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The Gene-Ecology of Crepis Nana Richardson and Crepis Elegans Hooker in Arctic and Alpine North America

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The Gene-Ecology of Crepis Nana Richardson and Crepis Elegans Hooker in Arctic and Alpine North America AN ABSTRACT OF THE THESIS OF ALLAN HERBERT LEGGE for the DOCTOR OFPHILOSOPHY (Name) (Degree) in GENETICS presented on May 7,1971 (Major) (Date) Title: THE GENE-ECOLOGY OF CREPIS NANARICHARDSON AND CREPIS ELEGANS HOOKER IN ARCTIC AND ALPINE NORTH AMERICA Abstract approved:Redacted for privacy Kenton Lee Chambers A gene-ecological transplant study was made on populationsof the Crepis nana and C. elegans complex from theArctic and Alpine of North America. Transplants were collected from theBrooks Range, Eagle Summit, and Alaska Range in Alaska, the RockyMountains in Alberta, the Olympic Mountains in Washington, the WallowaMoun- tains in Oregon, and the Sierra Nevada Mountains ofCalifornia. Chromosome numbers were found to be uniformly2N=14 throughout the entire range of both species. The morphological variability present in Crepis nanais shown to be ecotypical and correlatedwith habitat type.Crepis nana ssp. typica Babcock is here divided into a taprooted rivergravel ecotype, with inflated fistulous caudex, and a creepingrhizomatous talus eco- type, with narrowly inflated fistulous caudex, C. nana ssp.clivicola Legge. The subspecific concept of the creepingrhizomatous C. nana ssp. ramosa Babcock which lacks a fistulous caudex is enlarged. The pattern of major and minor ribs on achenes and the number of major ribs at the point of attachment to the receptacle are shown to be excellent ecotypic markers.All ecotypes were found to be naturally self-pollinating.Cross-pollinations between ecotypes re- vealed at low frequency a splitting of the fruit coats.This splitting was taken as a measure of hybrid vigor and heterosis and hence genetic compatibility.The suggestion is made that this may be mor- phological evidence for mitochondrial heterosis. Comparative growth chamber experiments on transplants from the field were used to show by statistical methods the degree of phenotypic plasticity and the genetically controlled differences in the achenes of both Crepis elegans and the ecotypes of Crepis nana. Discriminant functions were generated from achene characteristics using the program *BMDO5M on the CDC 3300 and shown to be statis- tically significant. The possible evolutionary history of both species is presented with special reference to the Bering Sea Land Bridge, the Alaskan refugium, central. Asia and Siberia. 01971 ALLAN HERBERT LEGGE ALLRIGHTS RESERVED The Gene-Ecology of Crepis nana Richardson and Crepis elegans Hooker in Arctic and Alpine North America by Allan Herbert Legge A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 1971 APPROVED: Redacted for privacy Professor of Botany in charge of major Redacted for privacy Director orthe Genetics "14stitute Redacted for privacy Dean of the Graduate School Date thesis is presented May 7,1971 Typed by Donna L. Olson for Allan Herbert Legge Dedicated to my Parents andGrandparents for their patiencewithout whose efforts this research might never havebeen conceived. ACKNOWLEDGEMENTS I wish to express my sincere thanks to Dr. K'nton L. Chambers for his patience, understanding and guidance during my graduate study.His constant encouragement and willingness to both listen and to express new ideas made the difficult times a littlebrighter. The author would like to acknowledge the following people who made this research possible:Dr. W. D. Billings of Duke University for his enthusiastic and continued encouragement; Dr. Arvo Kallio, University of Minnesota (formerly of the University of Alaska) and Dr. L. Viereck, University of Alaska, for making transplant storage facilities available in Alaska; Dr. M. E. Britton and Dr. Max Brewer for use of the NARL wanigan at Anaktuvuk Pass in1968; Mr. Roy Mil lice, Dome Petroleum Ltd., for logistic support to andfrom Melville Island in the Canadian Arctic Archipelago; Dr. Helge Irgens -Moller, Oregon State University, whose growth chambers made this study possible; Mr. Pete Sanchez, ChiefNaturalist, and Mr. Jerry Hassinger, Igloo Creek Ranger, Mt. McKinleyNational Park, without whose cooperation the polymorphism of the talus eco- type of Crepis nana might have gone undetected; the curatorsof the many herbaria from which specimens wereborrowed; the Park Per- sonnel of Olympic National Park, Washington; and Dr. Ralph Bogart who had confidence in my abilities when very few people did. I gratefully acknowledge my many friends who put upwith my interests and offered many helpful suggestions regardlessof how high they had to climb: Steve Mason, Robert Whitacre,Alice Lindahl, Dennis Green ley and Gail Foster. A very special thanks to Mr. Kay Fernald who taught me to appreciate