Paleobotany (eBook)

Front Cover 1
Paleobotany: The Biology and Evolution of Fossil Plants 4
Copyright Page 5
Contents 6
Preface 16
Acknowledgments 18
About the Authors 22
CHAPTER 1 Introduction to Paleobotany, How Fossil Plants are Formed 24
What Is Paleobotany? 24
The Objectives of Paleobotany 25
Reconstructing the Plants 25
Evolution of Plant Groups 26
Form and Function in Fossil Plants 27
Biostratigraphy and Correlation 27
Paleoecology: Plants in Their Environment 28
Determining Paleoclimate from Fossil Plants 29
Tree Rings 29
Nearest Living Relative 29
Leaf Physiognomy 30
Stomatal Index 30
Summary 30
Preservation: How Plant Fossils are Formed and Preserved 31
Depositional Environments of Fossil Plants 31
Compressions 33
Cuticle 36
Biofilms and Plant Fossil Preservation 39
Electron Microscopy 40
Confocal Microscopy 40
Maceration and Dégagement 40
Other Techniques 41
Coal and Charcoal 41
Impressions 44
Molds and Casts 45
Cellular Preservation 46
Permineralization 48
Peel Technique 48
Coal Balls 50
Other Permineralizations 52
Petrifaction 53
Unaltered Plant Material 53
Chemical Fossils 55
Ancient DNA 56
Mummification 56
Amber 56
Summary Discussion 57
Palynology 57
Geochronology and Biostratigraphy 59
Paleoecology 60
Absolute Dating 61
Geologic Timescale 62
Biological Correlation 63
Systematics and Classification 63
Nomenclature of Fossil Plants 64
Classification of Organisms 65
Background Reading 65
CHAPTER 2 Precambrian Life 66
The Origin of Life on Earth 67
Origin of Life: Theory and Biology 69
Earliest Record of Life on Earth 70
Historical Background 70
Earliest Records of Life: Paleoarchean (3.6–3.2 Ga) 70
Geochemistry 70
Microfossils (Body Fossils) 72
Isua Greenstone Belt, Greenland 72
Warrawoona Group, Australia 72
Barberton Greenstone Belt, South Africa 74
Stromatolites 75
Sedimentary Evidence 76
Mesoarchean–Neoarchean Life 77
Conclusions: Archean Life 78
Oxygenation of the Earth (2.45–2.2 Ga) 80
Proterozoic Life 82
Paleoproterozoic 82
Origin of Eukaryotes 84
Mesoproterozoic 87
Earliest Multicellular Life 87
Neoproterozoic 87
Bitter Springs Biota 88
Stromatolites 89
Other Microfossils 90
Doushantuo Formation 93
Conclusions 93
CHAPTER 3 Fungi, Bacteria, and Lichens 94
Fungi 94
Earliest Fossil Fungi 96
Systematics of Fungi 100
Chytridiomycota 100
Zygomycota 105
Glomeromycota 107
Ascomycota 113
Basidiomycota 116
Other Fungal Remains 120
Fungal Life-History Strategies 121
Saprotrophism 121
Parasitism 122
Mutualism 126
Fungi–Animal Interactions 128
Geologic Activities of Fungi 130
Epiphyllous Fungi 131
Fungal Spores 134
Fungal-like Organisms 135
Peronosporomycetes (Oomycota) 135
Eubacteria and Archaea 135
Archaea 136
Eubacteria 136
Cyanobacteria 138
Lichens 140
CHAPTER 4 Algae 144
Chlorophyta (Green Algae) 146
Prasinophyceae 147
Chlorophyceae 149
Volvocales 149
Tetrasporales 149
Chlorococcales 150
Ulvophyceae 151
Dasycladales 151
Receptaculitida and Cyclocrinales 153
Caulerpales 153
Taxa Incertae Sedis 156
Charophyceae 156
Charales 157
Zygnematales 161
Euglenophyta 161
Dinophyta (Dinoflagellates) 162
Heterokontophyta 164
Bacillariophyceae (Diatoms) 164
Dictyochophyceae (Silicoflagellates) 165
