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University Microfilms International 300 North 2««b Road Ann Arbor, Michigan 48106 USA S t John's Road, Tyler's Green High Wycombe, Bucks, England HP10 BHR I I 77-31,992 STOCKEY, Ruth Anne, 1950- MORPHOLOGY AND REPRODUCTIVE BIOLOGY OF CERRO CUADRADO FOSSIL CONIFERS. The Ohio State University, Ph.D., 1977 Botany University Microfilms International, Ann Arbor. Michi0an woe MORPHOLOGY AND REPRODUCTIVE BIOLOGY OF CERRO CUADRADO FOSSIL CONIFERS « DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University r By Ruth Anne Stockey, B.Sc., M.Sc. ****** The Ohio State University 1977 Reading Committee: Dr. Thomas N. Taylor Dr. Gary L. Floyd Dr. Barbara A. Schaal Dr. Valaymghat Raghavan Adviser Department of Bota: This publication is dedicated to the memory of the late Dr. Hans Tralau who provided much assistance and many of the specimens during this study, and whose untimely passing constitutes a great loss to the study of paleo­ botany. ii ACKNOWLEDGEMENTS The author wishes to thank the following people for the acquisition and loan of specimens and the use of lab­ oratory facilities: Dr. Eugene S. Richardson, Field Museum of Natural History, Chicago; Mr. Cedric Shute, British Museum of Natural History, London: Dr. Britta Lundblad, Naturhistoriska Riksmuseet, Stockholm; Dr. James M. Schopf and Dr. Stig Bergstrom, United States Geological Survey Coal Geology Laboratory and Orton Mu­ seum, The Ohio State University; Dr. John W. Hall, Uni­ versity of Minnesota; Dr. Francis M. Hueber, United States National Museum, Smithsonian Institution; Dr. Elizabeth McClintock, California Academy of Sciences, San Francisco; Mr. Frederick C. Boutin, Huntington Botanical Gardens, San Marino, CA; Dr. Enrique C. Clos, Jardin Agrobotanico de Santa Catalina, Buenos Aires; Dr. Garth Nikles, Forestry Department, Brisbane; Dr. Richard Gould and Mr. Ron Larson, Imbil Forestry Station, Queensland; M. Fran­ cois Guinaudeau, C. T. F. T. Parc Forestier, Noumea, New Caledonia. Special appreciation is expressed to Dr. Thomas N. Taylor for his guidance and support during this study. Supported in part by National Science Foundation grants BMS 75-14221 and BMS 74-21105. iii VITA January 3, 1950 Born - Harvey, Illinois 1972-1974. Teaching Assistant, Ohio University, Athens, Ohio 1974 ................... M.Sc., Ohio University, Athens, Ohio 1974-1977 Research and Teaching Assis­ tant The Ohio State Uni­ versity Columbus, Ohio PUBLICATIONS Stockey, R. A. 1974. Seeds and embryos Of Araucaria mirabilis. Amer. J. Bot. 61s 19-20 (Abstract). ______ . 1975a. Pararaucaria patagonica Wieland from the Cerro Cuadrado petrified forest (Jurassic) Argentina: Seeds and embryos. Akad. Nauk SSSR (XII International Botanical Congress Abstracts) p. 237. ______ . 1975b. Seeds and embryos of Araucaria mirabilis Amer. J. Bot. 62:856-868. ______ . 1975c. Morphology and anatomy of Pararaucaria patagonicas Seeds and embryos.' Bbt. Soc. Amer. Abstracts. Allen Press, Lawrence, Kansas p. 25. 1975d. Immature ovulate cones of Araucaria mirabilis. Bot. Soc. Amer, Abstracts'! Allen Press, Lawrence, Kansas, p. 25. ______ , and T.N. Taylor. 1976. Supposed silicified conifer seedlings from the Cerro Cuadrado petrified forest (Jurassic) Patagonia. Bo.t. Soc. Amer. Abstracts. Allen Press, Lawrence, Kansas, p. 32. Taylor, T. N. and R. A. Stockey. 1975. The morphological nature of Microspermopteris. Bot. Soc. Amer. iv Abstracts. Allen Press, Lawrence, Kansas, p. 25- 26. ______ , and . 