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The Paleozoic Era: Diversification of Plant and Animal Life 7 Published in 2011 by Britannica Educational Publishing (a trademark of Encyclopædia Britannica, Inc.) in association with Rosen Educational Services, LLC 29 East 21st Street, New York, NY 10010. Copyright © 2011 Encyclopædia Britannica, Inc. Britannica, Encyclopædia Britannica, and the Thistle logo are registered trademarks of Encyclopædia Britannica, Inc. All rights reserved. Rosen Educational Services materials copyright © 2011 Rosen Educational Services, LLC. All rights reserved. Distributed exclusively by Rosen Educational Services. For a listing of additional Britannica Educational Publishing titles, call toll free (800) 237-9932. First Edition Britannica Educational Publishing Michael I. Levy: Executive Editor J.E. Luebering: Senior Manager Marilyn L. Barton: Senior Coordinator, Production Control Steven Bosco: Director, Editorial Technologies Lisa S. Braucher: Senior Producer and Data Editor Yvette Charboneau: Senior Copy Editor Kathy Nakamura: Manager, Media Acquisition John P. Ra erty: Associate Editor, Life and Earth Sciences Rosen Educational Services Alexandra Hanson-Harding: Senior Editor Nelson Sá: Art Director Cindy Reiman: Photography Director Matthew Cauli: Designer, Cover Design Introduction by Catherine Vanderhoof Library of Congress Cataloging-in-Publication Data The Paleozoic era : diversifi cation of plant and animal life / edited by John P. Ra erty. p. cm.—(The geologic history of Earth) “In association with Britannica Educational Publishing, Rosen Educational Services.” Includes bibliographical references and index. ISBN 978-1-61530-196-6 (eBook) 1. Geology, Stratigraphic—Paleozoic. 2. Paleoecology—Paleozoic. I. Ra erty, John P. QE654.P2413 2010 561'.112—dc22 2009051123 On the cover: Plants such as ferns (above) and ancient creatures such as the trilobite (below) were common during the Paleozoic era. SSPL via Getty Images (above); Sinclair Stammers/Photolibrary/Getty Images (below and pages 5, 29, 36, 93, 138, 187, 245, 285, 325, 326, 329, 331) On page 18: Fallen pine trees in Emerald Lake, refl ecting the Burgess Shale, British Columbia, Canada. Shutterstock.com CONTENTS Introduction 18 Chapter 1: An Overview of Paleozoic Time 29 Paleozoic Life 30 Paleozoic Geography 32 31 Chapter 2: The Cambrian Period 36 The Cambrian Environment 38 Paleogeography 38 Paleoclimate 45 Cambrian Life 46 The Fossil Record of the Precambrian-Cambrian Transition 47 Photosynthetic Organisms 51 Fauna 52 Deposits with Soft-Bodied Organisms 56 Cambrian Extinction Events 59 Signifi cant Cambrian Life-Forms 60 Burgess Shale 60 57 Conodonts 61 Denticles 64 Graptolites 65 Lamp Shells 65 Lingulids 78 Obolus 78 Stromatoporida 79 Trilobites 79 Cambrian Geology 82 Types and Distribution 82 58 The Boundaries and 94 Subdivisions of the Cambrian System 85 The Economic Significance of Cambrian Deposits 87 The Correlation of Cambrian Strata 88 Stages of the Cambrian Period 90 Fortunian Stage 90 Drumian Stage 91 Guzhangian Stage 91 Paibian Stage 92 106 Chapter 3: The Ordovician Period 93 The Ordovician Environment 93 Paleogeography 96 Sea Level 97 The Circulation of the Ocean 97 Plate Tectonics 99 