Xanthophyceae (Yellow-Green Algae) 165
Phaeophyceae (Brown Algae) 166
Prymnesiophyta (Haptophytes) 167
Rhodophyta (Red Algae) 168
Solenoporaceans 169
Other Calcified Red Algae 172
Corallinales 172
Uncalcified Red Algae 173
Acritarcha (Acritarchs) 181
CHAPTER 5 Hornworts and Bryophytes 184
Early Fossil Evidence 186
Anthocerotophyta (Hornworts) 188
Bryophyta (Bryophytes) 189
Marchantiophytina (Liverworts or Hepatophytes) 190
Bryophytina (Mosses) 197
CHAPTER 6 The Move to the Land 202
Enigmatic Organisms 203
Nematophytes 203
Prototaxites 203
Nematothallus 206
Nematoplexus 206
Nematasketum diversiforme 206
Pachytheca 208
Spongiophytaceae 208
Spongiophyton 208
Orestovia 209
Other Enigmatic Organisms 209
Protosalvinia 209
Parka 211
Isolated Fragments: Clues to the Transition to Land? 212
Cuticle and Cuticle-Like Material 212
Spores and Spore Tetrads 212
Tubes 215
Land Plant Ancestors 216
The Transition to Land 217
Anchorage and Water Uptake 217
Structural Support and Water Transport 218
Protection Against Desiccation and Radiation 218
Gas Exchange 218
Reproduction on Land 219
Life History Biology 219
Homologous Theory 219
Antithetic Theory 219
Animals 221
A Fungal Partner 221
Conclusion 222
CHAPTER 7 Introduction to Vascular Plant Morphology and Anatomy 224
Plant Organography 225
Cell Types 226
Parenchyma 226
Collenchyma 226
Sclerenchyma 226
Tracheary Elements 227
Tracheids 227
Vessel Elements 229
Sieve Elements 229
Plant Tissues and Primary Growth 230
Xylem Tissue 230
Phloem Tissue 230
Meristems 231
Epidermis 231
Cuticle 232
Stomata 232
Trichomes 233
Anatomy of Stems and Roots 233
Arrangement of Primary Tissues 233
Primary Xylem Maturation Patterns 235
Secondary Development 235
Vascular Cambium 235
Cork Cambium (Phellogen) 236
Secondary Xylem 237
Secondary Phloem 239
Stele Types 239
Primitive Vascular Plants (Vascular Cryptogams) 239
Seed Plants 242
Leaf Morphology and Anatomy 244
Leaf Anatomy 244
Leaf Evolution 245
Further Reading 245
CHAPTER 8 Early Land Plants with Conducting Tissue 246
Conducting Elements in Early Land Plants 247
History of Discovery 248
Rhyniophytes 250
Rhynie Chert Plants 251
Aglaophyton major 252
Rhynia Gwynne-vaughanii 258
Horneophyton lignieri 260
Asteroxylon mackiei 261
Nothia aphylla 262
Trichopherophyton teuchansii 264
Ventarura lyonii 264
Gametophyte Generation 264
Other Rhyniophytes 269
Discussion: Rhyniophyte Evolution 274
Zosterophyllophytes 275
Zosterophyll Evolution 282
Trimerophytes 282
Trimerophyte Evolution 285
Early Land Plant Evolution 286
CHAPTER 9 Lycophyta 288
Evolution of the Microphyll 290
Drepanophycales 291
Protolepidodendrales 294
Lepidodendrales 302
Vegetative Features 305
Stem Surface and Leaf Bases 305
Stem Anatomy 308
Cortical Tissues 309
Stem Development 310
Leaves 312
Underground Organs 312
Development of Underground Organs 316
Reproductive Biology 317
Microsporangiate and Bisporangiate Cones 318
Megasporangiate Cones 320
Gametophytes 325
Sigillariaceae 326
Leaf Bases 327
Leaves 328
Stem Structure 328
Underground Organs 329
Reproductive Biology 329
Other Lepidodendrid Genera 330
Lycopodiales 333
Selaginellales 335
Pleuromeiales 339
