Papers to Commemorate the Life and Legacy of Thomas N. Taylor Edited by Michael Krings SNSB-Bavarian State Collection for Palaeontology and Geology Munich, Germany Ludvvig-Maxin1il ians-U niversitat Munchen Munich, Germany The University of Kansas La\vrence, Kansas, United States J. Carla Harper The University of Kansas La\vrence, Kansas, United States SNSB-Bavarian State Collection for Palaeontology and Geology Munich, Germany Nestor Ruben Cuneo Museo Paleontol6gico Egidio Feruglio, and National Research Counci l of Argentina Trelew, Chubut, Argentina Gar W. 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Library of Congress Cataloging-in-Publication Data A catalog record for this book is available fro1n Lhe Library of Congress British Library Cataloguing-in-Publication Data ,IJ.,. catalogue record for this book is available from the British Library ISBN: 978-0-12-813012-4 For information on all Acaden,ic Press publicalions visit our \vebsite at https:// www .elsevier.con1/books-and-jou rnals ! l , . \Vorki11g togetl1er to grow 1i braries in !\id '"L.S;,_~ER Book developi11g cot1ntries lnternanonaJ www.elsevier.com • www.bookaid.org Publisher: Andre Gerhard \1/olff Acquisi1ions Edilor: t\ nna Valutkevich Editorial Project Manager: Pat Gonzalez Production Project 1\1anager: Sreejith Vis~ anathan 1 Cover Designer: Mark Rogers Typeset by TNQ Technologies Marep1,1an, 3a1141,1114eHH1>u,'i asropcK1'1M npasoM Contents Contributors 2. Early D evonian Woody Plants and XII I About the Editors XVII Implications for the Early Evolution . Fore\.vord XXI of Vascular Cambia Acknowledgments XXV Patricia G. Gensel 1. Introduction 21 2. Background 21 Section I 2 .1 Previously Described Occurrences Early Land Plants: Innovations and of Secondary Xylem in Fossil Plants 22 3. Previously Described Early Devonian Taxa 22 Adaptations 3 .1 Armoricaphyton chateaupannense 23 3.2 Unnamed New Brunswick Plant A: 1. The Evolutionary Origin of the Plant Prelirninary Data 24 Spore in Relation to the Antithetic 3.3 Franhueberia gerriennii, Ernsian, Origin of the Plant Sporophyte Gaspe 26 3.4 Additional Occurrences of E1nsian Paul K. Strother, Wilson A. Tavlor , Plants W ith Secondary Xylem: 1. Introduction 3 Preliminary Data 26 2. The Antithetic Theory as a Scaffold for 4. Comparisons 29 Interpreting the Fossil Record of the 5. Discussion 30 Algal-Plant Transition 4 5. 1 lmpl ications 30 3. Early Cryptospore Morphology 6 5.2 Evolution of Cambiun1 31 3 .1 Sporogenesis and Cryptospore 5.3 lnforn1ation From Developn1ental M orphology 6 Studies 31 3.2 Early Cryptospore Topology 8 5.4 Psilophyton- One Taxon 3.3 Spore \Nal I Topology in Relation to or Several? 32 the Sporopol lenin Transfer 5.5 Size is Decoupled From Hypothesis 9 Secondary Tissues in These 3.4 The Evolutionary Developn1ental Early Plants 32 and Morphological Significance of 6. Conclusions 32 Laminated Spore Wall Acknowledgments 32 Structure 1 I References 32 4. Cryptospores and the Origin of Meiosis in Plants 13 3. Using Architecture M odeling of the 5. The Stratigraphic Record of the Devonian Tree Pseudosporochnus to Cryptospores as a Record of Sporophyte Evolution 15 Compute Its Biomass 6. Interpreting Ordovician-Silurian Anae1le Dambreville, Brigitte Meyer-Berthaud, Phytodebris as a Record of