Paleobotany, Second Edition: The Biology and Evolution of Fossil Plants PDF
Preview Paleobotany, Second Edition: The Biology and Evolution of Fossil Plants
PALEOBOTANY This page intentionally left blank PALEOBOTANY The Biology and Evolution of Fossil Plants Second Edition THOMAS N. TAYLOR Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Research Center, The University of Kansas, Lawrence, Kansas EDITH L. TAYLOR Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Research Center, The University of Kansas, Lawrence, Kansas MICHAEL KRINGS Bayerische Staatssammlung für Paläontologie und Geologie und GeoBio-CenterLMU, Munich, Germany AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD PARIS • SAN DIEGO • SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 32 Jamestown Road, London NW 1 7BY, UK 525 B Street, Suite 1900, San Diego, CA 92101-4495, USA 360 Park Avenue South, New York, NY 10010-1710, USA Copyright © 2009, Elsevier Inc. All rights reserved No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means electronic, mechanical, photocopying, recording or otherwise without the prior written permission of the publisher. 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Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN: 978-0-12-373972-8 For information on all Academic Press publications visit our website at www.elsevierdirect.com Typeset by Charon Tec Ltd., A Macmillan Company. (www.macmillansolutions.com) Printed and bound in the USA 09 10 11 10 9 8 7 6 5 4 3 2 1 CONTENTS Preface xv Chemical Fossils 32 Acknowledgments xvii Ancient DNA 33 Mummifi cation 33 About the Authors xxi Amber 33 Summary Discussion 34 1 CHAPTER Palynology 34 Introduction to Paleobotany, Geochronology and Biostratigraphy 36 How Fossil Plants are Formed 1 Paleoecology 37 Absolute Dating 38 What Is Paleobotany? 1 Geologic Timescale 39 The Objectives of Paleobotany 2 Biological Correlation 40 Reconstructing the Plants 2 Systematics and Classifi cation 40 Evolution of Plant Groups 3 Nomenclature of Fossil Plants 41 Form and Function in Fossil Plants 4 Classifi cation of Organisms 42 Biostratigraphy and Correlation 4 Background Reading 42 Paleoecology: Plants in Their Environment 5 Determining Paleoclimate from Fossil Plants 6 Tree Rings 6 2 CHAPTER Nearest Living Relative 6 Precambrian Life 43 Leaf Physiognomy 7 Stomatal Index 7 The Origin of Life on Earth 44 Summary 7 Origin of Life: Theory and Biology 46 Preservation: How Plant Fossils are Formed and Earliest Record of Life on Earth 47 Preserved 8 Historical Background 47 Depositional Environments of Fossil Plants 8 Earliest Records of Life: Paleoarchean (3.6–3.2 Ga) 47 Compressions 10 Geochemistry 47 Cuticle 13 Microfossils (Body Fossils) 49 Biofi lms and Plant Fossil Preservation 16 Isua Greenstone Belt, Greenland 49 Electron Microscopy 17 Warrawoona Group, Australia 49 Confocal Microscopy 17 Barberton Greenstone Belt, South Africa 51 Maceration and Dégagement 17 Stromatolites 52 Other Techniques 18 Sedimentary Evidence 53 Coal and Charcoal 18 Mesoarchean–Neoarchean Life 54 Impressions 21 Conclusions: Archean Life 55 Molds and Casts 22 Oxygenation of the Earth (2.45–2.2 Ga) 57 Cellular Preservation 23 Proterozoic Life 59 Permineralization 25 Paleoproterozoic 59 Peel Technique 25 Origin of Eukaryotes 61 Coal Balls 27 Mesoproterozoic 64 Other Permineralizations 29 Earliest Multicellular Life 64 Petrifaction 30 Neoproterozoic 64 Unaltered Plant Material 30 Bitter Springs Biota 65 vv vi contents Stromatolites 66 Dinophyta (Dinofl agellates) 139 Other Microfossils 67 Heterokontophyta 141 Doushantuo Formation 70 Bacillariophyceae (Diatoms) 141 Conclusions 70 Dictyochophyceae (Silicofl agellates) 142 Xanthophyceae (Yellow-Green Algae) 142 3 CHAPTER Phaeophyceae (Brown Algae) 143 Fungi, Bacteria, and Lichens 71 Prymnesiophyta (Haptophytes) 144 Rhodophyta (Red Algae) 145 Fungi 71 Solenoporaceans 146 Earliest Fossil Fungi 73 Other Calcifi ed Red Algae 149 Systematics of