Isoetales 343
Putative Lycopsids 348
Conclusions 349
CHAPTER 10 Sphenophytes 352
Pseudoborniales 354
Sphenophyllales 355
Devonian Sphenophyllales 356
Sphenophyllum 357
Leaves 357
Stem Anatomy 358
Roots 360
Reproductive Biology 360
Other Sphenophyllales 361
Ecology 364
Equisetales 365
Calamitaceae 366
Archaeocalamites 366
Calamites 368
Pith Casts 372
Stem Anatomy 373
Extraxylary Tissues 375
Growth and Development 375
Roots 376
Leaves 377
Other Calamitean Leaves 380
Reproductive Biology 381
Spores 389
Tchernoviaceae and Gondwanostachyaceae 391
Vegetative Body 391
Reproductive Biology 392
Equisetaceae 394
Forms with Uncertain Affinities 399
Sphenophyte Evolution 402
CHAPTER 11 Ferns and Early Fernlike Plants 406
Evolution of the Megaphyll 409
Cladoxylopsida 410
Pseudosporochnales 411
Calamophyton Plant 419
Iridopteridales 421
Phylogenetic Position of the Cladoxylopsids 423
Early Fernlike Plants 424
Rhacophytales 424
Rhacophyton 425
Other Taxa 426
Systematics of the Rhacophytales 427
Coenopterid Ferns 428
Stauropteridales 428
Zygopteridales 431
Zygopterid Evolution 440
Marattiales 441
Psaroniaceae: Vegetative Features 441
Psaronius Plant 441
Other Stem Taxa 448
Psaroniaceae: Reproductive Features 448
Paleozoic Compression Taxa 454
Mesozoic Marattialeans 456
Marattialean Evolution 457
Ophioglossales 458
Leptosporangiate Ferns 459
Osmundales 459
Paleozoic Stem Taxa 460
Guaireaceae 461
Mesozoic and Cenozoic Stem Taxa 461
Sterile and Fertile Foliage 463
Osmundalean Evolution 465
Botryopteridaceae 466
Vegetative Organs 466
Reproductive Organs 469
Other Genera 472
Anachoropteridaceae 472
Kaplanopteridaceae 474
Psalixochlaenaceae 475
Sermayaceae 476
Tedeleaceae 477
Skaaripteridaceae 480
Tempskyaceae 480
Schizaeaceae 482
Hymenophyllaceae 485
Gleicheniaceae 485
Dicksoniaceae 487
Cyatheaceae 488
Matoniaceae 489
Loxsomataceae 492
Dipteridaceae 492
Polypodiales 493
Salviniales 495
Marsileaceae 495
Salviniaceae 496
Conclusions 499
CHAPTER 12 Progymnosperms 502
Archaeopteridales 503
Archaeopteris Leaves 504
Archaeopterid Reproduction 506
Callixylon Stems 507
Other Archaeopterids 510
Aneurophytales 512
Aneurophyton 512
Tetraxylopteris 512
Triloboxylon 514
Rellimia 515
Other Aneurophytes 517
Protopityales 519
Noeggerathians 520
Progymnosperm Evolution 524
CHAPTER 13 Origin and Evolution of the Seed Habit 526
Homospory, Heterospory, and the Seed Habit 526
Homospory 526
Heterospory 527
Sporangia 527
Endospory 530
Lycopsid Heterospory 531
Seed Habit 531
Evolution of the Integument 532
Evolution of Pollen Capture 533
Pollen 534
Cupules 534
Cupulate Devonian Seeds 534
Reproductive Biology 540
Carboniferous Seeds 541
Pollen Chamber Function 546
Microgametophytes 547
Diversity of Early Seeds 548
Paleozoic Seeds with Embryos 549
CHAPTER 14 Paleozoic Seed Ferns 552
Calamopityales 554
Buteoxylonales 562
Lyginopteridales 563
Lyginopteris Plant 563
Vegetative Organs 563
Reproductive Structures 565
Other Lyginopterids: Vegetative Remains 569
Heterangium 570
Microspermopteris 573
Schopfiastrum 573
Pitys 574
Devonian–Mississippian Taxa 575
Problematic Lyginopterids 577
Other Lyginopterids: Seeds and Cupules 578
