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Molekulare Systematik Der Gattung Suaeda (Chenopodiaceae) Und

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Molekulare Systematik Der Gattung Suaeda (Chenopodiaceae) Und Molekulare Systematik der Gattung Suaeda (Chenopodiaceae) und Evolution des C4-Photosynthesesyndroms Inaugural-Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) im Fachbereich Naturwissenschaften der Universität Kassel vorgelegt von: Peter Wolfram Schütze aus Halle/Saale Kassel, November 2008 Betreuer: Prof. Dr. Kurt Weising, Prüfungskommission: Prof. Dr. Kurt Weising (1. Gutachter) Prof. Dr. Helmut Freitag (2. Gutachter) Prof. Dr. Ewald Langer (Beisitzer) Dr. Frank Blattner (Beisitzer) Tag der mündlichen Prüfung: 17. Februar 2009 2 Inhaltsverzeichnis Inhaltsverzeichnis 1. Einleitung ........................................................................................................................................ 5 1.1. Vorbemerkungen.................................................................................................................... 5 1.2. Charakteristik der Suaedoideae............................................................................................. 6 1.2.1. Systematischer Überblick.............................................................................................. 6 1.2.2. Biologie, Klassifikationsmerkmale und Verbreitung der Sippen.................................... 9 1.2.3. Besonderheiten im Photosyntheseweg....................................................................... 12 1.2.4. Evolutionäre Trends innerhalb der Suaedoideae........................................................ 14 1.2.5. Theorien zur Sippenevolution - eine Synthese ........................................................... 16 1.3. Themenentwicklung und Stand der Forschung ................................................................... 18 2. Material und Methoden ................................................................................................................. 22 2.1. Systematische Methoden und Prinzipien............................................................................. 22 2.1.1. Organismenbezogene Methoden................................................................................ 22 2.1.2. Molekülbasierte Methoden .......................................................................................... 22 2.2. Pflanzenmaterial, Kulturversuche ........................................................................................ 25 2.2.1. Auswahl der Arten ....................................................................................................... 25 2.2.2. Aufsammlung, Kulturversuche und Materialkonservierung......................................... 25 2.3. Gewinnung molekularer Merkmale ...................................................................................... 27 2.3.1. DNA-Isolation .............................................................................................................. 27 2.3.2. Gelelektrophorese ....................................................................................................... 28 2.3.3. DNA-Regionen und Primer.......................................................................................... 29 2.3.4. PCR-Methodik ............................................................................................................. 32 2.3.5. DNA-Sequenzierung ................................................................................................... 34 2.4. Auswertung der molekularen Daten..................................................................................... 36 2.4.1. Datenauswahl.............................................................................................................. 36 2.4.2. Sequenz-Alignment ..................................................................................................... 36 2.4.3. Phylogenetische Berechnungsverfahren .................................................................... 38 2.4.4. Molekulare Uhr und Zeitdatierung............................................................................... 45 2.4.5. Test- und Überprüfungsverfahren ............................................................................... 47 2.5. Bewertung morphologischer Merkmale ............................................................................... 49 3. Ergebnisse.................................................................................................................................... 51 3.1. Ausprägung und Variabilität der molekularen Merkmale..................................................... 51 3.1.1. DNA-Isolation, Amplifikation und Sequenzierung ....................................................... 51 3.1.2. Strukturmerkmale der untersuchten DNA-Regionen .................................................. 52 3.2. Phylogenetische Rekonstruktion.......................................................................................... 55 3.2.1 Stellung der Suaedoideae innerhalb der Chenopodiaceae ........................................ 55 3.2.2. ITS-Phylogenie der Suaedoideae ............................................................................... 59 3.2.3. Chloroplasten-Phylogenie der Suaedoideae .............................................................. 68 3.2.4. Phylogenie und Diversität der Sektion Brezia............................................................. 75 3.3. Altersdatierung und Molekulare Uhr .................................................................................... 81 3.4. Evolution von C4-Photosynthese und Blatttypen ................................................................. 87 3.5. Merkmalsentwicklung anhand molekularer Phylogenien..................................................... 93 3 Inhaltsverzeichnis 4. Diskussion .................................................................................................................................... 97 4.1. Qualität, Aussagekraft und Grenzen der Daten und Methoden........................................... 97 4.1.1. Auswahl und Umfang der Sippen................................................................................ 97 4.1.2. Vergleichende DNA-Sequenzierung ........................................................................... 98 4.1.3. Phylogenetische Rekonstruktionsverfahren und Datierungsmethoden ...................... 99 4.2. Inter- und intraspezifische Variabilität................................................................................ 100 4.3. Phylogenie und Artbildung innerhalb der Suaedoideae .................................................... 101 4.3.1. Molekulare Evolution ................................................................................................. 101 4.3.2. Radiation, Gruppenbildung und Speziation............................................................... 103 4.3.3. Hybridisierung und Polyploidisierung ........................................................................ 104 4.4. Merkmalsentwicklung innerhalb der Suaedoideae ............................................................ 107 4.4.1. Systematisch relevante Merkmale und ihre Evolution .............................................. 107 4.4.2 Evolution der C4-Photosynthese ............................................................................... 111 4.5. Ableitung systematischer Einheiten................................................................................... 111 4.5.1. Position und Gliederung der Suaedoideae ............................................................... 111 4.5.2. Arten und systematische Gruppen in der Gattung Suaeda ...................................... 112 4.5.2.1. Brezia-Gruppe ........................................................................................................... 112 4.5.2.2. Schoberia-Gruppe inkl. Alexandra lehmannii............................................................ 113 4.5.2.3. Fruticosa-Gruppe ...................................................................................................... 113 4.5.2.4. Physophora-Gruppe.................................................................................................. 113 4.5.2.5. Suaeda-Gruppe......................................................................................................... 114 4.5.2.6. Schanginia-Gruppe mit Borszczowia aralocaspica................................................... 114 4.5.2.7. Suaeda glauca .......................................................................................................... 114 4.5.2.8. Bienertia .................................................................................................................... 114 4.6. Vorschläge zur Neugliederung der Suaedoideae .............................................................. 115 5. Zusammenfassung ..................................................................................................................... 118 6. Literaturverzeichnis..................................................................................................................... 121 Anhang ................................................................................................................................................ 131 Anhang 1 Abbildungsverzeichnis............................................................................................... 131 Anhang 2 Tabellenverzeichnis................................................................................................... 132 Anhang 3 Abkürzungsverzeichnis: ............................................................................................ 133
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