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Microfilms International 300 N INFORMATION TO USERS This reproduction was made from a copy of a document sent to us for microfilming. While the most advanced technology has been used to photograph and reproduce this document, the quality of the reproduction is heavily dependent upon, the quality of the material submitted. The following explanation of techniques is provided to help clarify markings or notations which may appear on this reproduction. 1.The sign or “target’* for pages apparently lacking from the document photographed is “Missing Page(s)”. If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure complete continuity. 2. When an image on the film is obliterated with a round black mark, it is an indication of either blurred copy because of movement during exposure, duplicate copy, or copyrighted materials that should not have been filmed. For blurred pages, a good image of the page can be found in the adjacent frame. If copyrighted materials were deleted, a target note will appear listing the pages in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photographed, a definite method of “sectioning” the material has been followed. It is customary to begin filming at the upper left hand comer of a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again—beginning below the first row and continuing on until complete. 4. For illustrations that cannot be satisfactorily reproduced by xerographic means, photographic prints can be purchased at additional cost and inserted into your xerographic copy. These prints are available upon request from the Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In all cases the best available copy has been filmed. University Microfilms International 300 N. Zeeb Road Ann Arbor, Ml 48106 8426353 Bayer, Randall James EVOLUTIONARY INVESTIGATIONS IN ANTENNARIA GAERTNER (ASTERACEAE: INULEAE) The Ohio State University Ph.D. 1984 University Microfilms International300 N. Zeeb Road, Ann Arbor, Ml 48106 PLEASE NOTE: In all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark V . 1. Glossy photographs or pages ______ 2. Colored illustrations, paper or print _____ 3. Photographs with dark background _____ 4. Illustrations are poor copy ______ 5. Pages with black marks, not original copy ______ 6. Print shows through as there is text on both sides of page _____ 7. Indistinct, broken or small print on several pages ^ 8. Print exceeds margin requirements _____ 9. Tightly bound copy with print lost in spine ______ 10. Computer printout pages with indistinct print 11. Page(s) ____________lacking when material received, and not available from school or author. 12. Page(s) ____________seem to be missing in numbering only as text follows. 13. Two pages numbered . Text follows. 14. Curling and wrinkled pages ______ 15. Other________________________________________________________________________ University Microfilms International EVOLUTIONARY INVESTIGATIONS IN ANTENNARIA GAERTMIR (ASTERACEAEs INULEAE). DISSERTATION Presented in Partial Fullfillntenfc of the Requiranents for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University by Randall James Bayer, B.S., M.S. ***** The Ohio State University 1984 Reading Committees Approved bys Dr. Daniel J. Cra&rford Dr. Gary L. Floyd Dr. Katherine L. Gross Dr. Tod F. Stuessy idvisor Dept, of Botany Dedicated to Prof o Go Ledyard Stebbins, who first introduced me to the genus Mtennaria and whose continued support and invaluable discussions have provided a constant source of inspiration. ACKNOHLEDGMENTS Numerous people should bs acknowledged for their contributions during the tenure of this studio Dr. Daniel J. Crawford? ny major professor ? is gratefully acknowledged for his continued encouragenenfc and guidance for the past six years. Members of ray committee? Drs. Daniel J. Crawford, Gary L. Floyd. Katherine L. Gross? and Tod F. Stuessy are given special thanks for their comments and criticisms on this dissertation. This study was supported by grants from Sigma Xi? The American Alpine Club, and NSF doctoral dissertation improvenent grant #DEB~82fl0359. I am especially grateful to Joe L. Bruner and G. Ledyard Stebbins for their aid and companionship in securing many of the collections. The following people are also gratefully acknowledged for their collecting efforts from various regions: L. Bayer? P. Bierzychudek. R. Bittman? J. Canne? D. Crawford, M. Galligan, H. Iltis? J. LaDuke? R. Pilatcwski? R. Stebbins? K. Urbanska? R. Whitkus? and E. Williams. I also appreciate the invaluable discussions with my fella*? graduate students in systematica and ecology at Ohio State. Special thanks are extended to Dr. Robert Warmbrodt for his advise concerning illustrations contained in this dissertation. I also wish to thank my family for their continued support and encouraganent. iii VITA. July 13, 1955................Born - Buffalo, N w York* 1978.oeeooooooooooooeooooooe.B.S., Cornell University, Ithaca, New York. 1978-1979....................Graduate Teaching Assistant, Hie Ohio State University, Columbus. 