1976. Studies of Paleozoic seed ferns: Anatomy and morphology of Microspermopteris aphyllum. Amer. J. Bot. 63:1302-l3l0, Stockey, R. A. 1977. Reproductive biology’of the Cerro Cuadrado (Jurassic) fossil conifers: Pararaucaria patagonica Wieland. Amer. J. Bot. 65:734-746, FIELDS OF STUDY Major Field: Paleobotany Phylogeny and reproductive biology of the Araucaria- ceae and fossil, conifer groups. Professor Thomas N. Taylor. Carboniferous coal ball permineralized plants: anatomy, morphology, ontogeny, and phylogenetic trends. Professor Thomas N. Taylor. v LIST OF FIGURES Page Figures 1-7. Pararaucaria patagonica................. 103 Figures 8-13. Seed and cone-scale features. 105 Figures 14-20. Pararaucaria patagonica embryos. 107 Figures 21-27. Seedlings . ...................... 109 Figures 28-33. Seedlings .......................... Ill Figures 34-37. Seedlings .......................... 113 Figures 38-44. Seedlings ........... 115 Figures 45-53. Seedlings .... ........... .... 117 Figures 54-57. Seedlings .......................... 119 Figures 58-68. Araucaria mirabilis. ............... 121 Figures 69-76. Araucaria mirabilis.................... 123 Figures 77-82. Araucaria mirabilis.................... 125 Figures 83-90. Araucaria mirabilis............. 127 Figures 91-96. Araucaria mirabilis.................... 129 Figures 97-101. Araucaria mirabilis............ 131 TABLE OP CONTENTS Page ACKNOWLEDGEMENTS..................................... iii VITA...................................... ............ iv LIST OF F I G U R E S ......................................... vi INTRODUCTION. ........... 1 MATERIALS AND METHODS .......... i . 5 Directory of Specimens............... 5 Techniques................ ; . ................... 9 STRATIGRAPHY ................... 14 PARARAUCARIA PATAGONICA WIELAND ....................... 16 Introduction....................................... 16 Emended diagnosis . ............ 17 Description..................... 19 General features ..... ............... 19 Cone axis.....................................20 Cone-scale complex ............ 21 Seeds .......... ............... 23 Embryos............................. 27 Discussion......................................... 29 SEEDLINGS . ............ 41 Introduction....................................... 41 Fossil seedlings...................................42 Living araucarian seedlings ................... 44 Other fossil seedlings............................ 49 Discussion......................................... 52 ARAUCARIA MIRABILIS (SPEGAZZINI) WINDHAUSEN EMEND. C A L D E R ................. 59 Introduction. ....... ................... 59 Emended diagnosis . '........... 63 Description . 64 vii Page Cone structure............................ 64 Cone a x i s ..........................‘ 66 Resin canals. .......................... 67 Isolated cone axes.............. 69 Cone-scale complex. ........... 70 Ovules and seeds.......................... 71 Integuments. ‘. ............... 72 Ovule vascularization.............. 73 N u c e l l u s ............................ 74 Abortive ovules. ............... 76 Megaspore membrane ................. 77 Megagametophy te..................... 77 Embryos . k . ................... 78 Discussion ....................... 81 GENERAL DISCUSSION .... .......................... 96 APPENDIX ................... 103 LIST OF REFERENCES..................................... 133 viii INTRODUCTION The Cerro Cuadrado Petrified Forest of Patagonia has yielded some of the best fossil conifer remains known. Silicified cones, wood, twigs with leaf scars, and sup­ posed "seedlings” from the province of Santa Cruz, Argen­ tina have been scattered to various universities and mu­ seums around the world. The original material comes from a forty square mile area in southern Patagonia near the bases of three Volcanic peaks, the Cerro Cuadrado, Cerro Alto, and Cerro Madre e Hija. Since the 1930's specimens have been sold as curios and kept in private collections because of their beauty and excellent preservation (Wieland, 1935). The petrified forest was initially