Sphaerostoma 579
Salpingostoma 579
Conostoma 579
Coronostoma 580
Physostoma 580
Tyliosperma 581
Calathospermum 581
Gnetopsis 582
Megatheca 582
Other Lyginopterids: Pollen Organs 583
Incertae Sedis 586
Lyginopterid Evolution 588
Medullosales 589
Stems 589
Medullosa 589
Other Stem Taxa 592
Leaves (Fronds) 593
Roots 595
Growth Habit 595
Seeds 596
Pollen organs 604
Pollen 613
Medullosan Evolution 614
Callistophytales 616
Vegetative Organs 617
Reproductive Structures 618
Callistophytalean Evolution 621
Glossopteridales 621
Leaves 622
Glossopteris 622
Gangamopteris 626
Other Leaf Types 626
Stems and Roots 628
Ovulate Reproductive Structures 629
Permineralized Forms 629
Impression–Compression Specimens 632
What is the Glossopterid Ovulate Structure? 637
Pollen Organs 639
Glossopteris Habit and Habitat 641
Phylogenetic Position 641
CHAPTER 15 Mesozoic Seed Ferns 644
Caytoniales 645
Sagenopteris 645
Caytonanthus 646
Caytonia 647
Ruflorinia and Ktalenia 649
Corystospermales 650
Foliage 650
Stems 653
Pollen Organs 654
Ovulate Structures 657
Petriellales 660
Peltaspermales 662
Foliage 662
Reproductive Organs and Whole-Plant Concepts 666
Conclusions 671
CHAPTER 16 Late Paleozoic and Mesozoic Foliage 674
Late Paleozoic Foliage 675
Adiantites 678
Alethopteris 679
Aneimites 680
Aphlebia 681
Alloiopteris 681
Botrychiopsis 682
Callipteridium 682
Cardiopteridium 683
Cardiopteris (Fryopsis) 683
Charliea 683
Cyclopteris 684
Dicksoniites 685
Discopteris 687
Eremopteris 687
Ginkgophytopsis 687
Kankakeea 688
Karinopteris, Mariopteris, and Pseudomariopteris 688
Lesleya 692
Linopteris, Reticulopteris, and Barthelopteris 692
Lobatopteris 694
Lonchopteridium and Lonchopteris 695
Megalopteris 695
Neuropteris sensu lato 696
Laveinopteris 697
Macroneuropteris 697
Margaritopteris 697
Neuralethopteris 697
Neurocallipteris 698
Neurodontopteris 698
Neuropteris sensu stricto 698
Paripteris 698
Sphenoneuropteris 699
Neuropterid Growth Habit 699
Blanzyopteris 699
Nothorhacopteris 700
Odontopteris and Lescuropteris 700
Pecopteris 702
Rhodea (=Rhodeopteridium) 703
Sphenopteris 703
Sphenopteris sensu stricto 705
Eusphenopteris 705
Spiropteris 706
Taeniopteris 706
Tinsleya 708
Triphyllopteris, Genselia, and Charbeckia 708
Mesozoic Foliage 708
Anomozamites 710
Cladophlebis 710
Coniopteris 711
Ctenis 712
Deltolepis and Cycadolepis 712
Dictyophyllum 712
Dictyozamites 712
Doratophyllum 713
Macrotaeniopteris 713
Matonidium 713
Mesodescolea 713
Nilssonia 713
Nilssoniopteris 714
Otozamites 716
Pachypteris, Komlopteris, and Thinnfeldia 718
Phlebopteris 719
Pseudoctenis 719
Pseudocycas 720
Pterophyllum 720
Ptilophyllum 721
Ptilozamites 722
Ruflorinia 722
Taeniozamites 723
Ticoa 723
Wingatea 723
Yabeiella 723
Zamites 724
CHAPTER 17 Cycadophytes 726
Cycadales 726
Leaves and Petioles 729
Stems 730
Paleozoic Reproductive Structures 732
Triassic Cycads 738
Jurassic Cycads 741
Pollination Biology 744
Discussion: Cycad Evolution 744
Bennettitales 745
Cycadeoidaceae 748
Stem Anatomy 748
Reproductive Structures 751
Development 753
Williamsoniaceae 755
Ovulate Structures 757
Pollen Organs 761
Discussion: Bennettitales 762
CHAPTER 18 Ginkgophytes 766