Sporophyte Jean-Franc;ois Barczi, Anne-Laure Decon1beix, Evolution 17 Sebastien Griffon, Herve Rey 7. Conclusion 18 Acknowledgments 18 1. Introduction 35 References 18 2. Material 36 V Marepi.1an. 381UJ.1lll0HHbliii asropcKJ.1M npasoM vi Contents 3. Modeling Pseudosporochnus With 4. The Lirnitations of Charcoalified AmapSim 36 Lagerstatten 61 3 .1 AmapSirn Software 36 4.1 Taphonomy 61 3.2 Pseudosporoc hn us Architectural 4.2 Con1parisons Between D ifferent Unit 37 Preservation Types 62 3.3 Geometrical Parameters of 4.3 Diversity 63 Pseudosporochnus 38 5. Future Research 63 3.4 Pseudosporochnus Physiological 5 .1 New Localities 63 Age Indexing 40 5.2 Bulk Maceration and Sorting 63 4. Computing Biomass 40 5.3 Comprehensive Comparisons W ith 4.1 Biornass of a Pseudosporochnus the Dispersed Spore Record 64 Tree 40 5.4 Technological Advances in 4.2 Biomass of a Pseudosporochnus Anaton1 ica I Investigations 64 Stand 40 5.5 Geochemical Approaches 64 5. Results 40 Acknowledgments 65 5. I Simulating the Growth of References 65 Pseudosporochnus 40 5. 5.2 Carbon Al location in Early Tracheophyte Phylogeny: Pseudosporochnus 40 A Preliminary Assess,nent of 5.3 Carbon Content and Biomass of a Homologies Con1n1unity of Pseudosporochnus 41 Williarn L. Crepet, Karl J. Nik/as 6. Discussion 42 6.1 Con1puter Models of Fossil Plant 1. Introduction 69 Architecture 42 2. Materials and Methods 70 6.2 Model Accuracy 42 2. I Taxa Selection 70 6.3 Carbon Al location in 2.2 Characters and Character Pseudosporochnus 43 70 Assignments 6.4 Forest Biomass 44 2.3 Algorithn1s and Philosophy 76 7. Conclusions 45 3. Results 76 Supplementary Figure 46 4. Discussion 79 Acknowledgments 46 4.1 An Incomplete Fossi I Record 81 References 46 4.2 An Insufficient Nurnber of Characters and Character States 82 4. The Advantages and Frustrations of a 4.3 Rapid Radiations and Mosaic Plant Lagerstatte as Illustrated by a Evolution 82 New Taxon From the Lower 4.4 The Problern of Honiology 82 Devonian of the Welsh Borderland, UK 5. Conclusions 88 Appendices 88 Jennifer L. Morris, Dianne Edwards, References 91 John 8. Richardson Further Reading 92 1. Introduction 49 2. Geological Background and Previous Research 52 Section 11 2 .1 Geological Background 52 Late Paleozoic and Mesozoic Plants 2.2 Note,vorthy Advances by Previous and Floras Studies 53 3. New Research 54 6. Lower Permian Flora of the Sanzen- 3. I Material and Methods 54 bacher Ranch, Clay County, Texas 3.2 Morphology and Anatornical Descriptions 54 William A. DiMichele, Robert· W. Hook, 3.3 Comparisons and Systernatics 59 Hans Kerp, Carol L. Hotton, Cindy V. Looy, 3.4 Functional, Physiological, and Dan S. Chaney Evolutionary Conundrurns 60 1. Introduction 95 2. Geology 95 Marepi.1an. 381U"1lll0HHbliii asropcK"1M npasoM Contents vii 3. Collections 97 9. Pachytestopsis tayloriorum 4. Methods 98 gen. et sp. nov., an Anatomically 4.1 Macrofossils 98 Preserved G lossopterid Seed From 4.2 M icrofossils 101 the Lopingian of Queensland, s. Results 103 Australia 5 .1 Taxonomic Composition of the Flora 103 Stephen Mcloughlin, Benjamin Bornfleur, 5.2 Quantitative Composition of the Andrew N. Drinnan Flora 114 1. Introduction 155 6. Discussion 117 2. Geological Setting 155 6.1 Collecting 117 3. Material and Methods 156 6.2 