Fungi 77 Corallinales 149 Chytridiomycota 77 Uncalcifi ed Red Algae 150 Zygomycota 82 Acritarcha (Acritarchs) 158 Glomeromycota 84 Ascomycota 90 5 CHAPTER Basidiomycota 93 Hornworts and Bryophytes 161 Other Fungal Remains 97 Fungal Life-History Strategies 98 Early Fossil Evidence 163 Saprotrophism 98 Anthocerotophyta (Hornworts) 165 Parasitism 99 Bryophyta (Bryophytes) 166 Mutualism 103 Marchantiophytina (Liverworts or Hepatophytes) 167 Fungi–Animal Interactions 105 Bryophytina (Mosses) 174 Geologic Activities of Fungi 107 Epiphyllous Fungi 108 CHAPTER6 Fungal Spores 111 The Move to the Land 179 Fungal-like Organisms 112 Enigmatic Organisms 180 Peronosporomycetes (Oomycota) 112 Nematophytes 180 Eubacteria and Archaea 112 Prototaxites 180 Archaea 113 Nematothallus 183 Eubacteria 113 Nematoplexus 183 Cyanobacteria 115 Nematasketumdiversiforme 183 Lichens 117 Pachytheca 184 Spongiophytaceae 185 4 CHAPTER Spongiophyton 185 Algae 121 Orestovia 186 Chlorophyta (Green Algae) 123 Other Enigmatic Organisms 186 Prasinophyceae 124 Protosalvinia 186 Chlorophyceae 126 Parka 188 Volvocales 126 Isolated Fragments: Clues to the Transition to Land? 189 Tetrasporales 126 Cuticle and Cuticle-Like Material 189 Chlorococcales 127 Spores and Spore Tetrads 189 Ulvophyceae 128 Tubes 192 Dasycladales 128 Land Plant Ancestors 193 Receptaculitida and Cyclocrinales 130 The Transition to Land 194 Caulerpales 130 Anchorage and Water Uptake 194 Taxa Incertae Sedis 133 Structural Support and Water Transport 195 Charophyceae 133 Protection Against Desiccation and Radiation 195 Charales 134 Gas Exchange 195 Zygnematales 138 Reproduction on Land 196 Euglenophyta 138 Life History Biology 196 vii Contents Homologous Theory 196 Rhyniophytes 227 Antithetic Theory 196 Rhynie Chert Plants 228 Animals 198 Aglaophytonmajor 229 A Fungal Partner 198 Rhynia Gwynne-vaughanii 235 Conclusion 199 Horneophyton lignieri 237 Asteroxylon mackiei 238 Nothia aphylla 239 7 CHAPTER Trichopherophyton teuchansii 241 Introduction to Vascular Plant Morphology Ventarura lyonii 241 and Anatomy 201 Gametophyte Generation 241 Other Rhyniophytes 246 Plant Organography 202 Discussion: Rhyniophyte Evolution 251 Cell Types 203 Zosterophyllophytes 252 Parenchyma 203 Zosterophyll Evolution 259 Collenchyma 203 Trimerophytes 259 Sclerenchyma 203 Trimerophyte Evolution 262 Tracheary Elements 204 Early Land Plant Evolution 263 Tracheids 204 Vessel Elements 206 9 Sieve Elements 206 CHAPTER Lycophyta 265 Plant Tissues and Primary Growth 207 Xylem Tissue 207 Evolution of the Microphyll 267 Phloem Tissue 207 Drepanophycales 268 Meristems 208 Protolepidodendrales 271 Epidermis 208 Lepidodendrales 279 Cuticle 209 Vegetative Features 282 Stomata 209 Stem Surface and Leaf Bases 282 Trichomes 210 Stem Anatomy 285 Anatomy of Stems and Roots 210 Cortical Tissues 286 Arrangement of Primary Tissues 210 Stem Development 287 Primary Xylem Maturation Patterns 212 Leaves 289 Secondary Development 212 Underground Organs 289 Vascular Cambium 212 Development of Underground Organs 293 Cork Cambium (Phellogen) 213 Reproductive Biology 294 Secondary Xylem 214 Microsporangiate and Bisporangiate Cones 295 Secondary Phloem 216 Megasporangiate Cones 297 Stele Types 216 Gametophytes 302 Primitive Vascular Plants (Vascular Cryptogams) 216 Sigillariaceae 303 Seed Plants 219 Leaf Bases 304 Leaf Morphology and Anatomy 221 Leaves 305 Leaf Anatomy 221 Stem Structure 305 Leaf Evolution 222 Underground Organs 306 Further Reading 222 Reproductive Biology 306 Other Lepidodendrid Genera 307 8 Lycopodiales 310 CHAPTER Early Land Plants with Conducting Selaginellales 312 Pleuromeiales 316 Tissue 223 Isoetales 320 Conducting Elements in Early Land Plants 224 Putative Lycopsids 325 History of Discovery 225 Conclusions 326 viii contents 10 Zygopterid Evolution 417 CHAPTER Sphenophytes 329 Marattiales 418 Psaroniaceae: Vegetative Features 418 Pseudoborniales 331 Psaronius Plant 418 Sphenophyllales 332 Other Stem Taxa 