Vegetative reproduction and clonal diversity in pleurocarpous mosses (Bryophytina) of mesic habitats A combined molecular and morpho-anatomical study in Pseudoscleropodium purum (Hedw.) M. Fleisch. ex Broth. (Brachytheciaceae), Pleurozium schreberi (Brid.) Mitt. (Hylocomiaceae) and Rhytidiadelphus squarrosus (Hedw.) Warnst. (Hylocomiaceae) Dissertation zur Erlangung des akademischen Grades des Doktors der Naturwissenschaften (Dr. rer. nat.) eingereicht im Fachbereich Biologie, Chemie, Pharmazie der Freien Universität Berlin vorgelegt von Sebastian Fritz Geburtsort Berlin Berlin 2009 Die Arbeit wurde im Zeitraum Nov. 2005 – Nov. 2009 unter Leitung von Prof. Dr. W. Frey am Institut für Systematische Botanik und Pflanzengeographie, der Freien Universität Berlin angefertigt Gutachter: 1. Prof. Dr. Wolfgang Frey 2. Prof. Dr. Jürgen Schmitt Tag der mündlichen Prüfung 23.04.2010 Für meine Großeltern und Eltern, im Besonderen in Gedenken an meinen Großvater Bernhard Fritz der mich auf den Weg zur Botanik gebracht hat. Contents Contents INDEX OF FIGURES ......................................................................................................................... III INDEX OF TABLES ........................................................................................................................... VI ABBREVIATIONS ............................................................................................................................ VII 1 INTRODUCTION .............................................................................................................................. 1 2 REPRODUCTION AND DISPERSAL ............................................................................................ 3 2.1 Vegetative Reproduction .............................................................................................................................. 3 2.2 Diaspore dispersal ........................................................................................................................................ 5 3 MATERIALS AND METHODS ....................................................................................................... 7 3.1 Studied species ............................................................................................................................................. 7 3.1.1 Pseudoscleropodium purum ................................................................................................................. 7 3.1.1.1 Morphology .................................................................................................................................................... 7 3.1.1.2 Reproduction .................................................................................................................................................. 8 3.1.1.3 Ecology ........................................................................................................................................................... 9 3.1.1.4 Plant communities .......................................................................................................................................... 9 3.1.1.5 General distribution ........................................................................................................................................ 9 3.1.2 Pleurozium schreberi .......................................................................................................................... 11 3.1.2.1 Morphology .................................................................................................................................................. 11 3.1.2.2 Reproduction ................................................................................................................................................ 11 3.1.2.3 Ecology ......................................................................................................................................................... 12 3.1.2.4 Plant communities ........................................................................................................................................ 12 3.1.2.5 General distribution ...................................................................................................................................... 14 3.1.3 Rhytidiadelphus squarrosus................................................................................................................ 15 3.1.3.1 Morphology .................................................................................................................................................. 15 3.1.3.2 Reproduction ................................................................................................................................................ 15 3.1.3.3 Ecology ......................................................................................................................................................... 16 3.1.3.4 Plant communities ........................................................................................................................................ 17 3.1.3.5 General distribution ...................................................................................................................................... 17 3.2 Study areas ................................................................................................................................................. 18 3.2.1 Topography ......................................................................................................................................... 18 3.2.1.1 Thuringia (Plots Sil1, Sil2 and Sil3) ............................................................................................................. 18 3.2.1.2 Berlin (Plot B1) ............................................................................................................................................ 21 3.2.1.3 Brandenburg (Plots NH1 and Saarm1) ......................................................................................................... 21 3.2.2 Gap re-colonisation experiments ........................................................................................................ 22 3.2.3 Climate ............................................................................................................................................... 23 3.3 Sampling of plant material ......................................................................................................................... 24 3.3.1 Method of sampling ............................................................................................................................ 24 3.3.2 Foreign specimens .............................................................................................................................. 25 3.3.3 Identification of Rhytidiadelphus specimens ...................................................................................... 25 3.4 Morpho-anatomical analysis ...................................................................................................................... 27 3.5 Preparation of plant material for molecular analysis (AFLP) ..................................................................... 27 3.6 Molecular analysis - AFLP Fingerprinting ................................................................................................. 27 3.6.1 Method ................................................................................................................................................ 27 3.6.2 Used protocol ...................................................................................................................................... 27 3.7 Data scoring and analysis ........................................................................................................................... 31 4 RESULTS .......................................................................................................................................... 34 4.1 Morpho-anatomical analysis ...................................................................................................................... 34 4.1.1 Pseudoscleropodium purum ............................................................................................................... 34 4.1.1.1 Plot and patch descriptions ........................................................................................................................... 34 4.1.1.2 Generative reproduction ............................................................................................................................... 34 4.1.1.3 Vegetative reproduction ................................................................................................................................ 34 4.1.1.4 Gap re-colonisation ....................................................................................................................................... 36 -I- Contents 4.1.2 Pleurozium schreberi .......................................................................................................................... 38 4.1.2.1 Plot and patch descriptions ........................................................................................................................... 38 4.1.2.2 Generative reproduction ............................................................................................................................... 38 4.1.2.3 Vegetative reproduction ...............................................................................................................................