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PALEOBOTANY
The Biology and
Evolution of Fossil Plants
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 Staatssammlungfur Paldontologie und
Geologie und GeoBio-CenterLMV,
Munich, Germany
Wm P » AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD
PARIS SAN DIEGO
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* S A N FRANCISCO • SINGAPORE • SYDNEY • TOKYO
ELSEVIER
Academic Press is an imprint of Elsevier
CONTENTS
Preface xv
Acknowledgments xvii
About the Authors xxi
CHAPTER 1
Introduction to Paleobotany,
How Fossil Plants are Formed
/
What Is Paleobotany?
The Objectives of Paleobotany
Reconstructing the Plants
Evolution of Plant Groups
Form and Function in Fossil Plants
Biostratigraphy and Correlation
Paleoecology: Plants in Their Environment
Determining Paleoclimate from Fossil Plants
Tree Rings
Nearest Living Relative
Leaf Physiognomy
Stomatal Index
Summary
Preservation: How Plant Fossils are Formed and
Preserved
Depositional Environments ofjFossil Plants
Compressions
!
Cuticle
Biofilms and Plant Fossil Preservation
Electron Microscopy
Confocal Microscopy
Maceration and Degagement
Other Techniques
Coal and Charcoal
Impressions
Molds and Casts
Cellular Preservation
Permineralization
Peel Technique
Coal Balls
Other Permineralizations
Petrifaction
Unaltered Plant Material
/
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29
30
30
Chemical Fossils
Ancient DNA
Mummification
Amber
Summary Discussion
Palynology
Geochronology and Biostratigraphy
Paleoecology
Absolute Dating
Geologic Timescale
Biological Correlation
Systematics and Classification
Nomenclature of Fossil Plants
Classification of Organisms
Background Reading
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34
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38
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41
42
42
CHAPTER 2
Precambrian Life 43
The Origin of Life on Earth
Origin of Life: Theory and Biology
Earliest Record of Life on Earth
Historical Background
Earliest Records of Life: Paleoarchean (3.6-3.2 Ga)
Geochemistry
Microfossils (Body Fossils)
Isua Greenstone Belt, Greenland
Warrawoona Group, Australia
Barberton Greenstone Belt, South Africa
Stromatolites
Sedimentary Evidence
Mesoarchean-Neoarchean Life
Conclusions: Archean Life
Oxygenation of the Earth (2.45-2.2 Ga)
Proterozoic Life
Paleoproterozoic
Origin of Eukaryotes
Mesoproterozoic
Earliest Multicellular Life
Neoproterozoic
Bitter Springs Biota
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VI
CONTENTS
Stromatolites
Other Microfossils
Doushantuo Formation
Conclusions
66
67
70
70
CHAPTER 3
Fungi, Bacteria, and Lichens 71
Fungi
Earliest Fossil Fungi
Systematics of Fungi
Chytridiomycota
Zygomycota
Glomeromycota
Ascomycota
Basidiomycota
Other Fungal Remains
Fungal Life-History Strategies
Saprotrophism
Parasitism
Mutualism
Fungi-Animal Interactions
Geologic Activities of Fungi
Epiphyllous Fungi
Fungal Spores
Fungal-like Organisms
Peronosporomycetes (Oomycota)
Eubacteria and Archaea
Archaea
Eubacteria
Cyanobacteria
!