1980.........................M.S., Hie Ohio State University, Columbus. 1979-1982.... .... ........ Graduate Research Assistant, Hie Ohio State University, Columbus. 1982-1983....................Graduate Teaching Assistant, Hie Ohio State University, Columbus. 1980. ........ ...Sigma Xi Research Grant. 1980......... ..... ..... American Alpine Club Research Grant. 1982-1984 ... ............ NSF Doctoral Dissertation Improvement Grant. 1983-1984....................Presidential Fellow, Hie Ohio State University, Columbus. PUBLICATIONS Bayer, R.J. and G.L. Stebbins. 1981. Chromosome numbers of North American species of Antennaria Gaertner (Asteraceae? Inuleae). Amer. J. Bot. 68:1342-49. Crawford, D.J. and R.J. Bayer. 1981. Allozyme divergence in Coreopsis cyclocarpa (Ccmpositae). Syst. Bot. 6:373-379. Bayer, R.J. and G.L. Stebbins. 1982. A revised classification of Antennaria (Asteraceae: Inuleae) of the eastern United States. iv Sysfco Bofco 7:300-313. Bayer, R.J. and G.L. Stebbins. 1983. Distribution of sexual and asexual populations of Antennaria parlinii. Evolution. 37:555-561. Bayer, R.J. 1984. Chromosome numbers and taxonomic notes for North American species of M t e n m r i a (Asteraceae: Inuleae). Syst. Bot. 9:74-83. Price, Ho J., Crawford, D.J., and R.J. Bayer. In press, n contents of Caresegis. xmesensfiidea and Ck meseagis (Asteraceae), a progenitor-derivative species pair. Bot. Gaz. PAPERS READ AT EKQFESSIONAL MEETINGS: Bayer, R.J. and G.L. Stebbins. Population structure and asexual reproduction in Anfcsmaria Earlinii of Ohio. At ic-SEB II, Vancouver B.C., July 1980. Bayer, R.J. and G.L. Stebbins. Geographic distribution of chromosome numbers, sex ratios, and aponictic reproduction in Antennaria of the eastern United States. At IC-SEB II, Vancouver, B.C., July 1980. Stebbins. G.L. and R.J. Bayer. Sexual and asexual reproduction in Antennaria and the origin of sex. At IC-SEB II, Vancouver, B.C.. July 1980. Crawford, D.J. and R.J. Bayer. Allozyme variation in Coreopsis cyclocarpa (Ccmpositae; Heliantheae). At AIBS, Bloomington, Indiana. LaDuke, J., Nescm, G., Crawford, D., and R.J. Bayer. The biology of Trillium nivaie Riddell. At AIBS, Bloomington, Indiana. Bayer, R.J. On the possible origins of the heteroploid agamic Antennaria Parlinli complex of the eastern United States. At Hie Ohio Academy of Sciences meetings, April 19©. Bayer, R.J. The compilospecies Antennaria Eaglmli and its possible relationship with certain diploid species of the genus. At AIBS, Pennsylvania State University, August 19©. Bayer, R.J. The compilospecies Antennaria neodioisa and its possible relationship to diploid species of the genus. At AIBS, Grand Forks, North Dakota, August 19©. Bayer, R.J. Electrophoretic assessment of the genomic composition of the Antennaria Parlinii polyploid agamic complex. At AIBS, Fort Collins, Colorado, August 1984. FIELD OF STUDY Major Fields Systenatic Botany Evolutionary Studies in Mtennaria. Professor Daniel Cranford. TH3LE OF CONTENTS page v m . •••O...... 0 0 0 0 .00 iv LIST OF TABLES............. .. ix LIST OF FIGURES..................... ............... x INTRODUCTION ...... 1 CHAPTER I. INVESTIGATIONS INTO THE EVOLUTIONARY HISTORY OF THE POLYPLOID COMPLEXES IN ANTENNARIA (ASTERACEAE; INULEAE). I. THE NEODXQICA COMPLEX............ 4 Introduction. .... 4 Materials and Methods...................... 6 Results........ 9 Discussion .......... 17 Literature Cited........ 22 II. INVESTIGATIONS INTO THE EVOLUTIONARY HISTORY OF THE P0LYHO3ED COMPLEXES IN ANTENNARIA (ASTERACEAE; INULEAE) II. THE A. PARLINII COMPLEX.,................. 40 Introduction....................................... 40 Materials and Methods .... 43 Results.. .... 45 Discussion. .... 49 Literature Cited. ..... 58 III. ALLOZYME DIVERGENCE AMONG FIVE DIPLOID SPECIES OF ANTENNARIA AND THEIR ALLOEOLYELOID DERIVATIVES 74 Introduction. ....... 74 Materials and Methods............... 76 Results .............. 79 vii page Discussion........... ................. 82 Literature Cited............ 95 SUMMARY ......... 113 APMDIX A. Chromosome numbers and taxonomic notes for North
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