Macroflora-Palynoflora (~o,nparison 120 4. Results 156 6.3 Broader Context of the 5. Discussion 165 Sanzenbacher Flora 121 5 .1 Affinities of the Seeds 165 7. Conclusions 123 5.2 Cornparison With Other Pern1ian Acknowledgments 124 Gondwanan Seeds 166 References 124 5.3 (~lossopterid Seed Character States 172 5.4 Associated Pollen 172 7. Permian Ginkgophytes of Angaraland 5.5 Taphonomy 175 6. Conclusions 175 Serge V. Naugolnykh Acknowledgments 176 1. Introduction 127 References 176 2. Historical Background 130 Appendix 1 178 3. Material and Methods 130 4. Results 131 10. A Triassic Mystery Solved: Fertile 4.1 Leaves of Psygmophylloids 131 Pekinopteris From the Triassic of 4.2 Sphenobaiera-Like Leaves 133 North Carolina, United States 4.3 Grovvth Forms 133 4.4 Fen,ale Reproductive ()rgans 137 Brian Ax.smith, Judith Skog, Christian Pott 4.5 Male Fructi fications/Pol I in i ferous 1. Introduction 179 Organs 137 2. Geological Setting 180 5. Discussion 139 3. Material and Methods 180 Acknowledgments 141 3 .1 Discovery and Nature of the References 141 Specin1ens 180 3.2 Preservation and Methods of 8. G lossopterid Plant Remains in Analysis 180 Permineralization: What Do They 4. Systematic Paleontology 180 Tell Us? 5. Discussion 181 5 .1 Structure and Life H istory 181 Harufurni Nishida, Kathleen B. Pigg, 5.2 Con,parisons and Potential M elanie L. Devore a·) 1 Phylogenetic lmpl ications . .;, 1. Introduction 145 6. Conclusion 185 2. Material and Methods 146 Acknowledgment<; 185 3. Results 146 References 186 4. Discussion 148 4.1 Comparison VVith Previously 11 . Enigmatic, Structurally Preserved Described, Anatomically Preserved Stems From the Triassic of Central Glossopterid Ovulate Structures Europe: A Fern or Not a Fern? and Ovules 148 Jean Ga/tier, Carla J. Harper, Ronny R6l3/er, 4.2 Morphological Comparison Evelyn Kustatscher, Michael Krings of Glossopterid and Angiospern, Reproductive O rgans 151 1. Introduction 187 Acknowledgments 152 2. Historical Background 188 References 153 3. Geological Settings 190 Marep1-1an. 3a!Ul-1lll0HHbliii asropcK1-1M npasoM viii Contents 4. Material and Methods 190 13. Aerodynamics of Fossil Pollen: s. Systematic Paleontology 196 Implications for Understanding 6. Results 197 Pollination Biology in Extinct Plants 6.1 Description 197 Lisa Grega, Adam Novotny, Christopher Stabile, 7. Discussion 202 Mackenzie L. Taylor, Charles P. Daghlian, 7 .1 Taxonorny 202 Jeffrey M. Osborn 7.2 Systematic ,~Jfinities 202 7.3 Growth Habit and Ecology 204 1. Introduction 253 7.4 Stelar Morphology of the Primary 2. Material and Methods 254 Vascular System 205 2 .1 Fossi I Materia I 254 8. Conclusions 207 2.2 Physical Scale Models 254 Acknowledgments 207 2 ·:1 Settling V1d ocity Experiments 255 References 207 2.4 Visualizing Wake Flow 257 2.5 Quantitatively Modeling Non- spherical Particle Drag at Low Reynolds Nu,nbers 257 Section 111 2.6 Calculating Drag c: oefficients and Paleobiogeography, Biology, and Shape Factors 258 Phylogenetic Relationships of Plants 3. Results 258 3 .1 Pollen Geo1netrv and Orientation 258 I 12. A Comprehensive Assessment of 3.2 Aerodynamic Coefficients 259 the Fossil Record of Liverworts in 3.3 Validation of Existing Mathe1nat- Amber ical 1\t\odels to Predict Settling Velocity 263 Jochen Heinrichs, Kathrin