Lichens
71
73
77
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82
84
90
93
97
98
98
99
103
105
107
108
111
112
112
112
113
113
115
117
CHAPTER 4
Algae
121
Chlorophyta (Green Algae)
,
Prasinophyceae
Chlorophyceae
Volvocales
Tetrasporales
Chlorococcales
Ulvophyceae
Dasycladales
Receptaculitida and Cyclocrinales
Caulerpales
Taxa Incertae Sedis
Charophyceae
Charales
Zygnematales
Euglenophyta
123
124
126
126
126
127
128
128
130
130
133
133
134
138
138
Dinophyta (Dinoflagellates)
Heterokontophyta
Bacillariophyceae (Diatoms)
Dictyochophyceae (Silicoflagellates)
Xanthophyceae (Yellow-Green Algae)
Phaeophyceae (Brown Algae)
Prymnesiophyta (Haptophytes)
Rhodophyta (Red Algae)
Solenoporaceans
Other Calcified Red Algae
Corallinales
Uncalcified Red Algae
Acritarcha (Acritarchs)
CHAPTER 5
Hornworts and Bryophytes 161
Early Fossil Evidence
Anthocerotophyta (Hornworts)
Bryophyta (Bryophytes)
Marchantiophytina (Liverworts or Hepatophytes)
Bryophytina (Mosses)
CHAPTER 6
The Move to the Land 179
Enigmatic Organisms
Nematophytes
Prototaxites
Nematothallus
Nematoplexus
Nematasketum diversiforme
Pachytheca
Spongiophytaceae
Spongiophyton
Orestovia
Other Enigmatic Organisms
Protosalvinia
Parka
Isolated Fragments: Clues to the Transition to Land?
Cuticle and Cuticle-Like Material
Spores and Spore Tetrads
Tubes
Land Plant Ancestors
The Transition to Land
Anchorage and Water Uptake
Structural Support and Water Transport
Protection Against Desiccation and Radiation
Gas Exchange
Reproduction on Land
Life History Biology
CONTENTS
Homologous Theory
Antithetic Theory
Animals
A Fungal Partner
Conclusion
196
196
198
198
199
CHAPTER 7
Introduction to Vascular Plant Morphology
and Anatomy 201
Plant Organography
Cell Types
Parenchyma
Collenchyma
Sclerenchyma
Tracheary Elements
Tracheids
Vessel Elements
Sieve Elements
Plant Tissues and Primary Growth
Xylem Tissue
Phloem Tissue
Meristems
Epidermis
Cuticle
Stomata
Trichomes
Anatomy of Stems and Roots
Arrangement of Primary Tissues
Primary Xylem Maturation Patterns
Secondary Development
k
Vascular Cambium
Cork Cambium (Phellogen)
Secondary Xylem
Secondary Phloem
Stele Types
,
Primitive Vascular Plants (Vascular Cryptogams)
Seed .Plants
Leaf Morphology and Anatomy
Leaf Anatomy
Leaf Evolution
Further Reading
202
203
203
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203
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204
206
206
207
207
207
208
208
209
209
210
210
210
212
212
212
213
214
216
216
216
219
221
221
222
222
CHAPTER 8
Early Land Plants with Conducting
Tissue 223
Conducting Elements in Early Land Plants
History of Discovery
224
225
Rhyniophytes
Rhynie Chert Plants
Aglaophyton major
Rhynia Gwynne-vaughanii
Horneophyton lignieri
Asteroxylon mackiei
Nothia aphylla
Trichopherophyton teuchansii
Ventarura lyonii
Gametophyte Generation
Other Rhyniophytes
Discussion: Rhyniophyte Evolution
Zosterophyllophytes
Zosterophyll Evolution
Trimerophytes
Trimerophyte Evolution
Early Land Plant Evolution
vu
227
228
229
235
237
238
239
241
241
241
246
251
252
259
259
262
263
CHAPTER 9
Lycophyta
265
Evolution of the Microphyll
Drepanophycales
Protolepidodendrales
Lepidodendrales
Vegetative Features
