Table Of ContentPALEOBOTANY
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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
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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