Feldberg, 3.4 Effect of Geometry Variations of Julia Bechteler, Ledis Regalado, Gothania on Settling Velocity 263 Matthew A.M. Renner, Alfons Schafer- 4. Discussion 264 Verwimp, Carsten Gro"hn, Patrick Muller, 4.1 Aerodynan1ic Implications of Harald Schneider, Michael Krings Pollen Geon1etry 264 1. Introduction 213 4.2 Aerodynamic lmpl ications 2. Material and Methods 217 of Sacci 266 2 .1 Amber Localities Yielding 4.3 Monoletes 268 s. Conclusions 268 Liven,vort Fossils and Their Ackno\vledgments 268 Geological Age 217 References 268 2.2 ln1agi ng of Fossi Is 218 2.3 Divergence Time Estin1ates 218 14. Escapia gen. nov.: Morphological 2.4 Lineage-Through-Ti me Plots and Coding of Ecological Preferences 219 Evolution, Paleogeographic 3. Results 219 Diversification, and the 3 .1 Systen1atic Paleontology 225 Environmental D istribution of 4. Discussion 225 Marattialean Ferns Through Time 4.1 Identification of Incompletely Gar W Rothwell, Michael A. Millay, Preserved Amber Fossils 225 Ruth A. Stockey 4.2 Classification of Extant Liverv,orts 234 4.3 Independent Evidence Derived 1. Introduction 271 From Molecular Chronograms 238 2. Material and Methods 272 4.4 Amber Fossil Record and the 3. Systematic Paleontology 272 ('.retaceous Terrestrial Revolution 239 3. I Systernatics 272 4.5 Hovv Complete Is the t\mber 3.2 Description 274 Fossi I Record of Liven,vorts? 240 3.3 Occurrences of M arattiales s. Conclusions and Future Prospects 241 Through Time 278 Acknowledgments 241 4. Discussion 281 References 241 4.1 Relationships of Escapia Appendix: rbcL Accession Numbers 249 chri.stensenioides 281 Marepi.1an. 3a!U"1lll0HHbliii asropcK"1M npasoM Contents ix 4.2 Characterization of the Families 16. Why Are Bryophytes So Rare in the Psaroniaceae and Fossil Record? A Spotlight on M arattiaceae 284 Taphonomy and Fossil Preservation 4.3 Aphlebia, Stipules, and Pulvini in Alexandru M .F. Tomescu, Benjamin Bo,nfleur, Eusporangiate Ferns and Fern-Like Alexander C. Bippus, Adolfina Savoretti Plants 285 4.4 Stratigraphic Range of the Fa,nily 1. Introduction 375 Psaroniaceae 285 2. Challenges to Traditional Views of the 4.5 Arrival of the Family M arattiaceae 286 Bryophyte Fossil Record 376 4.6 Floristic Turnover of Psaroniaceae 2 .1 Observations and Experimental to M arattiaceae 286 Data on Extant Bryophytes 376 4.7 Paleoecology of the Psaroniaceae 287 2.2 Stratigraphic Range of Bryophytes 377 4.8 Paleogeographid Paleoclimatic 2.3 M odes of Fossi I Preservation 381 Distribution of Marattiales 288 2.4 Overview of the Bryophyte 5. Conclusions 288 M acrofossil Record 384 References 289 2.5 The Fossilization Potential of Further Reading 293 Bryophytes Is On Par W ith That of Appendix 1: Synangium Characters of Tracheophytes 386 Escapia christensenioides Specimens 293 3. Bryophyte Taphonomy 386 Appendix 2: Paleontological 3 .1 Necrology 387 Occurrences of Marattialean Ferns 294 3.2 Biostratinomy 387 3.3 D iagenesis 388 15. Heterosporous Ferns From 3.4 Exquisite Bryophyte Fossi I Patagonia: The Case of Azol/a Preservation 389 3.5 A Taphonornic Surnmary of C. Facundo De Benedetti, Mana def Zamaloa, Bryophyte Fossilization 390 Marta A. Gandolfo, Nestor Ruben Ciineo 4. Hun1an Bias 391 1. Introduction 361 5. Conclusions and Future Outlook 392 2. Material and Methods 