GENETIC AND EPIGENETIC VARIATION OF TYPICAL GRASSLAND SPECIES – HABITAT SPECIFIC PROCESSES AND IMPLICATIONS FOR CONSERVATION DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (D R. RER . NAT .) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG VORGELEGT VON ELLEN PAGEL AUS HAMBURG IM JAHR 2020 II PROMOTIONSGESUCH WURDE EINGEREICHT AM : 20.02.2020 DIE ARBEIT WURDE ANGELEITET VON : PROF . DR. CHRISTOPH REISCH UNTERSCHRIFT : ……………………………….. ELLEN PAGEL III IV Für meine Familie „Die Natur muß gefühlt werden, wer nur sieht und abstrahirt, kann ein Menschenalter, im Lebensgedränge der glühenden Tropenwelt, Pflanzen und Thiere zergliedern, er wird die Natur zu beschreiben glauben, ihr selbst aber ewig fremd sein.“ Alexander v. Humboldt, 1810 V VI TABLE OF CONTENTS DECLARATION OF MANUSCRIPTS ................................ ................................ ............................. IX ACKNOWLEDGEMENTS ................................ ................................ ................................ .......... XI SUMMARY ................................ ................................ ................................ ........................ XIII CHAPTER ONE : GENERAL INTRODUCTION ................................ ................................ .................. 1 FROM HABITATS TO WITHIN -SPECIES DIVERSITY ............................................................................... 2 SEMI -NATURAL GRASSLANDS – HOTSPOTS OF BIODIVERSITY IN CENTRAL EUROPE ................................... 3 TWO CENTURIES OF GRASSLAND HISTORY ON THE SWABIAN ALB AND THE WÜRTTEMBERGIAN ALLGÄU ..... 5 CONSERVATION GENETICS AND LANDSCAPE ECOLOGY IN GRASSLAND ECOSYSTEMS ................................. 7 THESIS OUTLINE AND RESEARCH QUESTIONS ................................................................................... 9 CHAPTER TWO : GENETIC VARIATION IN OAT -GRASS MEADOW SPECIES ................................ ........... 11 ABSTRACT ............................................................................................................................. 12 INTRODUCTION ...................................................................................................................... 13 METHODS ............................................................................................................................. 14 RESULTS ............................................................................................................................... 20 DISCUSSION ........................................................................................................................... 25 CHAPTER THREE : GENETIC VARIATION IN LITTER MEADOW SPECIE S ................................ ................ 29 ABSTRACT ............................................................................................................................. 30 INTRODUCTION ...................................................................................................................... 31 METHODS ............................................................................................................................. 33 RESULTS ............................................................................................................................... 35 DISCUSSION ........................................................................................................................... 39 VII CHAPTER FOUR : GENETIC AND EPIGENETIC VARIATION IN LINUM CATHARTICUM .............................. 43 ABSTRACT ............................................................................................................................. 44 INTRODUCTION ...................................................................................................................... 45 METHODS ............................................................................................................................. 46 RESULTS ............................................................................................................................... 49 DISCUSSION ........................................................................................................................... 53 CHAPTER FIVE : ‘G O WITH THE FLOW ’ – GENERAL DISCUSSION , CONCLUSION & PERSPECTIVES .......... 57 EFFECT OF LAND USE HISTORY , LANDSCAPE VARIABLES AND HABITAT QUALITY ..................................... 58 EPIGENETIC VARIATION IN THE VIEW OF GRASSLAND CONSERVATION ................................................. 59 CONSERVATION GENETICS IN EUROPEAN GRASSLANDS ................................................................... 60 GENETIC CONSERVATION AREAS ................................................................................................. 