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 xiAcknowledgements xvChapter 1 History 1Introduction 1Scientific Context 4Early Studies (First Half of the Nineteenth Century) 8C.G. Ehrenberg and J. Müller 8Second Half of the Nineteenth Century to ca. 1920 13E. Haeckel and his Disciples 13Legacy of Early Studies 16Early Twentieth Century (ca. 1920–1940) 17The Early New Period (ca. 1940–1970) 20The Origins of Radiolarian Biostratigraphy: 1940s to 1950s 20Deep-Sea Drilling 21Taxonomy 25Biology 27Mid New Period (1970–2000) 28Current Period (2000-Present) 37Chapter 2 Biology 41General Characteristics of Planktonic Protist Biology 41Physical Characteristics of the Pelagic Ocean 42Plankton Taxa 46Ecologic and Behavioral Constraints due to Small Body Size 46Basic Radiolarian Cellular Structure 48Skeleton 53Skeleton Formation and Growth 55Size 59Colonial Forms 59Life Cycle 60Longevity 62Motility 63Feeding 63Predators 65Abundance and Role in Carbon Cycle 66Symbiosis 67Bioluminescence 68Summary 69Chapter 3 Ecology 71Introduction 71Biogeography 75Vertical Distribution 83Tropical Submergence 86Longitudinal Gradients and Upwelling Assemblages 89Latitudinal Gradients 90Coastal Gradients 90Seasonal Variability 91Interannual Variability 93Chapter 4 Genetics 95Introduction 95Molecular Phylogenetic Position of “Radiolarians” within Eukaryotes 96Molecular Studies of Radiolarian’s Position within Eukaryotes 97Relationships of Radiolarian Clades 98Origination Times of Radiolarian Clades 102Family-Level Phylogeny 102Spumellaria (Shell-Bearing Radiolarians) 105Collodaria (Colonial or Naked Radiolarians) 105Nassellaria 106Acantharia 107Microevolution of Radiolaria 107Diversity of Pico-Radiolarian Material 111Transcriptomics of Radiolaria 112Methodology 113DNA Extraction 114Reproductive Cell Method 114Dissecting Cell Method 114PCR 114Summary 114Chapter 5 Taxonomy and Fossil Record 117Introduction 117PART 1 - Radiolarian Taxonomy 118Principles of Species-Level Taxonomy 118Rules for Describing and Naming Species 121Current Status of Descriptive Radiolarian Taxonomy 124Principles of Higher-Level Taxonomy 129Haeckel and the Beginnings of Higher-Level Radiolarian Taxonomy 129Biologic Systematics 132Higher-Level Taxonomy in Radiolaria 134The Observational Basis of Taxonomy: Structures of the Radiolarian Shell 136Higher-Level Taxonomy in this Book 139Formal Classification of Polycystina 143Cenozoic Taxa 143Order Spumellaria Ehrenberg 1876 143Family Actinommidae Haeckel 1862 145Family Heliodiscidae Haeckel 1881 149Family Coccodiscidae Haeckel 1862, emend. Sanfilippo and Riedel 1980 151Family Pyloniidae Haeckel 1881 153Family Lithelidae Haeckel 1862 155Family Tholonidae Haeckel 1887 156Family Spongodiscidae Haeckel 1862 156Order Nassellaria Ehrenberg 1876 160Family Plagiacanthidae Hertwig 1879 162Family Trissocyclidae (Haeckel) Goll 1968[superfamily Acanthodesmiacea] 163Family Theoperidae Haeckel 1881 163Family Artostrobiidae Riedel 1967 167Family Pterocoryithidae (Haeckel) Moore 1972 167Family Carpocaniidae (Haeckel) Riedel, 1967 [Carpocaniinae] 171Family Cannobotryidae Haeckel, 1881 173Superfamily Collodaria 173Family Collosphaeridae Müller, 1858 175Family Sphaerozoidae Haeckel, 1862 175Family Collophidiidae Biard and Suzuki, in Biard et al., 2015 177Order Entactinaria 183Family Orosphaeridae Haeckel, 1887 183Family Saturnalidae Deflandre 1953 184Mesozoic and Paleozoic Taxa 185Species-Level Variation in Radiolaria 185PART 2 - Summary of the Radiolarian Fossil Record 193Cambrian and Ordovician 194Silurian to the Lower Carboniferous 195Late Paleozoic to Late Mesozoic Siliceous Sedimentation 196Mass Extinctions at the End of the Paleozoic Era 197Basal Mesozoic Scarcity of Radiolarian Fossils and Faunal Turnover (Early Triassic) 200Triassic 201Triassic–Jurassic Boundary Mass Extinction 204Jurassic 205Early