Paleozoic Record 767
Ginkgophyte Wood 770
Ginkgophyte Foliage 770
Pollen-Producing Structures 773
Ginkgophyte Plants 773
Ginkgoaceae 773
Karkeniaceae 775
Umaltolepidiaceae 775
Yimaiaceae 775
Schmeissneriaceae 776
Taxa with Uncertain Affinities 777
Conclusions 778
CHAPTER 19 Gymnosperms with Obscure Affinities 780
Gigantopteridales 781
Vegetative Remains 781
Reproductive Organs 785
Vojnovskyales 786
Czekanowskiales 788
Iraniales 791
Pentoxylales 791
Hermanophytales 796
Gnetales 798
Extant Genera 799
Ephedra 799
Gnetum 799
Welwitschia 799
Extant Reproductive Structures 800
Fossil Gnetophyte Pollen 800
Gnetophyte Megafossils 801
Putative Gnetophytes 804
Dirhopalostachyaceae 808
CHAPTER 20 Cordaitales 810
Vegetative Features 811
Stems 811
Foliage 814
Roots 817
Reproductive Features 818
Reproductive Organs 818
Seeds 821
Angaran Cordaites 824
Phylogenetic Position and Origin of the Cordaites 826
CHAPTER 21 Conifers 828
Early Conifers 829
Voltziales 830
Utrechtiaceae 831
Utrechtia 832
Ernestiodendron 832
Ortiseia 832
Otovicia 833
Moyliostrobus 834
Other Taxa 834
Thucydiaceae 837
Emporiaceae 838
Majonicaceae 839
Ullmanniaceae 842
Bartheliaceae 843
Other Voltzialeans 843
Ferugliocladaceae 846
Buriadiaceae 849
Pollen Cones 849
Summary Discussion: Voltzialeans 851
Coniferales 853
Palissyaceae 853
Cheirolepidiaceae 854
Summary Discussion: Cheirolepidiaceae 860
Podocarpaceae 861
Summary Discussion: Podocarpaceae 866
Araucariaceae 866
Summary Discussion: Araucariaceae 871
Cupressaceae 872
Cunninghamioideae 873
Taiwanioideae 874
Athrotaxoideae 874
Sequoioideae 875
Taxodioideae 877
Cupressoideae 880
Cupressaceous Wood 882
Summary Discussion: Cupressaceae 882
Sciadopityaceae 883
Pararaucariaceae 884
Pinaceae 884
Pinoideae 886
Genus Pinus 887
Pinus Wood 889
Larix 889
Piceoideae 890
Abietoideae 890
Summary Discussion: Pinaceae 891
Cephalotaxaceae 891
Taxaceae 892
Summary Discussion: Cephalotaxaceae and Taxaceae 892
Conclusions 893
CHAPTER 22 Flowering Plants 896
Angiosperm Origins 899
Origin of the Flower 900
Pseudanthial Theory 900
Euanthial Theory 901
Microsporangial Theories 901
Transitional–Combination Theory 901
Habit 902
Ecological Considerations 902
Site of Origin 903
Pre-Cretaceous Fossil Evidence 903
Sanmiguelia 904
Furcula 905
Problematospermum 906
Pre-Cretaceous Pollen 906
Dispersed Pollen 907
Early Angiosperm Evidence 908
Pollen 908
Pollen Evolution 912
Evidence from Leaves 912
Angiosperm Ancestors 916
Caytoniales 917
Czekanowskiales 918
Glossopteridales 918
Bennettitales 918
Pentoxylales 918
Gigantopteridales 918
Phylogenetic Analyses and Angiosperm Origins 918
Selected Angiosperm Families 920
Basal Angiosperms 921
Amborellaceae 921
Hydatellaceae 921
Archaefructaceae 921
Chloranthaceae 922
Nymphaeales 924
Nymphaeaceae 924
Austrobaileyales 925
Austrobaileyaceae 925
Illiciaceae 925
Schisandraceae 926
Ceratophyllales 927
Ceratophyllaceae 927
Magnoliids 927
Canellales 927
Winteraceae 927
Laurales 929
Calycanthaceae 929
Lauraceae 929
Magnoliales 931
Annonaceae 931
Magnoliaceae 932
Myristicaceae 937
Piperales 938
Lactoridaceae 938
Saururaceae 938
Monocotyledons 940
Alismatales 940
Alismataceae 940