61 FUTURE PERSPECTIVES ............................................................................................................. 61 REFERENCES ................................ ................................ ................................ ...................... 65 SUPPLEMENTARY MATERIAL ................................ ................................ ................................ 83 DECLARATION OF MANUSCRIPTS Submitted manuscripts included in the thesis: Chapter Two was submitted with the thesis’ author as lead author: Pagel E , Lehmair T A, Poschlod P & Reisch C. Genetic variation of typical plant species in ancient and recent oat-grass meadows: investigating the effect of land use history, landscape and vegetation struc- ture. Under review at Journal of Applied Ecology . Chapter Three was submitted with the thesis’ author as co-lead author: Lehmair T A, Pagel E , Poschlod P & Reisch C. Conservation of intraspecific diversity in litter meadows – the role of habitat age, landscape configuration and environmental factors. In revisions at Conservation Genetics . Chapter Four was submitted with the thesis’ author as lead author: Pagel E , Poschlod P & Reisch C. Habitat matters – strong genetic and epigenetic differentiation among populations of Linum catharticum from dry and wet grasslands. In revision at Ecology and Evolution . Published and submitted manuscripts not included in the thesis: Lehmair T A, Pagel E , Poschlod P & Reisch C. The impact of habitat age, landscape structure, habitat quality, and population size on the genetic variation of typical calcareous grassland plant species. Sub- mitted to Landscape Ecology . Krickl P, Gilck F, Lehmair T A, Pagel E , Rubanschi S, Simmel J, Straubinger C, Fischer S & Poschlod P. Die Ausstellung „Ein Leben für die Botanik – Einzigartiger Zeitzeuge des Wandels von Lebensraum- und Artenvielfalt“. Hoppea, Denkschr. Regensb. Bot. Ges . 79 (2018): 87 – 112 IX X ACKNOWLEDGEMENTS First and foremost, I would like to express my deepest gratitude to Christoph Reisch and Peter Poschlod. They gave me the opportunity to develop my skills in the “Grassland”-project and warmly welcomed me into the ‘deepest South of Germany far away from the ‘cold northern German winds’. Christoph calmly guided me through the good and the more difficult times. He supported me in devel- oping my knowledge in molecular ecology, conservation genetics and many other things. I value his mentorship highly and am very glad he gave me so many opportunities to grow. I also want to express my gratitude to Peter, for his enthusiasm and sharing is great knowledge freely and for his feedback on the manuscripts and reports during my PhD. Special thanks to my colleagues from the “Grassland”-project Theresa Lehmair, Cornelia Straubinger, Eva Wagner, Lina Begemann & Patrizia Krickl, who helped building the framework for this thesis. A special and deep thank you is due to Lina, who was a great and necessary emotional and professional support during the last months of my thesis. Thanks also to Petra Schitko for guided me in the lab and providing all the materials and assistance during analysing my data. Sabine Fischer taught me the art of using ArcGIS and helped me a lot during the digitization of the endless historic maps. Thank you for your patience and answering even the silly questions. I also want to thank the Theoretical Ecology group led by Florian Hartig for their advice and help with analysing my data and R in general. All other colleagues I thank for creating such a great working environment and making me feel at home at the University of Regensburg. Many thanks also to the Bachelor and Master students that helped with the realization of this project, either as part of their thesis or as a student helper: Simone Fabian, Lukas Feichtmeier, Christina Fischer, Helene Kleijn, Laura Krassini, Christina Manhart, Franziska Parzefall, Marco Peschke, Laila Pfättisch, Annika Schmid, Leonie Schulz, Sebastian Segieth, Jakob Speigl & Vivien Tkcocz. My family and friends have always been there for me in every possible way and supported me through the good and the bad times of my PhD years. My parents were always the support in the background and supported my decisions and encouraged me to study in many different places and gather the ex- periences I made during my Bachelor, Master and PhD studies. Special gratitude is due to my late grandfather, who has first installed an interest in nature in me and build the starting block of me as a biologist and ecologist. XI XII SUMMARY European semi-natural grasslands are among the Similarly, Chapter Three deals with the genetic most species rich habitats in the world. They were variation in litter meadow plant species (Angelica historically