Stem Surface and Leaf Bases
Stem Anatomy
Cortical Tissues
Stem Development
Leaves
Underground Organs
Development of Underground Organs
Reproductive Biology
Microsporangiate and Bisporangiate Cones
Megasporangiate Cones
Gametophytes
Sigillariaceae
Leaf Bases
Leaves
Stem Structure
Underground Organs
Reproductive Biology
Other Lepidodendrid Genera
Lycopodiales
Selaginellales
Pleuromeiales
Isoetales
Putative Lycopsids
Conclusions
267
268
271
279
282
282
285
286
287
289
289
293
294
295
297
302
303
304
305
305
306
306
307
310
312
316
320
325
326
VIU
CONTENTS
CHAPTER 1 0
Sphenophytes 329
Pseudoborniales
Sphenophyllales
Devonian Sphenophyllales
Sphenophyllum
Leaves
Stem Anatomy
Roots
Reproductive Biology
Other Sphenophyllales
Ecology
Equisetales
Calamitaceae
Archaeocalamites
Catamites
Pith Casts
Stem Anatomy
Extraxylary Tissues
Growth and Development
Roots
Leaves
Other Calamitean Leaves
Reproductive Biology
Spores
Tchernoviaceae and Gondwanostachyaceae
Vegetative Body
Reproductive Biology
Equisetaceae
Forms with Uncertain Affinities
Sphenophyte Evolution
331
332
333
334
334
335
337
337
338
341
342
343
343
345
349
350
352
352
353
354
357
358
366
368
368
369
371
376
379
CHAPTER 11
Ferns and Early Fernlike Plants 383
<
Evolution of the Megaphyll
Cladoxylopsida
Pseudosporochnales
Calamophyton Plant
Iridopteridales
Phylogenetic Position of the Cladoxylopsids
Early Fernlike Plants
Rhacophytales
Rhacophyton
Other Taxa
Systematics of the Rhacophytales
Coenopterid Ferns
Stauropteridales
Zygopteridales
386
387
388
396
398
400
401
401
402
403
404
405
405
408
Zygoptend Evolution
Marattiales
Psaroniaceae: Vegetative Features
Psaronius Plant
Other Stem Taxa
Psaroniaceae: Reproductive Features
Paleozoic Compression Taxa
Mesozoic Marattialeans
Marattialean Evolution
Ophioglossales
Leptosporangiate Ferns
Osmundales
Paleozoic Stem Taxa
Guaireaceae
Mesozoic and Cenozoic Stem Taxa
Sterile and Fertile Foliage
Osmundalean Evolution
Botryopteridaceae
Vegetative Organs
Reproductive Organs
Other Genera
Anachoropteridaceae
Kaplanopteridaceae
Psalixochlaenaceae
Sermayaceae
Tedeleaceae
Skaaripteridaceae
Tempskyaceae
Schizaeaceae
Hymenophyllaceae
Gleicheniaceae
Dicksoniaceae
Cyatheaceae
Matoniaceae
Loxsomataceae
Dipteridaceae
Polypodiales
Salviniales
Marsileaceae
Salviniaceae
Conclusions
CHAPTER 1 2
Progymnosperms 479
Archaeopteridales
Archaeopteris Leaves
Archaeopterid Reproduction
Callixylon Stems
Other Archaeopterids
CONTENTS
Aneurophytales
Aneurophyton
Tetraxylopteris
Triloboxylon
Rellimia
Other Aneurophytes
Protopityales
Noeggerathians
Progymnosperm Evolution
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489
489
491
492
494
496
497
501
CHAPTER 1 3
Origin and Evolution of the Seed Habit
Homospory, Heterospory, and the Seed Habit
Homospory
.r
J
Heterospory
Sporangia
Endospory
Lycopsid Heterospory
Seed Habit
Evolution of the Integument
Evolution of Pollen Capture
Pollen
Cupules
Cupulate Devonian Seeds
Reproductive Biology
Carboniferous Seeds
Pollen Chamber Function
Microgametophytes
Diversity of Early Seeds
Paleozoic Seeds with Embryos
CHAPTER 1 4
Paleozoic Seed Ferns
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Calamopityales
Buteoxylonales
Lyginopteridales
.
