Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) an der Fakultät für Biologie der Ludwig-Maximilians-Universität München PHYLOGENY, BIOGEOGRAPHY, CLASSIFICATION, AND AMBER FOSSILS OF THE LIVERWORT FAMILIES LEJEUNEACEAE AND RADULACEAE Julia Maria Theresa Bechteler München, 07. Dezember 2017 ii für meine Familie iii iv PREFACE

 Statutory declaration Erklärung Diese Dissertation wurde im Sinne von §12 der Promotionsordnung von Prof. Dr. Jochen Heinrichs betreut. Ich erkläre hiermit, dass die Dissertation nicht einer anderen Prüfungskommission vorgelegt worden ist und dass ich mich nicht anderweitig einer Doktorprüfung ohne Erfolg unterzogen habe. Eidesstattliche Erklärung Ich versichere hiermit an Eides statt, dass die vorgelegte Dissertation von mir selbstständig und ohne unerlaubte Hilfe angefertigt wurde. Julia Bechteler, 07. Dezember 2017 (Unterschrift) 1. Gutachter: Prof. Dr. Susanne S. Renner 2. Gutachter: Prof. Dr. Wolfgang Frank Tag der Abgabe: 07. Dezember 2017 Tag der mündlichen Prüfung: 08. Februar 2018 v vi Declaration of contribution In this thesis, I present the results from my doctoral research, carried out in Munich from August 2014 to November 2017 under the guidance of Prof. Dr. Jochen Heinrichs. My thesis resulted in four manuscripts, presented in Chapters 2 to 5, of which all have been published. I generated all data and conducted all analyses myself except for Chapter 1 for which J. Heinrichs and A. Schäfer-Verwimp contributed the character matrix and G.E. Lee the drawing in Fig. 2. For Chapter 5, I conducted the analyses based on datasets provided by M.A.M. Renner, and photographs by A.R. Schmidt and J. Heinrichs. Writing and discussion involved collaboration with J. Heinrichs, with input from the co-authors. Julia Bechteler Prof. Susanne S. Renner (Signature) (Signature) List of publications Peer-reviewed journal articles BECHTELER, J., LEE, G.E., SCHÄFER-VERWIMP, A., PÓCS, T., PERALTA, D.F., RENNER, M.A.M., SCHNEIDER, H., HEINRICHS, J. 2016. Towards a monophyletic classification of Lejeuneaceae IV: reinstatement of Allorgella, transfer of Microlejeunea aphanella to Vitalianthus and refinements of the subtribal classification. Plant Systematics and Evolution, 302, 187–201. BECHTELER, J., LEE, G.E., SCHÄFER-VERWIMP, A., RENNER, M.A.M., PERALTA, D.F., HEINRICHS, J. 2016. Towards a monophyletic classification of Lejeuneaceae V: the systematic position of Pictolejeunea. Phytotaxa, 280, 259–270. BECHTELER, J., SCHÄFER-VERWIMP, A., LEE, G.E., FELDBERG, K., PÉREZ ESCOBAR, O.A., PÓCS, T., PERALTA, D.F., RENNER, M.A.M., HEINRICHS, J. 2017. Geographical structure, narrow species ranges and Cenozoic diversification in a pantropical clade of epiphyllous leafy liverworts. Ecology and Evolution, 7, 638–653. vii BECHTELER, J., SCHMIDT, A.R., RENNER, M.A.M., WANG, B., PÉREZ- ESCOBAR, O.A., SCHÄFER-VERWIMP, A., FELDBERG, K., HEINRICHS, J. 2017. A Burmese amber fossil of Radula (Porellales, Jungermanniopsida) provides insights into the Cretaceous evolution of epiphytic lineages of leafy liverworts. Fossil Record, 20, 201–213. Funding My research was partially supported by the Deutsche Forschungsgemeinschaft (DFG HE 3584/6, 2016-2018). viii CONTENTS Preface ......................................................................................................................................v Statutory Declaration .................................................................................................v Erklärung ....................................................................................................................v Eidesstattliche Erklärung ...........................................................................................v Declaration of contribution .................................................................................... vii List of publications ................................................................................................. vii Funding ................................................................................................................... viii Contents.............................................................................................................................ix Summary.............................................................................................................................1 Chapter 1: General Introduction ............................................................................................3 1.1 Epiphyllous liverworts ........................................................................................7 1.2 The study systems: Lejeuneaceae and Radulaceae ...........................................9 1.3 Evolution of Lejeuneaceae and Radulaceae.................................................... 11 1.4 Fossil record of Lejeuneaceae and Radulaceae .............................................. 12 1.5 Specific research questions .............................................................................. 14 Chapter 2: Towards a monophyletic classification of Lejeuneaceae IV: Reinstatement of Allorgella, transfer of Microlejeunea aphanella to Vitalianthus and refinements of the subtribal classification ..................................................... 17 Chapter 3: Towards a monophyletic classification of Lejeuneaceae V: The systematic position of Pictolejeunea ................................................................................ 35 Chapter 4: Geographical structure, narrow species ranges and Cenozoic diversification in a pantropical clade of epiphyllous leafy liverworts .................................. 49 Chapter 5: Burmese amber fossil of Radula (Porellales, Jungermanniopsida) provides insights into the Cretaceous evolution of epiphytic lineages of leafy liverworts ......................................................................................................... 67 Chapter 6: General Discussion ............................................................................................ 93 6.1 Phylogeny and classification of Lejeuneeae (Lejeuneaceae)......................... 95 6.2 Evolution of epiphyllous liverworts: The case of Leptolejeunea .................. 96 6.3 A Cretaceous amber fossil belonging to the genus Radula (Radulaceae) .... 99 6.4 General conclusion and perspective .............................................................. 101 References ........................................................................................................................... 103 Acknowledgements ............................................................................................................ 119 Curriculum vitae................................................................................................................ 121 ix x