361 Ackno\.vledgments 393 2 .1 Stratigraphy 361 References 393 2.2 Palynological Techniques 362 Appendix ·1 399 2.3 Tern1inology and Classification 362 3. Results 363 17. Fossil Seeds With Affinities to 3 .1 Systematic Palaebotany 363 Austrobaileyales and Nymphaeales 3.2 Specific Diagnosis 363 From the Early Cretaceous (Early to 3.3 DE:scription '.163 Middle Albian) of Virginia and 4. Discussion 366 Maryland, USA: New Evidence for 4.1 Comparison With Modern Azolla Extensive Extinction Near the Base Species 367 of the Angiosperm Tree 4.2 Comparison W ith Other Fossil Azolla Species 367 Else Marie Friis, Peter R. Crane, 4.3 Taxonomy, Systernatics, and Kaj R. Pedersen Divergence Tirne Estin1ates 369 1. Introduction 417 4.4 Paleogeographical Remarks 369 2. Geological Setting 418 4.5 Paleoenvironmental Remarks 370 3. Material and Methods 418 5. Conclusions 371 4. Results 418 Acknowledgments 371 5. Discussion 430 References 371 .5. 1 Distinguishing Features for Nitaspermum 430 Marep1-1an. 381Ul-1lll0HHbliii asropcK1-1M npasoM x Contents 5.2 Comparison With Other Fossi I 15. Plant Cell Walls as Defense Against Seeds 432 Endoparasitism 452 5.3 Con,parison \,Vith Seeds of Extant 16. Endoparasitic Prokaryotes in Plants 452 Angiosperms and Systematic 17. Building on the Reactive Oxygen Assignment of the Fossil Seeds 433 Defensive System in Animals 453 6. Conclusion 434 18. Multicellularity as Defense Against Acknowledgrnents 434 Endoparasites 453 References 434 19. Evolution of Apoptosis 454 20. Conclusions 454 Acknowledgments 455 Section IV References 455 Further Reading 460 Fossil Microorganisms 19. Fossils of Arbuscular Mycorrhizal 18. Reactive Oxygen Defense Against Fungi Give Insights Into the History Cellular Endoparasites and the of a Successful Partnership W ith Origin of Eukaryotes Plants James F. White, Jr., Kathryn Kingsley, Mark C. Brundrett, Christopher Walker, Carla }. Harper, Satish K. Vern1a, Lara Brindisi, Carla}. Harper, Michael Krings Qiang Chen, Xiaoqian Chang, April Micci, Marshall Bergen 1. Introduction 461 2. The Early History of Mycorrhizal Fungi 463 1. Introduction 439 3. Fungal Associations of Fossil Plants and 1. 1 J'v\odels for Eukaryogenesis 439 Their Living Equivalents 465 1.2 Overview of Proposed Model 440 4. Comparing Mycorrhizas in Fossil and 2. Frequent Invasion of Eukaryotic Cells Modern Land Plants 474 by Diverse Microbes 440 Ackno,vledgments 475 2 .1 Bacterial Entry Into Eukaryotic References 476 Cells 440 2.2 Bacteria Are Progra,ned for 20. Looking for Arbuscular Mycorrhizal Endoparasitism 44 1 Fungi in the Fossil Record: 3. Were Archaea Victims of Endoparasi- An Illustrated Guide tism by Bacteria? 441 4. The Fossil Record of Endoparasitisrn 444 Christopher Walker, Carla J. Harper, 5. Reactive Oxygen Defense Response Mark C. Brundrett, Michael Krings of the Host Cell 446 6. ER and NADPH-Oxidase 449 1. Introduction 481 2. Materials Used to Illustrate the Guide 4f32 7. Membrane Sterols to Protect 3. Life Cycle of Extant AMF 4f32 Eukaryote Membranes From Reactive Oxygen Damage 449 4. Structural Characteristics of AMF 482 8. Evolution of the Cytoskeleton 4.1 Arbuscules and Hyphal Coils 484 Network of Eukaryotic Cells 450 4.2 Vesicles 485 9. Evolution of the Nuclear Envelope 450 4.3 Spore Morphology, Development, 