acantharia 107

Microevolution of Radiolaria 107

Diversity of Pico-Radiolarian Material 111

Transcriptomics of Radiolaria 112

Methodology 113

DNA Extraction 114

Reproductive Cell Method 114

Dissecting Cell Method 114

PCR 114

Summary 114

Chapter 5 Taxonomy and Fossil Record 117

Introduction 117

PART 1 - Radiolarian Taxonomy 118

Principles of Species-Level Taxonomy 118

Rules for Describing and Naming Species 121

Current Status of Descriptive Radiolarian Taxonomy 124

Principles of Higher-Level Taxonomy 129

Haeckel and the Beginnings of Higher-Level Radiolarian Taxonomy 129

Biologic Systematics 132

Higher-Level Taxonomy in Radiolaria 134

The Observational Basis of Taxonomy: Structures of the Radiolarian Shell 136

Higher-Level Taxonomy in this Book 139

Formal Classification of Polycystina 143

Cenozoic Taxa 143

Order Spumellaria Ehrenberg 1876 143

Family Actinommidae Haeckel 1862 145

Family Heliodiscidae Haeckel 1881 149

Family Coccodiscidae Haeckel 1862, emend. Sanfilippo and Riedel 1980 151

Family Pyloniidae Haeckel 1881 153

Family Lithelidae Haeckel 1862 155

Family Tholonidae Haeckel 1887 156

Family Spongodiscidae Haeckel 1862 156

Order Nassellaria Ehrenberg 1876 160

Family Plagiacanthidae Hertwig 1879 162

Family Trissocyclidae (Haeckel) Goll 1968

[superfamily Acanthodesmiacea] 163

Family Theoperidae Haeckel 1881 163

Family Artostrobiidae Riedel 1967 167

Family Pterocoryithidae (Haeckel) Moore 1972 167

Family Carpocaniidae (Haeckel) Riedel, 1967 [Carpocaniinae] 171

Family Cannobotryidae Haeckel, 1881 173

Superfamily Collodaria 173

Family Collosphaeridae Müller, 1858 175

Family Sphaerozoidae Haeckel, 1862 175

Family Collophidiidae Biard and Suzuki, in Biard et al., 2015 177

Order Entactinaria 183

Family Orosphaeridae Haeckel, 1887 183

Family Saturnalidae Deflandre 1953 184

Mesozoic and Paleozoic Taxa 185

Species-Level Variation in Radiolaria 185

PART 2 - Summary of the Radiolarian Fossil Record 193

Cambrian and Ordovician 194

Silurian to the Lower Carboniferous 195

Late Paleozoic to Late Mesozoic Siliceous Sedimentation 196

Mass Extinctions at the End of the Paleozoic Era 197

Basal Mesozoic Scarcity of Radiolarian Fossils and Faunal Turnover (Early Triassic) 200

Triassic 201

Triassic–Jurassic Boundary Mass Extinction 204

Jurassic 205

Early and Middle Jurassic Radiolaria 205

Late Jurassic–Early Cretaceous 208

Cretaceous 208

The K/T Extinction Event and Early Paleocene 212

Cenozoic 214

Chapter 6 Preservation and Methods 217

Introduction 217

Preservation 218

Geographic Variation in Preservation 222

Diagenesis 222

Loss of Rock Record 224

Differences between Modern and Ancient Oceans 224

Quality of Radiolarian Fossil Record 225

Methods 227

Collecting Material from the Water Column 228

Collecting Sediments 231

Collecting Lithified Material from Sections on Land 236

Recovering Radiolarians from Samples 238

Extracting Radiolarians with Intact Protoplasm 238

Extracting Radiolarian Skeletons 238

Separation of Radiolarians from other Chemically Resistant Similar-Sized Components of Residue 242

Mounting Radiolarians 243

Live Preparations 245

Dissection and Serial Sectioning 246

Imaging Radiolarians 247

Visualization (enhanced imagery) 248

Morphometrics 249

Automatic Identification 249

Chapter 7 Paleoceanography 253

Introduction 253

Radiolarians as Tracers of Water Masses 259

Assemblage-Based Methods of Paleoceanographic Analysis 259

Non-temperature Uses of Assemblage Analyses 268

Radiolarians in Bulk: Summary Indices and Non-Taxonomic Uses of Radiolarians in Paleoceanography 273

Chapter 8 Radiolarian Biostratigraphy 281

Introduction 281

Biostratigraphy in Shallow Marine Rocks: General Aspects 283

Biostratigraphy in Deep-Sea Sediment Sections 285

Other Types of Geochronologic Information 287

Radiometric Dating and Absolute Age 287

Paleomagnetic Stratigraphy 288

Stable Isotope Stratigraphy 290

Cyclostratigraphy 291

Quantitative Biostratigraphy 292

Cenozoic Radiolarian Stratigraphy 295

History of Development 296

Tropical Cenozoic Radiolarian Stratigraphy 297

Subtropical North Atlantic to Arctic 299

North Pacific 302

Southern Ocean 305

History 305

Characteristics 307

Important Sections 307

Important Species 307

Mesozoic Radiolarian Stratigraphy 308

Cretaceous 308

Europe and Southwest North America 311

Low-Latitude Western part of Mesotethys 311

Mid-Ltitude Northern Part of Mesotethys 311

Russian Epicontinental Seas 312

East Margin of the Mid-Latitude Pacific 312

Northwest Pacific 312

Other Regions 313

The Jurassic–Cretaceous Boundary

(Tithonian–Berriasian Boundary) 313

Jurassic 314

Middle and Late Jurassic 314

Lower Jurassic 316

Triassic–Jurassic Boundary 316

Triassic 316

Latest Triassic (Rhaetian) 317

Carnian and Norian 318

Late Olenekian to Ladinian 318

Basal Triassic (Induan) and Permian–Triassic (P–T) boundary 318

Paleozoic Radiolarian Stratigraphy 319

Permian 319

Carboniferous 321

Devonian and Silurian 321

Ordovician and Cambrian 325

Chapter 9 Evolution 327

Introduction and General Principles 327

Features of the Deep-Sea Microfossil Record Relevant to the Study of Evolution 330

Microevolution 331

Pattern and Processes 332

Examples of Microevolution 333

Cladogenesis 333

Anagenesis 339

Extinction 344

Hybridization 344

Macroevolution 346

Definitions and Theory 346

Theories of Diversity and Evolution 348

Macroevolutionary Patterns in Radiolaria 349

Origin of Radiolarians 349

Origin of Collodaria and Colonial Radiolaria 352

Origin of Higher Taxa within Radiolaria – General Comments 354

Diversity History of Radiolarians 354

Methods of Diversity Reconstruction 354

Other Problems of Diversity Reconstruction 358

Data for Diversity Reconstruction 358

Global Phanerozoic Diversity 358

Paleozoic 363

Mesozoic 364

Cretaceous–Tertiary Boundary 368

Cenozoic 372

Other Aspects of Cenozoic Radiolarian Macroevolutionary Change 382

Phanerozoic Diversity – A More Modest View 386

Summary Discussion 388

References 393

Index 461