and Middle Jurassic Radiolaria 205Late Jurassic–Early Cretaceous 208Cretaceous 208The K/T Extinction Event and Early Paleocene 212Cenozoic 214Chapter 6 Preservation and Methods 217Introduction 217Preservation 218Geographic Variation in Preservation 222Diagenesis 222Loss of Rock Record 224Differences between Modern and Ancient Oceans 224Quality of Radiolarian Fossil Record 225Methods 227Collecting Material from the Water Column 228Collecting Sediments 231Collecting Lithified Material from Sections on Land 236Recovering Radiolarians from Samples 238Extracting Radiolarians with Intact Protoplasm 238Extracting Radiolarian Skeletons 238Separation of Radiolarians from other Chemically Resistant Similar-Sized Components of Residue 242Mounting Radiolarians 243Live Preparations 245Dissection and Serial Sectioning 246Imaging Radiolarians 247Visualization (enhanced imagery) 248Morphometrics 249Automatic Identification 249Chapter 7 Paleoceanography 253Introduction 253Radiolarians as Tracers of Water Masses 259Assemblage-Based Methods of Paleoceanographic Analysis 259Non-temperature Uses of Assemblage Analyses 268Radiolarians in Bulk: Summary Indices and Non-Taxonomic Uses of Radiolarians in Paleoceanography 273Chapter 8 Radiolarian Biostratigraphy 281Introduction 281Biostratigraphy in Shallow Marine Rocks: General Aspects 283Biostratigraphy in Deep-Sea Sediment Sections 285Other Types of Geochronologic Information 287Radiometric Dating and Absolute Age 287Paleomagnetic Stratigraphy 288Stable Isotope Stratigraphy 290Cyclostratigraphy 291Quantitative Biostratigraphy 292Cenozoic Radiolarian Stratigraphy 295History of Development 296Tropical Cenozoic Radiolarian Stratigraphy 297Subtropical North Atlantic to Arctic 299North Pacific 302Southern Ocean 305History 305Characteristics 307Important Sections 307Important Species 307Mesozoic Radiolarian Stratigraphy 308Cretaceous 308Europe and Southwest North America 311Low-Latitude Western part of Mesotethys 311Mid-Ltitude Northern Part of Mesotethys 311Russian Epicontinental Seas 312East Margin of the Mid-Latitude Pacific 312Northwest Pacific 312Other Regions 313The Jurassic–Cretaceous Boundary(Tithonian–Berriasian Boundary) 313Jurassic 314Middle and Late Jurassic 314Lower Jurassic 316Triassic–Jurassic Boundary 316Triassic 316Latest Triassic (Rhaetian) 317Carnian and Norian 318Late Olenekian to Ladinian 318Basal Triassic (Induan) and Permian–Triassic (P–T) boundary 318Paleozoic Radiolarian Stratigraphy 319Permian 319Carboniferous 321Devonian and Silurian 321Ordovician and Cambrian 325Chapter 9 Evolution 327Introduction and General Principles 327Features of the Deep-Sea Microfossil Record Relevant to the Study of Evolution 330Microevolution 331Pattern and Processes 332Examples of Microevolution 333Cladogenesis 333Anagenesis 339Extinction 344Hybridization 344Macroevolution 346Definitions and Theory 346Theories of Diversity and Evolution 348Macroevolutionary Patterns in Radiolaria 349Origin of Radiolarians 349Origin of Collodaria and Colonial Radiolaria 352Origin of Higher Taxa within Radiolaria – General Comments 354Diversity History of Radiolarians 354Methods of Diversity Reconstruction 354Other Problems of Diversity Reconstruction 358Data for Diversity Reconstruction 358Global Phanerozoic Diversity 358Paleozoic 363Mesozoic 364Cretaceous–Tertiary Boundary 368Cenozoic 372Other Aspects of Cenozoic Radiolarian Macroevolutionary Change 382Phanerozoic Diversity – A More Modest View 386Summary Discussion 388References 393Index 461

“Paleobiology of the Polycystine Radiolaria is well worth the purchase price and should be in the personal library of all protistologists working on marine forms.” Journal of Eukaryotic Microbiology“A welcome addition to the literature in a field that is rich in potential for interdisciplinary research.” Journal of Plankton Research