SYSTEMATICS, PHYLOGENY AND BIOGEOGRAPHY OF COUSINIA (ASTERACEAE) Dissertation Zur Erlangung des Grades Doktor der Naturwissenschaften am Fachbereich Biologie der Johannes Gutenberg‐Universität Mainz Iraj Mehregan geb. in Gachsaran, Iran Mainz, 2008 Dekan: 1. Berichterstatter: 2. Berichterstatter: Tag der mündlichen Prüfung: 10. 07. 2008 II Kapitel 2 (chapter 2) dieser Arbeit ist im Druck bei “Taxon“: López‐Vinyallonga, S., Mehregan, I.*, Garcia‐Jacas, N., Tscherneva, O., Susanna, A. & Kadereit, J. W.*: Phylogeny and evolution of the Arctium‐Cousinia complex (Compositae, Cardueae‐Carduinae). * Von den Autoren Mehregan, I und Kadereit, J. W.: Die Generation der ITS‐Sequenzen von 113 Taxa (Appendix 1), die Bayesische Analyse der ITS‐ und rpS4‐trnT‐trnL‐Sequenzen, das Rechnen der Molekularen Uhr sowie der Partition‐Hemogenity Test und die Analyse des Cousinioid Clade wurde in Rahmen dieser Dissertation ausgeführt. Das Manuskript wurde in Zusammenarbeit aller Autoren geschrieben. Kapitel 3 (chapter 3) diese Arbeit wird bei “Willdenowia” eingereicht: Mehregan, I. & Kadereit, J. W.: The role of hybridization in the evolution of Cousinia s.s. (Asteraceae). Kapitel 4 (chapter 4) dieser Arbeit ist im Druck bei “Willdenowia“: Mehregan, I. & Kadereit, J. W.: Taxonomic revision of Cousinia sect. Cynaroideae. III Contents SUMMARY............................................................................................................................................................................1 ZUSAMMENFASSUNG .....................................................................................................................................................2 CHAPTER 1 ...........................................................................................................................................................................3 GENERAL INTRODUCTION............................................................................................................................................3 1.1 COUSINIA WITHIN ASTERACEAE........................................................................................................................3 1.2 TAXONOMY OF COUSINIA ....................................................................................................................................5 1.3 CENTRES OF BIODIVERSITY ..................................................................................................................................6 1.4 HYBRIDIZATION ......................................................................................................................................................7 1.5 SPECIES NUMBERS IN COUSINIA .........................................................................................................................8 1.6 AIMS OF THIS WORK ...............................................................................................................................................8 1.7 ACKNOWLEDGEMENTS ........................................................................................................................................9 1.8 REFERENCES ...........................................................................................................................................................10 CHAPTER 2 .........................................................................................................................................................................13 PHYLOGENY AND EVOLUTION OF THE ARCTIUM‐COUSINIA COMPLEX (COMPOSITAE, CARDUEAE‐CARDUINAE).............................................................................................................................................13 2.1 ABSTRACT................................................................................................................................................................13 2.2 INTRODUCTION.....................................................................................................................................................14 2.3 MATERIAL AND METHODS ................................................................................................................................17 2.3.1 Plant material .....................................................................................................................................................17 2.3.2 DNA extraction, amplification and sequencing..................................................................................................17 2.3.3 Analyses..............................................................................................................................................................18 2.4 RESULTS....................................................................................................................................................................20 2.4.1 Partition homogeneity test..................................................................................................................................20 2.4.2 Phylogenetic analysis..........................................................................................................................................20 2.4.3 Molecular Clock Analysis ...................................................................................................................................27 2.5 DISCUSSION ............................................................................................................................................................29 2.5.1 The Arctium‐Cousinia complex ..........................................................................................................................29 2.5.2 The Arctioid clade ...............................................................................................................................................30 2.5.3 The Cousinioid clade ...........................................................................................................................................32 2.5.4 Cousinia tenella...................................................................................................................................................36 2.6 ACKNOWLEDGEMENTS ......................................................................................................................................37 2.7 LITERATURE CITED...............................................................................................................................................37 2.8 APPENDIX 1.............................................................................................................................................................42 2.9 APPENDIX 2.............................................................................................................................................................46 CHAPTER 3 .........................................................................................................................................................................48 THE ROLE OF HYBRIDIZATION IN THE EVOLUTION OF COUSINIA S.S. .....................................................48 3.1 ABSTRACT................................................................................................................................................................48 3.2 INTRODUCTION.....................................................................................................................................................49 3.3 MATERIAL AND METHODS.................................................................................................................................50 3.4 RESULTS....................................................................................................................................................................50 3.5 DISCUSSION.............................................................................................................................................................54 3.6 ACKNOWLEDGEMENTS ......................................................................................................................................60 3.7 REFERENCES ...........................................................................................................................................................60 2.8 APPENDIX 1 .............................................................................................................................................................62 CHAPTER 4 .........................................................................................................................................................................68 TAXONOMIC REVISION OF COUSINIA SECT. CYNAROIDEAE .........................................................................68 4.1 ABSTRACT................................................................................................................................................................68 IV 4.2 INTRODUCTION.....................................................................................................................................................69 4.2.1 Taxonomic history of C. sect. Cynaroideae .........................................................................................................69 4.2.2 Monophyly of C. sect. Cynaroideae.....................................................................................................................70 4.2.3 Geographical distribution, biology, chromosome