1 o. Evolution of Autophagy 450 and Germination 486 11. Evolution of M itochondria 450 4.4 Auxiliary Cells 497 12. Massive Genon,es as a Legacy of 4.5 Spore Wall 497 Endo parasitism 451 4.6 /\r1ycel ia I Structure 506 s. 13. Chloroplast Acquisition 45 1 Fine Root Endophytes 508 6. Sumrnary and O utlook 514 14. Fungal Cell Walls as an Adaptation to Exclude Prokaryotic Endoparasites 451 Acknowledgments 515 References 515 Marepi.1an. 3a!U"1lll0HHbliii asropcK"1M npasoM Contents xi 21. Exceptional Preservation of Sessile, 4. Lichens in Time: The Fungal and Eukaryote Molecular Clock Revisited 563 Long-Stalked Microorganisms in the 5. Were the Ediacarans Lichens? 569 Lower Devonian W indyfield Chert 6. Protolichens: A Narro\v Prernise With (Scotland) 519 Broad Implications? 573 Michael Krings, Carla }. Harper, Hans Kerp, 7. Are Aspergillus and Penicillium Derived Edith L. Taylor From Lichenized Ancestors? 576 8. Lichens: Multiple Gains of a Successful 1. Introduction 519 Lifestyle 577 2. Geological Setting, Material, and 9. Conclusions 579 Methods 520 Acknowledgments 580 3. Results 520 References 580 4. Discussion 520 4.1 Affinities 520 4.2 Preservation in a Conservation Section V Trap 522 5. Conclusions 523 Antarctic Paleobotany Acknowledgments 524 References 524 24. Polar Regions of the Mesozoic- Paleogene Greenhouse World as 22. Morphological Convergence in Refugia for Relict Plant Groups Forest Microfungi Provides a Proxy Benjarnin Bomfleur, Patrick Blomenkemper, for Paleogene Forest Structure Hans Kerp, Stephen Mcloughlin Jouko Rikkinen, Alexander R. Schmidt 1. Introduction 593 1. Introduction 527 1. 1 The Dicroidium Plants 593 2. Morphology, Distribution, and Ecology 2. Material and M ethods 594 of Extant Calicioids 529 2 .1 D ispersed Cuticles Fron1 Section 2 .1 Morphology and Anaton1y 529 Peak 595 2.2 Distribution 532 2.2 Dispersed Cuticles From Shafer 2.3 Ecology 533 Peak 595 3. Fossil Calicioids Preserved in Amber 535 2.3 The Dicroicfium Assen1blage From 4. Discussion 540 Tirnber Peak 595 Acknowledgments 543 3. Results 595 References 543 4. Discussion 599 4.1 Relictual Dicroidiu,n Occurrences 23. Ediacarans, Protolichens, and in the Jurassic of Antarctica 599 Lichen-Derived Penicillium: 4.2 Distribution of Dicroidiurn A Critical Reassessment of the Through Space and Time 599 Evolution of Lichenization in Fungi 4.3 Pl ant Rel ictual ism in the High Latitudes of a Greenhouse \A/orld 600 Robert Liicking, Matthew P. Nelsen 4.4 Analogies With Low- and Mid-Latitude J\1ontane Refugia 605 1. Introduction 551 s. Conclusions 605 2. What Constitutes a (Fossil) Lichen? 552 Ackno,-vledgments 607 3. Lichens and Lichen Symbiosis in the References 607 Fossil Record 553 3 .1 Subfossils 553 25. Leaf Venation Density and 3.2 Brovvn c:oal (Lignite) Fossils .558 Calculated Physiological 3.3 Arnber Fossi Is .558 3.4 Three-Dimensional Perrnineralized Characteristics of Fossil Leaves and Charcoalified Fossils 559 From the Per,nian of Gondwana 613 3.5 Adpression (Compression, Andrew B. Schwende1nann ln1pression) Fossi Is, Casts, and M olds 560 1. Introduction 61 ·) ) J 3.6 Biochemical Signatures 561 2. Materials and M ethods 614 3.7 Evaluation of the Fossil Record of 2 .1 Stratigraphy 614 Lichens 561 Marep1-1an. 3a!Ul-1lll0HHbliii asropcK1-1M npasoM