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Cytotype Associations, Ecological Divergence and Genetic Variation in the Apomictic Complex Paspalum Intermedium Munro Ex Morong (Poaceae)

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Cytotype Associations, Ecological Divergence and Genetic Variation in the Apomictic Complex Paspalum Intermedium Munro Ex Morong (Poaceae) Cytotype Associations, Ecological Divergence and Genetic Variation in the Apomictic Complex Paspalum intermedium Munro Ex Morong (Poaceae) Dissertation for the award of the degree “Doctor of Philosophy” Ph.D. Division of Mathematics and Natural Sciences of the Georg-August-Universität Göttingen within the doctoral program Biology of the Georg-August University School of Science (GAUSS) Submitted by Piyal Karunarathne Göttingen, 2018 THESIS COMMITTEE Prof. Dr. Elvira Hörandl Department of Systematics, Biodiversity and Evolution of Plants (with herbarium) Albrecht-von-Haller Institute for Plant Sciences University of Göttingen, Germany Prof. Dr. Holger Kreft Department of Biodiversity, Macroecology & Biogeography Faculty of Forest Sciences and Forest Ecology University of Göttingen, Germany Dr. Diego Hojsgaard Department of Systematics, Biodiversity and Evolution of Plants (with herbarium) Albrecht-von-Haller Institute for Plant Sciences University of Göttingen, Germany MEMBERS OF THE EXAMINATION BOARD Reviewer Prof. Dr. Elvira Hörandl Department of Systematics, Biodiversity and Evolution of Plants (with herbarium), Albrecht-von-Haller Institute for Plant Sciences University of Göttingen, Germany Second Prof. Dr. Holger Kreft reviewer Department of Biodiversity, Macroecology & Biogeography Faculty of Forest Sciences and Forest Ecology University of Göttingen, Germany Further members of the Examination Board Prof. Dr. Stefan Scheu J.F. Blumenbach Institute of Zoology and Anthropology University of Göttingen Prof. Dr. Mark Maraun J.F. Blumenbach Institute of Zoology and Anthropology University of Göttingen Prof. Dr. Thomas Friedl Dept. EPSAG University of Göttingen Dr. Sven Bradler J.F. Blumenbach-Institut für Zoologie und Anthropologie University of Göttingen ii Acknowledgments I would like to place on record my sincere gratitude to Dr. Diego Hojsgaard for choosing me to carry out this research project, his excellent supervision, tremendous help and advice, and for remaining unflappable despite my various crisis. My thanks to Prof. Elvira Hörandl and Prof. Holger Kreft for invaluable discussions throughout the project to improve its quality. Funding for this project came mainly from the German Research foundation (Deutsche Forschungsgemeinschaft) and National Scientific and Technical Research Council of Argentina (DFG–MINCyT-CONICET bilateral collaboration [HO5462-1/1 and RD- 20150202-0167 to D.H. and E.M.]). Field work and plant cultivation assessment would have been impossible without the invaluable support from Eric Martinez, Ana Honfi, Verena Reutemann and Adriana Glücksberg. I thank Silvia Friedrichs and Sabine Schmidt for maintenance of live plant stocks. Special thanks to Gabriel Rua for allowing me to use herbarium facilities at Faculty of Agronomy, University of Buenos Aires. My family and friends have always been there for me whenever I needed help, advice and moral support. Finally I would like to express my gratitude to everyone else who helped me in many ways to accomplish my goals. Piyal Karunarathne 2018, Göttingen iii Paspalum intermedium Photo by Diego Hojsgaard iv TABLE OF CONTENT ACKNOWLEDGMENTS ............................................................................................................... III TABLE OF CONTENT .............................................................................................................. V LIST OF TABLES .................................................................................................................... VII TABLE OF FIGURES.............................................................................................................. VIII ABSTRACT .............................................................................................................................. 1 CHAPTER 1 .............................................................................................................................. 3 1. INTRODUCTION ............................................................................................................... 3 1.1. Polyploidization and Plant Evolution ....................................................................... 3 1.2. Apomixis ................................................................................................................ 9 1.3. Paspalum intermedium Munro ex Morong ............................................................. 16 1.4. Aims of the Project ............................................................................................... 17 CHAPTER 2 ............................................................................................................................ 19 2. DISTRIBUTION PATTERNS AND ECOLOGICAL DIFFERENTIATION IN DIFFERENT CYTOTYPES OF PASPALUM INTERMEDIUM ..................................................................... 19 2.1. ABSTRACT ........................................................................................................... 19 2.2. INTRODUCTION ................................................................................................. 20 2.3. MATERIALS AND METHODS ............................................................................... 21 2.4. RESULTS .............................................................................................................. 27 2.5. DISCUSSION ....................................................................................................... 34 CHAPTER 3 ............................................................................................................................ 39 3. EVOLUTIONARY IMPLICATIONS OF A TUG OF WAR BETWEEN SEXUAL AND APOMICTIC REPRODUCTIVE MODES IN PASPALUM INTERMEDIUM (POACEAE) LEADING TO FITNESS VARIATION IN THE POLYPLOID COMPLEX ..................................................................................................... 39 3.1. ABSTRACT ........................................................................................................... 39 3.2. INTRODUCTION ................................................................................................. 40 3.3. MATERIALS AND METHODS ............................................................................... 42 3.4. RESULTS .............................................................................................................. 47 3.5. DISCUSSION ....................................................................................................... 55 CHAPTER 4 ............................................................................................................................ 61 4. POPULATION GENETIC STRUCTURE ANALYSIS ECHOES THE DISTRIBUTION, COEXISTENCE AND NICHE DIVERGENCE OF CYTOTYPES IN THE POLYPLOID GRASS SPECIES PASPALUM INTERMEDIUM (POACEAE)61 4.1. ABSTRACT ........................................................................................................... 61 v 4.2. INTRODUCTION ................................................................................................. 62 4.3. MATERIALS AND METHODS ............................................................................... 65 4.4. RESULTS .............................................................................................................. 68 4.5. DISCUSSION ....................................................................................................... 72 4.6. CONCLUSIONS ................................................................................................... 78 CHAPTER 5 ............................................................................................................................ 79 5. GENERAL DISCUSSION .................................................................................................... 79 5.1. Cytotype Composition in Paspalum intermedium .................................................. 79 5.2. Spatial and Geographical Distribution of Cytotypes and Their Ecological Importance80 5.3. Reproductive Modes, Competition and Reproductive Fitness ................................. 81 5.4. Population Structure and Genetic Composition ..................................................... 82 6. LITERATURE CITED ........................................................................................................... 84 SUPPLEMENTARY DATA ...................................................................................................... 99 LIST OF PUBLICATIONS ........................................................................................................... 123 CURRICULUM VITAE .............................................................................................................. 123 vi LIST OF TABLES Table 2.1. Summary of ecoregions, collection sites, number of individuals and ploidy levels of the P. intermedium plants analysed. ........................................................................................ 22 Table 2.2. Bioclimatic and environmental variables retained (after multivariate analysis and binomial logistic regression) for cytotype distribution and niche analysis in P. intermedium, its significance values, and PCA contributions. ....................................................................... 26 Table 2.3. Reproductive mode variation between cytotypes of P. intermedium in the study area .. 31 Table 3.1. Proportions of meiotic/sexual and apomictic at embryo sac and seed stages in studied populations of P.intermedium. Proportions were calculated as a fraction of all the observed ES/seeds. Chi-squared values
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