photography, to Mr. E. J. Henkel whosedrafting skill is responsible for the excellent base maps, to Mrs.Jackie Atzett for her fine artistic handling of the achene illustrationsand Mr. Al Soeldner for his assistance with the Scanning ElectronMicroscope. Funds for this research were probided in part by National Defense Education Act Title IV Fellowship #66-04460. TABLE OF CONTENTS Page INTRODUCTION 1 METHODS AND MATERIALS 5 Field Site Determinations 5 Transplanting Procedure 5 Artificial Environments and Programing of Growth Chambers 8 Coding System for Plants and Population 14 Cross - Pollination Technique 16 Achene Collection and Storage 17 Achene Photography 17 Cytological Technique 18 Scanning Electron Microscopy (SEM) 18 Study of Herbarium Specimens 19 Statistical Analysis 20 Description of Programs 21 RESULTS AND DISCUSSION 22 General Habitat Descriptions 22 Achene Variation 40 Achene Germination and Environmental Adaptations 54 Breeding Systems and Floral Induction 60 Fruit-Set Variation 69 Statistical Analysis of Achene Variation 72 Discriminant Analysis 127 Genetic Relationships 144 TAXONOMY 151 Chromosome Counts 151 Description of the Species and Subspecies 154 CONCLUSION 177 BIBLIOGRAPHY 184 APPENDIX I.Mean achene weights in milligrams from populations of C. nana and C. elegans grown under different environmental conditions. 194 LIST OF TABLES Table Page 1. Populations and plants used in artificial environments. 15 2. Field collecting sites. 24 3. Comparison of the rib types found in achenes of Crepis nana and Crepis elegans. 48 4. Comparison of achene ribbing patterns from selected populations of Crepis nana and Crepis elegans in North America, and Crepis nana in Asia. 49 5. Summary of the relationships of Crepis nana and Crepis elegans to their ecology as seen in the achene ribbing patterns. 53 6. One factor analysis of variance, each population over all environments (*ANOVA12). 94 7. One factor analysis of variance, each ecotype over each environment (*ANOVA12). 99 8. One factor analysis of variance, each environment over all populations (*ANOVA12). 105 9. Summary of one-factor analysis of variance in component percentages for the achene character LMax. 109 10. Summary of one-factor analysis of variance in component percentages for the achene character LNeck' 111 11. Summary of one - factor analysis of variance in component percentages for the achene character 112 Max' 12. Summary of one-factor analysis of variance in component percentages for the achene character 114 Table Page 13. Summary of one-factor analysis of variance in component percentages for the achene character WBD. 116 14. Unbalanced nested analysis of variance(*NESTA0V) for ecotypes of Crepis nana in environment 1. 121 15. Unbalanced nested analysis of variance (*NESTA0V) for ecotypes of Crepis nana in environment 2. 122 16. Unbalanced nested analysis of variance(*NESTA0V) for ecotypes of Crepis nana in environment 4. 123 17. Balanced nested analysis of variance(*NESTA0V) for populations of Crepis nana in environment 1. 124 18. Balanced nested analysis of variance(*NESTA0V) for populations of Crepis elegans in environment 1. 125 19. Two factor analysis of variance (*ANOVA12) of the three ecotypes of Crepis nana in three environ- ments. 126 20. Populations of plants used in discriminant analysis . 131 21. Split fruit coats observed in crosses between ecotypes of Crepis nana in several artifical environments. 149 22. Chromosome counts in Crepis nana and Crepis elegans. 152 LIST OF FIGURES Figure Page 1. Map showing the distribution of Crepis nana Richards. in North America. 2 2. Map showing the distribution of Crepis elegans Hook, in North America. 3 3. Hygrothermographs for three artificial environments showing photoperiod and thermoperiod. 11 4. Hygrothermographs for three artificial environments showing a constant temperature of 14. 5° C but variable photoperiods. 12 5. Map showing collecting and transplant sites for Crepis nana and Crepis elegans in North America. 23 b. Illustrations of Crepis nana habitats in Alaska. 27 7. Illustrations of Crepis elegans showing habitat, growth forms, and relationship to Crepis nana. 30 8. The Crepis nana study population at Donnelly, Alaska, showing the biennial nature of the flowering response over a two-year period. 32 9. Illustrations of Crepis nana habitats other than in Alaska. 36 10. Illustration of the achene type of Crepis nana ssp. nana. 43 11A. Illustration of the achene type of Crepis nana ssp. clivicola. 44 11B. Illustration of the achene type of Crepis nana ssp. clivicola. 45 Figure Page 1 2. Illustration of the achene type of Crepis nana ssp. ramosa. 46 13. Illustration of the achene type of Crepis elegans. 47 14. Scanning electron microscope (SEM) micro- graphs of the basal attachment points of achenes of Crepis nana and Crepis elegans. 55 1 5A&B. Habitat daylength variation
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