425 Devonian Sphenophyllales 333 Psaroniaceae: Reproductive Features 425 Sphenophyllum 334 Paleozoic Compression Taxa 431 Leaves 334 Mesozoic Marattialeans 433 Stem Anatomy 335 Marattialean Evolution 434 Roots 337 Ophioglossales 435 Reproductive Biology 337 Leptosporangiate Ferns 436 Other Sphenophyllales 338 Osmundales 436 Ecology 341 Paleozoic Stem Taxa 437 Equisetales 342 Guaireaceae 438 Calamitaceae 343 Mesozoic and Cenozoic Stem Taxa 438 Archaeocalamites 343 Sterile and Fertile Foliage 440 Calamites 345 Osmundalean Evolution 442 Pith Casts 349 Botryopteridaceae 443 Stem Anatomy 350 Vegetative Organs 443 Extraxylary Tissues 352 Reproductive Organs 446 Growth and Development 352 Other Genera 449 Roots 353 Anachoropteridaceae 449 Leaves 354 Kaplanopteridaceae 451 Other Calamitean Leaves 357 Psalixochlaenaceae 452 Reproductive Biology 358 Sermayaceae 453 Spores 366 Tedeleaceae 454 Tchernoviaceae and Gondwanostachyaceae 368 Skaaripteridaceae 457 Vegetative Body 368 Tempskyaceae 457 Reproductive Biology 369 Schizaeaceae 459 Equisetaceae 371 Hymenophyllaceae 462 Forms with Uncertain Affi nities 376 Gleicheniaceae 462 Sphenophyte Evolution 379 Dicksoniaceae 464 Cyatheaceae 465 11 Matoniaceae 466 CHAPTER Ferns and Early Fernlike Plants 383 Loxsomataceae 469 Dipteridaceae 469 Evolution of the Megaphyll 386 Polypodiales 470 Cladoxylopsida 387 Salviniales 472 Pseudosporochnales 388 Marsileaceae 472 Calamophyton Plant 396 Salviniaceae 473 Iridopteridales 398 Conclusions 476 Phylogenetic Position of the Cladoxylopsids 400 Early Fernlike Plants 401 12 CHAPTER Rhacophytales 401 Progymnosperms 479 Rhacophyton 402 Other Taxa 403 Archaeopteridales 480 Systematics of the Rhacophytales 404 Archaeopteris Leaves 481 Coenopterid Ferns 405 Archaeopterid Reproduction 483 Stauropteridales 405 Callixylon Stems 484 Zygopteridales 408 Other Archaeopterids 487 ix Contents Aneurophytales 489 Conostoma 556 Aneurophyton 489 Coronostoma 557 Tetraxylopteris 489 Physostoma 557 Triloboxylon 491 Tyliosperma 558 Rellimia 492 Calathospermum 558 Other Aneurophytes 494 Gnetopsis 559 Protopityales 496 Megatheca 559 Noeggerathians 497 Other Lyginopterids: Pollen Organs 560 Progymnosperm Evolution 501 Incertae Sedis 563 Lyginopterid Evolution 565 13 CHAPTER Medullosales 566 Origin and Evolution of the Seed Habit 503 Stems 566 Medullosa 566 Homospory, Heterospory, and the Seed Habit 503 Other Stem Taxa 569 Homospory 503 Leaves (Fronds) 570 Heterospory 504 Roots 572 Sporangia 504 Growth Habit 572 Endospory 507 Seeds 573 Lycopsid Heterospory 508 Pollen organs 581 Seed Habit 508 Pollen 590 Evolution of the Integument 509 Medullosan Evolution 591 Evolution of Pollen Capture 510 Callistophytales 593 Pollen 511 Vegetative Organs 594 Cupules 511 Reproductive Structures 595 Cupulate Devonian Seeds 511 Callistophytalean Evolution 598 Reproductive Biology 517 Glossopteridales 598 Carboniferous Seeds 518 Leaves 599 Pollen Chamber Function 523 Glossopteris 599 Microgametophytes 524 Gangamopteris 603 Diversity of Early Seeds 525 Other Leaf Types 603 Paleozoic Seeds with Embryos 526 Stems and Roots 605 14 Ovulate Reproductive Structures 606 CHAPTER Paleozoic Seed Ferns 529 Permineralized Forms 606 Impression–Compression Specimens 609 Calamopityales 531 What is the Glossopterid Ovulate Structure? 614 Buteoxylonales 539 Pollen Organs 616 Lyginopteridales 540 Glossopteris Habit and Habitat 618 Lyginopteris Plant 540 Phylogenetic Position 618 Vegetative Organs 540 Reproductive Structures 542 15 CHAPTER Other Lyginopterids: Vegetative Remains 546 Mesozoic Seed Ferns 621 Heterangium 547 Microspermopteris 550 Caytoniales 622 Schopfi astrum 550 Sagenopteris 622 Pitys 551 Caytonanthus 623 Devonian–Mississippian Taxa 552 Caytonia 624 Problematic Lyginopterids 554 Rufl orinia and K talenia 626 Other Lyginopterids: Seeds and Cupules 555 Corystospermales 627 Sphaerostoma 556 Foliage 627 Salpingostoma 556 Stems 630
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