Lyginopteris Plant
Vegetative Organs
Reproductive Structures
Other Lyginopterids: Vegetative Remains
Heterangium
Microspermopteris
Schopfiastrum
Pitys
Devonian-Mississippian Taxa
Problematic Lyginopterids
Other Lyginopterids: Seeds and Cupules
Sphaerostoma
Salpingostoma
531
539
540
540
540
542
546
547
550
550
551
552
554
555
556
556
Conostoma
Coronostoma
Physostoma
Tyliosperma
Calathospermum
Gnetopsis
Megatheca
Other Lyginopterids: Pollen Organs
Incertae Sedis
Lyginopterid Evolution
Medullosales
Stems
Medullosa
Other Stem Taxa
Leaves (Fronds)
Roots
Growth Habit
Seeds
Pollen organs
Pollen
Medullosan Evolution
Callistophytales
Vegetative Organs
Reproductive Structures
Callistophytalean Evolution
Glossopteridales
Leaves
Glossopteris
Gangamopteris
Other Leaf Types
Stems and Roots
Ovulate Reproductive Structures
Permineralized Forms
Impression-Compression Specimens
What is the Glossopterid Ovulate Structure?
Pollen Organs
Glossopteris Habit and Habitat
Phylogenetic Position
IX
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557
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573
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603
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606
606
609
614
616
618
618
CHAPTER 1 5
Mesozoic Seed Ferns
Caytoniales
Sagenopteris
Caytonanthus
Caytonia
Ruflorinia and Ktalenia
Corystospermales
Foliage
Stems
621
622
622
623
624
626
627
627
630
CONTENTS
Pollen Organs
Ovulate Structures
Petriellales
Peltaspermales
Foliage
Reproductive Organs and Whole-Plant
Concepts
Conclusions
631
634
637
639
639
643
648
1 ZT
CHAPTER I D
Late Paleozoic and Mesozoic Foliage
651
Late Paleozoic Foliage
Adiantites
Alethopteris
Aneimites
Aphlebia
Alloiopteris
Botrychiopsis
Callipteridium
Cardiopteridium
Cardiopteris (Fryopsis)
Charliea
Cyclopteris
Dicksoniites
Discopteris
Eremopteris
Ginkgophytopsis
Kankakeea
Karinopteris, Mariopteris, and Pseudomariopteris
Lesleya
i
Linopteris, Reticulopteris, and Barthelopteris
Lobatopteris
Lonchopteridium and Lonchopteris
Megalopteris
Neuropteris sensu lato
,
Laveinopteris
I Macroneuropteris
Margaritopteris
Neuralethopteris
Neurocallipteris
Neurodontopteris
Neuropteris sensu stricto
Paripteris
Sphenoneuropteris
Neuropterid Growth Habit
Blanzyopteris
Nothorhacopteris
Odontopteris and Lescuropteris
Pecopteris
652
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656
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662
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679
Rhodea (=Rhodeopteridium)
Sphenopteris
Sphenopteris sensu stricto
Eusphenopteris
Spiropteris
Taeniopteris
Tinsleya
Triphyllopteris, Genselia, and Charbeckia
Mesozoic Foliage
Anomozamites
Cladophlebis
Coniopteris
Ctenis
Deltolepis and Cycadolepis
Dictyophyllum
Dictyozamites
Doratophyllum
Macrotaeniopteris
Matonidium
Mesodescolea
Nilssonia
Nilssoniopteris
Otozamites
Pachypteris, Komlopteris, and Thinnfeldia
Phlebopteris
Pseudoctenis
Pseudocycas
Pterophyllum
Ptilophyllum
Ptilozamites
Ruflorinia
Taeniozamites
Ticoa
Wingatea
Yabeiella
Zamites
CHAPTER 1 7
Cycadophytes
703
Cycadales
