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Tree diversity effects on species richness and infestation of foliar fungal pathogens in European tree diversity experiments Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Naturwissenschaftlichen Fakultät I – Biowissenschaften – der Martin-Luther-Universität Halle-Wittenberg, vorgelegt von Frau Dipl.-Biol. Lydia Hantsch geboren am 27.05.1986 in Löbau Gutachter 1. Prof. Helge Bruelheide (MLU Halle-Wittenberg) 2. Prof. Uwe Braun (MLU Halle-Wittenberg) 3. Prof. Christopher C Mundt (Oregon State University) Halle (Saale), 24.09.2013 (Einreichungsdatum) Datum der Verteidigung: 10.12.2013 “When we try to pick out anything by itself, we find it hitched to everything else in the universe.” John Muir (1838-1914, American environmentalist, naturalist, writer and scientist) | 3 CONTENTS Summary ................................................................................................................................... 6 Zusammenfassung .................................................................................................................... 8 1. Introduction ........................................................................................................................ 11 1.1 Tree diversity effects on foliar fungal pathogens .......................................................... 13 1.2 Foliar fungal pathogen analyses in tree diversity experiments ...................................... 19 1.3 Objectives of the thesis .................................................................................................. 28 2. Species richness and species identity effects on occurrence of foliar fungal pathogens in a tree diversity experiment ........................................................................................... 31 2.1 Abstract .......................................................................................................................... 31 2.2 Introduction ................................................................................................................... 32 2.3 Material and Methods .................................................................................................... 35 2.4 Results ........................................................................................................................... 38 2.5 Discussion ...................................................................................................................... 40 2.6 Acknowledgments ......................................................................................................... 42 2.7 Appendices .................................................................................................................... 43 3. Effects of tree diversity, host and non-host proportion on foliar fungal pathogens .... 45 3.1 Abstract .......................................................................................................................... 45 3.2 Introduction ................................................................................................................... 46 3.3 Material and Methods .................................................................................................... 50 3.4 Results ........................................................................................................................... 53 3.5 Discussion ...................................................................................................................... 58 3.6 Acknowledgments ......................................................................................................... 64 4. No plant functional diversity effects on foliar fungal pathogens in experimental tree communities ....................................................................................................................... 65 4.1 Abstract .......................................................................................................................... 65 4.2 Introduction ................................................................................................................... 66 4.3 Material and Methods .................................................................................................... 69 4.4 Results ........................................................................................................................... 72 4.5 Discussion ...................................................................................................................... 78 4.6 Acknowledgments ......................................................................................................... 82 4.7 Appendices .................................................................................................................... 82 CONTENTS | 4 5. Tree clone diversity effects on foliar fungal pathogens in a birch-clone diversity experiment .......................................................................................................................... 87 5.1 Abstract .......................................................................................................................... 87 5.2 Introduction ................................................................................................................... 88 5.3 Material and Methods .................................................................................................... 91 5.4 Results ........................................................................................................................... 93 5.5 Discussion .................................................................................................................... 101 5.6 Acknowledgments ....................................................................................................... 104 6. Synthesis ............................................................................................................................ 105 6.1 General discussion ....................................................................................................... 105 6.2 Outlook ........................................................................................................................ 115 References ............................................................................................................................. 117 Acknowledgements ............................................................................................................... 132 Appendix ............................................................................................................................... 134 Curriculum vitae ................................................................................................................ 134 List of publications ............................................................................................................ 136 Conference contributions ................................................................................................... 137 Eigenständigkeitserklärung ............................................................................................... 138 | 5 SUMMARY Foliar fungal pathogens are responsible for a broad range of damage on trees. The top-down control of foliar fungal pathogens on plant diversity and composition has been extensively investigated, in particular the effects on biomass and yield losses in natural and agricultural ecosystems. Conversely, comprehensive knowledge on whether plant ecosystem diversity and composition control foliar fungal pathogen richness and infestation is still missing. This applies in particular to forest ecosystems. An experimental set-up is required to systematically investigate such bottom-up effects, since plant diversity experiments comprise a plant diversity gradient, as well as controlled environmental conditions. This thesis is aimed at surveying the influence of tree diversity on richness and infestation of foliar fungal pathogens. The particular experimental design of different European tree diversity experiments offered the potential to consider different aspects of tree diversity. Tree species diversity was assessed at the BIOTREE experimental site at Kaltenborn (BK) and the Kreinitz experiment (K), whereas functional tree diversity was investigated at the BIOTREE experimental site at Bechstedt (BB) and tree clone diversity at the Satakunta experiment (S). Considering further aspects of tree diversity, i.e. tree species and tree clone identity effects (BK, BB, K, S), as well as density effects of host and non-host tree species (K) and tree clones (S) were surveyed. Pathogen species richness and pathogen load of foliar fungal pathogens were determined in a microscopic and macroscopic approach. Here, four different spatial scales were investigated: The community level examined the impact of the whole tree community, including all fungus species across all tree species/tree clones. The local neighbourhood focussed beyond that on the influence of the composition of the nearest neighbour tree individuals on a target tree individual. With this focus, the individual level considered all fungus species on all target individuals across all tree species/tree clones, the tree species/tree clone level regarded all fungus species on all target individuals of a particular tree species/tree clone and the fungus species level focused on a particular fungus species across all target individuals of its tree host. In the first hypothesis it is assumed