Paleoclimate 101 Ordovician Life 104 107 Marine Organisms 104 Microfossils and Plankton 104 Invertebrates 105 Early Fishes 107 Reefs 108 Terrestrial Organisms 109 Earliest Land Plants 109 Animals 110 The Ordovician Radiation 110 The Exploitation of Habitats 112 Causes of the Ordovician Radiation 114 111 Ordovician Extinction Events 114 The Mass Extinction at the End of the Ordovician 114 Regional Extinctions Within the Ordovician 116 Faunal Provinces 117 Significant Ordovician Life-Forms 118 Bellerophon 118 Bumastus 119 Byssonychia 119 Calymene 119 Ceraurus 119 Climacograptus 120 Clonograptus 120 Constellaria 120 Cryptolithus 120 Cryptostomata 121 Cystoids 121 Didymograptus 121 Diplograptus 122 Echinosphaerites 122 Favosites 122 Hallopora 122 Halysites 123 Hesperorthis 123 Horn Corals 123 Isotelus 124 Leptaena 124 Lituites 124 Lophospira 124 Loxonema 125 139 Maclurites 125 Modiolopsis 125 Neoprioniodiform 125 Phyllograptus 126 Platystrophia 126 Plectoceras 126 Prasopora 126 Resserella 127 Rhynchotrema 127 Streptelasma 127 Strephomena 127 Tabulata 128 Tetragraptus 128 Trepostomata 128 Trochonema 128 Ordovician Geology 129 Significant Geologic Events 129 The Economic Significance of Ordovician Deposits 131 The Major Subdivisions of the Ordovician System 132 The Correlation of Ordovician Strata 132 Stages of the Ordovician Period 133 Tremadocian Stage 134 Floian Stage 134 Dapingian Stage 135 Darriwilian Stage 135 Sandbian Stage 136 Katian Stage 136 Hirnantian Stage 137 Chapter 4: The Silurian Period 138 The Silurian Environment 138 Paleogeography 140 Laurentia 141 Baltica 142 Siberia, Kazakhstania, and Other Continents 143 Gondwana 143 Paleoclimate 144 Silurian Life 146 Pentamerid Communities 147 Reef Mounds and Coral Biostromes 149 Fishes 150 Vascular Land Plants 151 Silurian Extinction Events 151 Significant Silurian Life-Forms 152 Atrypa 153 Baragwanathia 153 Birkenia 154 Chonetes 154 Conchidium 154 Cyathocrinites 155 Cystiphyllum 155 Deiphon 155 Eospirifer 155 Giant Water Scorpions 156 Goniophora 157 Leptodesma 157 Monograptus 157 Nuculana 158 Ozarkodiniforms 158 Phacops 158 Platyceras 158 Preferns 159 Rhynchotreta 160 Silurian Geology 160 The Economic Significance of Silurian Deposits 161 The Major Subdivisions of the Silurian System 163 147 Significant Geologic 189 Events 164 The Effects of Late Ordovician Glaciation 164 Silurian Sea Level 165 The Occurance and Distribution of Silurian Deposits 166 Evaporites 167 Clastic Wedges 168 Platform Margins 170 Tillites 171 Volcanic Rocks 172 The Correlation of Silurian Strata 173 Graptolites 173 Brachiopods 174 Conodonts 175 Isotope Stratigraphy 176 Establishing Silurian Boundaries 177 The Work of Roderick Murchison and Other Researchers 177 Geochemical Analyses 179 Silurian Boundaries 180 Stages of the Silurian Period 181 Rhuddanian Stage 182 Aeronian Stage 182 Telychian Stage 183 Sheinwoodian Stage 183 Homerian Stage 184 Gorstian Stage 184 Ludfordian Stage 185 Pridoli Series 185 Chapter 5: The Devonian Period 187 The Devonian Environment 190 Paleogeography 190 Paleoclimate 192 Devonian Life 192 Invertebrates 193 Vertebrates 195 Plants 197 Faunal Realms and Migrations 199 Devonian Extinction Events 200 Significant Devonian Life-Forms 202 Ammonoids 202 