Leaves and Petioles
Stems
Paleozoic Reproductive Structures
Triassic Cycads
Jurassic Cycads
Pollination Biology
Discussion: Cycad Evolution
Bennettitales
Cycadeoidaceae
CONTENTS
Stem Anatomy
Reproductive Structures
Development
Williamsoniaceae
Ovulate Structures
Pollen Organs
Discussion: Bennettitales
725
728
730
732
734
738
739
794
795
795
798
801
803
CHAPTER 2 1
CHAPTER 18
Ginkgophytes
Conifers
743
Paleozoic Record
Ginkgophyte Wood
Ginkgophyte Foliage
Pollen-Producing Structures
Ginkgophyte Plants
Ginkgoaceae
Karkeniaceae
Umaltolepidiaceae
Yimaiaceae
Schmeissneriaceae
Taxa with Uncertain Affinities
Conclusions
744
747
747
750
750
750
752
752
752
753
754
755
CHAPTER 1 9
Gymnosperms with Obscure Affinities
Gigantopteri dales
Vegetative Remains
Reproductive Organs
Vojnovskyales
Czekanowskiales
Iraniales
[
Pentoxylales
'
Hermanophytales
Gnetales
Extant Genera
Ephedra
Gnetum
Welwitschia
Extant Reproductive Structures
Fossil Gnetophyte Pollen
Gnetophyte Megafossils
Putative Gnetophyte s
Dirhopalostachyaceae
,
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768
768
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781
785
CHAPTER 2 0
Cordaitales
Roots
Reproductive Features
Reproductive Organs
Seeds
Angaran Cordaites
Phylogenetic Position and Orijnnof the Cordaites
XI
787
Vegetative Features
Stems
Foliage
788
788
791
805
Early Conifers
Voltziales
Utrechtiaceae
Utrechtia
Ernestiodendron
Ortiseia
Otovicia
Moyliostrobus
Other Taxa
Thucydiaceae
Emporiaceae
Majonicaceae
Ullmanniaceae
Bartheliaceae
Other Voltzialeans
Ferugliocladaceae
Buriadiaceae
Pollen Cones
Summary Discussion: Voltzialeans
Coniferales
Palissyaceae
Cheirolepidiaceae
Summary Discussion: Cheirolepidiaceae
Podocarpaceae
Summary Discussion: Podocarpaceae
Araucariaceae
Summary Discussion: Araucariaceae
Cupressaceae
Cunninghamioideae
Taiwanioideae
Athrotaxoideae
Sequoioideae
Taxodioideae
Cupressoideae
Cupressaceous Wood
Summary Discussion: Cupressaceae
Sciadopityaceae
Pararaucariaceae
Pinaceae
Pinoideae
806
807
808
809
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816
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820
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863
xii
CONTENTS
Genus Pinus
Pinus Wood
Larix
Piceoideae
Abietoideae
Summary Discussion: Pinaceae
Cephalotaxaceae
Taxaceae
Summary Discussion: Cephalotaxaceae and
Taxaceae
Conclusions
864
866
866
867
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868
868
869
869
870
CHAPTER 11
Flowering Plants
873
Angiosperm Origins
Origin of the Flower
Pseudanthial Theory
Euanthial Theory
Microsporangial Theories
Transitional-Combination Theory
Habit
Ecological Considerations
Site of Origin
Pre-Cretaceous Fossil Evidence
Sanmiguelia
Furcula
Problematospermum
Pre-Cretaceous Pollen
Dispersed Pollen
Early Angiosperm Evidence
Pollen
Pollen Evolution
Evidence from Leaves
Angiosperm Ancestors
Caytoniales
Czekanowskiales
Glossopteri dales
Bennettitales
Pentoxylales
Gigantopteridales
Phylogenetic Analyses and Angiosperm Origins
Selected Angiosperm Families
Basal Angiosperms
Amborellaceae
Hydatellaceae