Araceae 940
Hydrocharitaceae 943
Zosteraceae (Seagrasses) 943
Asparagales 944
Agapanthaceae 944
Hemerocallidaceae 944
Orchidaceae 944
Dioscoreales 945
Dioscoreaceae 945
Liliales 945
Petermanniaceae 945
Pandanales 946
Pandanaceae 946
Triuridaceae 946
Commelinids 946
Arecales 946
Arecaceae (=Palmae) 946
Commelinales 948
Commelinaceae 948
Poales 948
Cyperaceae 948
Poaceae (Gramineae) 949
Zingiberales 951
Musaceae 951
Zingiberaceae 952
Eudicots 952
Buxaceae 953
Trochodendraceae 954
Proteales 956
Nelumbonaceae 956
Proteaceae 958
Platanaceae 960
Ranunculales 963
Berberidaceae 963
Ranunculaceae 963
Core Eudicots 964
Gunnerales 964
Gunneraceae 964
Caryophyllales 964
Phytolaccaceae 964
Saxifragales 965
Cercidiphyllaceae 965
Haloragaceae 966
Hamamelidaceae 968
Iteaceae 968
Saxifragaceae 969
Rosids 969
Vitaceae 970
Myrtales 971
Lythraceae 971
Trapaceae 971
Myrtaceae 971
Onagraceae 973
Eurosids I (Fabids) 973
Fabales 973
Fabaceae (=Leguminosae) 973
Fagales 976
Betulaceae 976
Casuarinaceae 978
Fagaceae 979
Juglandaceae 984
Myricaceae 989
Nothofagaceae 989
Malpighiales 990
Clusiaceae 990
Euphorbiaceae 991
Salicaceae 993
Malpighiaceae 993
Oxalidales 993
Cunoniaceae 993
Elaeocarpaceae 994
Rosales 994
Moraceae 994
Rhamnaceae 994
Rosaceae 994
Ulmaceae 996
Eurosids II (Malvids) 999
Brassicales 999
Capparaceae 999
Malvales 999
Tiliaceae 999
Sapindales 999
Anacardiaceae 1000
Meliaceae 1001
Rutaceae 1001
Sapindaceae 1002
Asterids 1004
Cornales 1004
Cornaceae 1004
Curtisiaceae 1007
Hydrangeaceae 1007
Ericales 1008
Ebenaceae 1008
Ericaceae 1008
Theaceae 1008
Euasterids I (Lamiids) 1009
Icacinaceae 1009
Garryales 1010
Eucommiaceae 1010
Gentianales 1010
Gentianaceae 1010
Rubiaceae 1010
Lamiales 1011
Avicenniaceae 1011
Byblidaceae 1011
Lentibulariaceae 1011
Oleaceae 1011
Solanales 1011
Solanaceae 1011
Euasterids II (Campanulids) 1011
Bruniaceae 1011
Quintiniaceae 1011
Apiales 1012
Araliaceae 1012
Aquifoliales 1012
Aquifoliaceae 1012
Asterales 1013
Asteraceae (=Compositae) 1013
Menyanthaceae 1014
Dipsacales 1014
Caprifoliaceae 1014
Cenozoic Floras 1014
Conclusions 1019
CHAPTER 23 Interactions Between Plants and Animals 1022
Early Terrestrial Ecosystem Associations 1024
Animals on Land 1024
Early Plant–Animal Associations 1024
Herbivory 1026
Defenses Against Herbivory 1027
Mechanical Protection 1028
Chemical Defenses 1029
Fossil Evidence of Herbivory 1030
Coprolites 1030
Gut Contents 1034
Marginal Feeding 1034
Defoliation 1036
Leaf Miners 1036
Wound Tissue 1038
Interactions with Vertebrates 1039
Herbivory 1039
Dentition 1041
Coprolites and Stomach Contents 1041
Dispersal 1041
Plants as Habitat 1042
Other Plant–Animal Interactions 1044
Mimicry 1044
Pollination 1045
Conclusions 1047
Appendix 1: Classification of Organisms 1050
Glossary 1054
A 1054
B 1055
C 1056
D 1058
E 1058
F 1059
G 1060
H 1060
I 1061
K 1061
L 1061
M 1062
N 1063
O 1063
P 1064
Q 1066
R 1066
S 1067
T 1069
U 1070
V 1070
W 1070
X 1070
Z 1070
References 1072
Index 1222
A 1222
B 1224
C 1226
D 1229
E 1230
F 1232
G 1233
H 1234
I 1235
J 1235
K 1235
L 1236
M 1238
N 1240
O 1241
P 1241
Q 1246
R 1246
S 1247
T 1250
U 1251
V 1251
W 1252
X 1252
Y 1253
Z 1253