Antiarchs 203 Archaeopteris 203 Arctolepis 204 Arthrodires 204 196 Bactrites 205 Bothriolepis 205 Cephalaspis 206 Cheirolepis 206 Cladoselache 207 Climatius 207 Dinichthys 207 Dipterus 208 Endothyra 208 Eusthenopteron 209 Greenops 209 Heliophyllum 210 Hyenia 210 Ichthyostega 210 Monoplacophorans 211 Mucrospirifer 212 Osteolepis 212 Ostracoderms 213 Palaeospondylus 213 211 Placoderms 213 216 Polygnathiforms 214 Pteraspis 215 Rensselaeria 215 Rhipidistia 215 Rhynie Plants 216 Schizodus 217 Sphenophyllum 217 Stringocephalus 218 Stropheodonta 218 Theodossia 218 Tiktaalik Roseae 218 Tornoceras 220 Trepospira 220 Tropidoleptus 220 Worthenia 221 Xenacanthus 221 Devonian Geology 221 Significant Geologic Events 222 The Economic Significance of Devonian Deposits 224 222 The Major Subdivisions of the Devonian System 225 Establishing Devonian Boundaries 225 The Occurance and Distribution of Devonian Deposits 226 Sediment Types 229 Europe 230 Asia 234 Southern Hemisphere 235 North America 237 The Correlation of Devonian Strata 238 Stages of the Devonian Period 239 Lochkovian Stage 239 Pragian Stage 240 Emsian Stage 241 Eifelian Stage 241 Givetian Stage 242 258 Frasnian Stage 243 Famennian Stage 244 Chapter 6: The Carboniferous Period 245 The Carboniferous Environment 245 Paleogeography 246 Paleoclimate 247 Carboniferous Life 248 Invertebrates 249 Plants 251 Fishes 252 Amphibians and Early Reptiles 253 Significant Carboniferous Life-Forms 254 Calamites 254 Composita 255 Cordaitales 255 Cryptoblastus 255 Dictyoclostus 256 Dielasma 256 Diplovertebron 256 Edaphosaurus 257 Euphemites 257 Fenestella 258 Fusulina 258 Fusulinella 259 Fusulinids 259 Gastrioceras 260 Labyrinthodonts 260 Lebachia 261 Lepidodendron 261 Linoproductus 262 Lophophyllum 263 Myalina 263 Neospirifer 263 262 Nuculopsis 264 Pentremites 264 Phillipsia 264 Platycrinites 264 Seed Ferns 265 Sigillaria 266 Wedekindellina 266 Carboniferous Geology 267 Significant Geologic Events 268 The Major Subdivisions of the Carboniferous System 269 The Economic Significance of Carboniferous Deposits 270 The Occurance and Distribution of Carboniferous Deposits 270 Mississippian Limestones 271 Pennsylvanian Cyclothems, Tillites, and Turbidites 273 The Correlation of Carboniferous Strata 276 265 Missippian Subsystem 276 Pennsylvanian Subsystem 277 Stages of the Carboniferous Period 278 Mississippian Subperiod 278 Tournaisian Stage 279 Vise´an Stage 280 Serpukhovian Stage 281 Pennsylvanian Subperiod 282 Bashkirian Stage 282 Moscovian Stage 283 Kasimovian Stage 283 Gzhelian Stage 284 Chapter 7: The Permian Period 285 The Permian Environment 286 Paleogeography 286 Paleoclimate 288 Permian Life 289 The Emergence of Important Reptiles 291 Permian Mass Extinction 293 Temperature Crises and the Permian Extinction 294 The Alteration of the Carbon Cycle 295 Other Potential Causes of the Permian Extinction 296 287 Significant Permian Life-Forms 296 Bradysaurus 396 Cacops 297 292 Captorhinus 297 Cynodonts 298 Diadectes 299 Dimetrodon 300 Eryops 300 Glossopteris 301 Leptodus 302 Limnoscelis 302 Mesosaurus 303 Moschops 303 Parafusulina 304 Paraschwagerina 304 Pseudoschwagerina 304 298 Schwaserina 305 Seymouria 305 Tapinocephalus 306
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