Archaefructaceae
Chloranthaceae
Nymphaeales
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877
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885
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889
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894
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901
Nymphaeaceae
Austrobaileyales
Austrobaileyaceae
Illiciaceae
Schisandraceae
Ceratophyllales
Ceratophyllaceae
Magnoliids
Canellales
Winteraceae
Laurales
Calycanthaceae
Lauraceae
Magnoliales
Annonaceae
Magnoliaceae
Myristicaceae
Piperales
Lactoridaceae
Saururaceae
Monocotyledons
Alismatales
Alismataceae
Araceae
Hydrocharitaceae
Zosteraceae (Seagrasses)
Asparagales
Agapanthaceae
Hemerocallidaceae
Orchidaceae
Dioscoreales
Dioscoreaceae
Liliales
Petermanniaceae
Pandanales
Pandanaceae
Triuridaceae
Commelinids
Arecales
Arecaceae (=Palmae)
Commelinales
Commelinaceae
Poales
Cyperaceae
Poaceae (Gramineae)
Zingiberales
Musaceae
Zingiberaceae
Eudicots
CONTENTS
Buxaceae
Trochodendraceae
Proteales
Nelumbonaceae
Proteaceae
Platanaceae
Ranunculales
Berberidaceae
Ranunculaceae
Core Eudicots
Gunnerales
Gunneraceae
Caryophyllales
Phytolaccaceae
Saxifragales
Cercidiphyllaceae
Haloragaceae
Hamamelidaceae
Iteaceae
Saxifragaceae
Rosids
Vitaceae
Myrtales
Lythraceae
Trapaceae
Myrtaceae
Onagraceae
Eurosids I (Fabids)
Fabales
Fabaceae (=Leguminosae)
Fagales
Betulaceae
\
Casuarinaceae
)
Fagaceae
Juglandaceae
Myricaceae
Nothofagaceae
Malpighiales
Clusiaceae
Euphorbiaceae
Salicaceae
Malpighiaceae
Oxalidales
Cunoniaceae
Elaeocarpaceae
Rosales
Moraceae
Rhamnaceae
Rosaceae
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971
Ulmaceae
Eurosids II (Malvids)
Brassicales
Capparaceae
Malvales
Tiliaceae
Sapindales
Anacardiaceae
Meliaceae
Rutaceae
Sapindaceae
Asterids
Cornales
Cornaceae
Curtisiaceae
Hydrangeaceae
Ericales
Ebenaceae
Ericaceae
Theaceae
Euasterids I (Lamiids)
Icacinaceae
Garryales
Eucommiaceae
Gentianales
Gentianaceae
Rubiaceae
Lamiales
Avicenniaceae
Byblidaceae
Lentibulariaceae
Oleaceae
Solanales
Solanaceae
Euasterids II (Campanulids)
Bruniaceae
Quintiniaceae
Apiales
Araliaceae
Aquifoliales
Aquifoliaceae
Asterales
Asteraceae (=Compositae)
Menyanthaceae
Dipsacales
Caprifoliaceae
Cenozoic Floras
Conclusions
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991
991
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CONTENTS
CHAPTER 2 3
Interactions Between Plants and Animals 999
Early Terrestnal Ecosystem Associations
Animals on Land
Early Plant-Animal Associations
Herbivory
Defenses Against Herbivory
Mechanical Protection
Chemical Defenses
Fossil Evidence of Herbivory
Coprolites
Gut Contents
Marginal Feeding
Defoliation
Leaf Miners
Wound Tissue
1001
1001
1001
1003
1004
1005
1006
1007
1007
1011
1011
1013
1013
1015
Interactions with Vertebrates
Herbivory
Dentition
Coprolites and Stomach Contents
Dispersal
Plants as Habitat
Other Plant-Animal Interactions
Mimicry
Pollination
Conclusions
1016
1016
1018
1018
1018
1019
1021
1021
1022
1024
Appendix 1: Classification of Organisms
Glossary
References
Index
1027
1031
1049
1199