Polycystine radiolaria are exclusively marine protists and are found in all ocean waters, from polar regions to the tropics, and at all water depths. There are approximately 600 distinct described living species and several thousand fossil species of polycystines. Radiolarians in general, and polycystines in particular, have recently been shown to be a major component of the living plankton and important to the oceanic carbon cycle. As fossils radiolarians are also fairly common, and often occur in sediments where other types of fossils are absent. This has made them very valuable for certain types of geologic research, particularly estimating the geologic age of the sediments containing them, and as guides to past oceanic water conditions. As our current understanding of the biology, and even taxonomy of the living fauna is still very incomplete, evolutionary studies based on living polycystines are still rare. However, the common occurrence of numerous specimens for many species, and in a wide variety of oceanic environments, provides an excellent opportunity to study the processes of biologic evolution in the fossil record.
Paleobiology of the Polycystine Radiolaria is the first major book on radiolarians to appear in the western literature since 2001. Focusing on living and fossil siliceous shelled radiolarians, it is notable for its emphasis not upon morphologic or taxonomic detail but on concepts and applications. The book attempts to provide a balanced, critical review of what is known of the biology, ecology, and fossil record of the group, as well as their use in evolutionary, biostratigraphic and paleoceanographic research. Full chapters on the history of study, and molecular biology, are the first ever in book form.
Written for an audience of advanced undergraduate to doctoral students, as well as for a broad range of professionals in the biological and Earth sciences, Paleobiology of the Polycystine Radiolaria summarizes current understanding of the marine planktonic protist group polycystine radiolaria, both in living and fossil form.
Inhoudsopgave
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
Over de auteur
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 Ph D 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.
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