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Ml 48106-1346 USA 313/761-4700 800/521-0600 I Order Number 8013706 Comparative analysis of sporodcrm ultrastructure in fossil and extant lycopods Taylor, Wilson Anthony, Ph.D. The Ohio State University, 1989 UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 COMPARATIVE ANALYSIS OF SPORODERM ULTRASTRUCTURE IN FOSSIL AND EXTANT LYCOPODS DISSERTATION Presented in partial fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Wilson Anthony Taylor, B.S., M.S. The Ohio State University 1989 Dissertation committee: Approved by T.N. Taylor V. Raghavan D.J. Crawford Adviser I F.D. Sack Departmentleeartment of BotanyRotanv* To Dad from Boo ii ACKNOWLEDGEMENTS I would like to thank the members of my reading committee, Drs. Daniel Crawford, Valayamghat Raghavan, and Fred Sack, for their time and effort. I am also grateful to the staff at the Royal Botanic Gardens, Kew, for their support during specimen collection and transport. I am especially indebted to Dr. Thomas N. Taylor for his financial support and scholarly guidance throughout all phases of this investigation. And finally, I recognize the loving suport of my wife, Robin, without whom none of this would have been possible. iii VITA June 3, 1960 ........ Born - Indiana, Pennsylvania 1982 ................ B.S., Geoscience Department, Indiana University of Pennsylvania, Indiana, Pennsylvania 1984 . ............ M.S., Department of Botany, The Ohio State University, Columbus, Ohio Field of Study: Botany PUBLICATIONS 1986a Ultrastructure of Sphenophyllalean spores. Review of Palaeobotany and Palynology 47y 105-128. 1986b Spore wall ultrastructure of Protosalyinia. American Journal of Botany 74? 437-443 (with T.N Taylor). 1987 Subunit construction of the spore wall in fossil and living lycopods. Pollen et Spores 29? 241-248. (with T.N. Taylor) 1988 Ultrastructural analysis of selected Cretaceous megaspores from Argentina. Journal of Micropalaeontology 7? 73-87. (with T.N. Taylor) 1989a Comparative ultrastructural analysis of fossil and living gymnoBperm cuticles. Review of Paoaeobotany and Palynology (with T.N. Taylor) 1989b Comparative ultrastructural analysis of microspores from fossil and living Selaginella. (with T.N. Taylor) 1989c Ultrastructural and developmental analysis of Carboniferous lycopod megaspore walls. Review of Palaeobotany and Palynology. 1984a Ultrastructure of sphenopsid spores. Ohio Journal of Science 84; 7. 1984b Spore wall ultrastructure in the Sphenophyllales. 2nd International Organization of Paleobotany Conference, Edmonton, Alberta. Abstracts of Contributed Papers, Poster Sessions. 1984c An ultrastructural investigation of sphenophyllalean spores. American Association of Stratigraphic Palynologists, Arlington, Virginia. AASP, Inc. Program and Abstracts; 21. 1985a Spore ultrastructure of the enigmatic alga Protosalyinia. Ohio Journal of Science 85; 18. (with T.N. Taylor) 1985b Spore and thallus ultrastructure of the enigmatic alga Protosalvinia. American Journal of Botany 72; 902. 1985c An ultrastructural comparison of Cretaceous and extant ginkgoalean cuticles. Botanical Society of America, Gainesville, Florida. 1986a Spore wall architecture of a megaspore from the Cretaceous of Argentina. Mid-continent Paleobotanical Colloquium IV, Ann Arbor, Michigan. 1986b Ultrastructural stasis within the Selaginellales. American Journal of Botany 73; 712. 1987 Evolutionary and developmental significance of megaspore wall ultrastructure. American Journal of Botany 74; 692. (Cookson Award) 1988 Developmental and structural aspects of lycopod megaspore wall ultrastructure. American Journal of Botany 75(6) Part 2; 50-51. v TABLE OF CONTENTS ACKNOWLEDGEMENTS..................................... ili VITA ............................................... iv LIST OF T A B L E S .................................. ix LIST OF P L A T E S ..................................... X CHAPTER I INTRODUCTION .............. 1 Literature survey ................................. 4 CHAPTER II MATERIALS AND METHODS .......... 16 M a t e r i a l s ......................................... 16 M e t h o d s ........................................... 17 Terminology ....................................... 22 CHAPTER III WALL DEVELOPMENT IN FOSSIL SYSTEMS . 24 Introduction.......... 24 R e s u l t s ........................................... 28 Mazocarpon - functional megaspores ........ 29 Mazocarpon aborted megaspores .............. 30 Lepidocarpon functional megaspores .......... 36 Lepidocarpon aborted megaspores ............ 36 Lagenoisporites ............................ 40 Valvisisporites ............................ 44 Discussion......................................... 46 Mazocarpon............................ 46 Valvisisporites ............................ 48 Lepidocarpon.......... 49 Lagenoisporites ............................ 51 Morphogenetic changes through wall development 52 Correlation of ultrastructure with dispersive s t r a t e g y .............................. 53 CHAPTER IV WALL ULTRASTRUCTURE IN SELAGINELLA . 56 Introduction ....................................... 56 Terms and concepts................................. 59 vi Species descriptions ............................... 65 Selaginella flabellata type ............ 65 Selaginella flabellata ....... 65 Selaginella u s t a .................. 66 Selaginella viridangula .......... 66 Selaginella inaequalifolia type ........ 72 Selaginella inaequalifolia ........ 72 Selaginella pallescens ............ 73 Selaginella pulcherrima .......... 74 Selaginella argentea type ............ 82 Selaginella argentea .............. 82 Selaginella ornata type ................ 83 Selaginella ornata ................ 83 Selaginella erythropus type ............ 87 Selaginella erythropus ............ 87 Selaginella pilifera .............. 88 Selaginella brevipes type .............. 92 Selaginella brevipes .............. 92 Selaginella elmeri ................ 93 Selaginella plana ................ 93 Selaginella fissidentoides type ........ 97 Selaginella fissidentoides ........ 97 Selaginella yemensis ................ 100 Selaginella intermedia type .............. 101 Selaginella intermedia .............. 101 Selaginella frondosa type ................ 104 Selaginella frondosa ................ 104 Selaginella lyallii type ................ 108 Selaginella lyallii ......... 108 Selaginella willdenovii ........... 109 D i s c u s s i o n ......................................... 115 Observations on basic megaspore types .... 117 Internal wall l a y e r s........................... 119 Surface layers ................................ 120 CHAPTER V WALL DEVELOPMENT IN SELAGINELLA .... 123 Introduction ...................................... 123 Description......................................... 128 Selaginella sulcata .......................... 128 Selaginella galeottii .... 136 D i s c u s s i o n .................................. 157 S. sulcata - wall d e v e l o p m e n t ................. 157 Structural basis for expansion ................ 158 Structural basis for unit enlargement .... 159 S. galeottii - wall d e v e l o p m e n t ...............160 Published descriptions of mature megaspores of S, galeottii ........ ........... 164 The role of the t a p e t u m ....................... 166 The m e s o s p o r e ................................. 168 Control of sporoderm patterning - sporophytic or gametophytic ............................ 170 vii Concordance with proposed models of sporoderm morphogenesis ........................... 170 Correlations with the ordered wall construction t y p e ..................................... 173 Implications on the nature of sporopollenin . 174 CONCLUSIONS ......................................... 176 APPENDIX A - GLOSSARY OF T E R M S ....................... 182 APPENDIX B - LIST OF SPECIMENS......................