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Lamiaceae, Nepetoideae, Mentheae) Unter Besonderer Berücksichtigung Des Satureja-Komplexes

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Lamiaceae, Nepetoideae, Mentheae) Unter Besonderer Berücksichtigung Des Satureja-Komplexes Phylogenetische und taxonomische Untersuchungen an der Subtribus Menthinae (Lamiaceae, Nepetoideae, Mentheae) unter besonderer Berücksichtigung des Satureja-Komplexes Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München zur Erlangung des Doktorgrades vorgelegt von Christian Bräuchler aus Mühldorf, Deutschland München, 5. Mai 2009 1. Gutachter: Prof. Dr. Günther Heubl 2. Gutachter: Prof. Dr. Jürke Grau Tag der mündlichen Prüfung: 22. Juni 2009 Für Verena, Felix und Sebastian Abkürzungsverzeichnis: Anmerkung: Abkürzungen von Pflanzennamen-Autoren und Zeitschriftentiteln sowie Herbariumakronyme sind nicht aufgeführt, da sie sich entsprechend der gängigen Praxis nach den Standards richten, die in IPNI (2009) und BPH online (2009), sowie Index Herbariorum (Holmgren & Holmgren, 1998) angegeben sind. 2n diploider Chromosomensatz 5,8s 5,8 Svedberg-Einheiten (=Sedimentationskoeffizient bei Ultrazentrifugation) Abb. Abbildung AFLP Amplified Fragment Length Polymorphism al. alii (lateinisch für: der, die, das andere/die anderen; Abkürzung bei Autorenzitat) äther. ätherisch bp basepair (englisch für: Basenpaar; bei gepaarten DNA-Strängen) bzw. beziehungsweise BGM Botanischer Garten München ca. circa (lateinisch für: ungefähr, in etwa) CB Christian Bräuchler Co. Company cp chloroplast (englisch für: Chloroplasten-; Vorsilbe zur Spezifizierung der DNA-Herkunft) d.h. das heißt DNA Deoxy RiboNucleic Acid (englisch für: Desoxyribonukleinsäure) ed./eds. editor/editors (englisch für: der/die Herausgeber) engl. englisch EST Expressed Sequence Tag etc. et cetera (lateinisch für: und übriges) fps2 nukleäres low copy Gen gapC nukleäres low copy Gen gen. genus (lateinisch für: Gattung) inkl. inklusive ISSR Inter Single Sequence Repeats ITS Internal Transcribed Spacer k.A. keine Angabe LMU Ludwig-Maximilians-Universität (München) matK codierender Bereich des Chloroplastengenoms max. maximal Mt. Mount (englisch für: Berg) N. Numero (lateinisch für: Nummer) ndhF codierender Bereich des Chloroplastengenoms nov. novus/a/um (lateinisch für: neuer/e/es) pers. Mitt. persönliche Mitteilung P. page p.p. pro parte (lateinisch für: zum Teil) Pp. pages rbcL codierender Bereich des Chloroplastengenoms rDNA ribosomale DNA REM Raster-Elektronen-Mikroskop RJB Real Jardín Botanico (Madrid) IV RNA RiboNucleic Acid (englisch für: Ribonukleinsäure) s. siehe sect. sectio (lateinisch für: Sektion; Standardabkürzung für die taxonomische Rangstufe „sectio“) s.l. sensu lato (lateinisch für: im weit(er)en Sinn) sp./spp. species (Singular/Plural; lateinisch für: Art) ssp./subsp. subspecies (alternative verwendete Abkürzungen, lateinisch für: Unterart) s.str. sensu stricto (lateinisch für: im eng(er)en Sinn) t. tabula (lateinisch für: (Abbildungs-)Tafel) Tab. Tabelle TM Trademark trnK Intergenerischer Spacer des Chloroplastengenoms trnL-F Intergenerischer Spacer des Chloroplastengenoms trnT-L Intergenerischer Spacer des Chloroplastengenoms u.a. unter anderem v.a. vor allem vgl. vergleiche vol. volume (englisch für: Band) z.B. zum Beispiel z.T. zum Teil V Inhaltsverzeichnis ___________________________________________________________________________ Inhaltsverzeichnis 1. Einleitung ..................................................................................................................... 1 1.1 Abgrenzung und infrafamiliäre Gliederung der Lamiaceae ................................. 1 1.2 Die Subtribus Menthinae und ihre Charakterisierung ........................................... 4 1.3 Historischer Abriss über die Taxonomie des Satureja Komplexes ....................... 13 1.4 Einfluß unterschiedlicher taxonomischer Konzepte für den Satureja Komplex .. 14 auf Florenwerke des 20. Jahrhunderts 1.5 Neuere Arbeiten an den Menthinae und dem Satureja Komplex .......................... 15 1.6 Fragestellung und Zielsetzung .................................................................................. 16 2. Ergebnisse und Diskussion ..................................................................................... 17 2.1 Materialbeschaffung .................................................................................................. 17 2.2 Methodenoptimierung ............................................................................................... 17 2.3 Manuskripte ............................................................................................................... 18 2.3.1 Zusammenfassung Manuskript 1: Bräuchler, C., Meimberg, H., Abele, T. 18 & Heubl, G. 2005: Polyphyly of the genus Micromeria (Lamiaceae) – evidence from cpDNA sequence data. Taxon 54(3): 639-650. 2.3.2 Zusammenfassung Manuskript 2: Bräuchler, C., Meimberg, H. & Heubl, .. 19 G. 2006: New names in Old World Clinopodium – the transfer of the species of Micromeria sect. Pseudomelissa to Clinopodium. Taxon 55(4): 977-981. 2.3.3 Zusammenfassung Manuskript 3: Meimberg, H., Abele, T., Bräuchler, C., 19 McKay, J., K., Pérez de Paz, P. & Heubl, G. 2006: Molecular evidence for adaptive radiation of Micromeria Benth. (Lamiaceae) on the Canary Islands as inferred from chloroplast and nuclear DNA sequences and ISSR fingerprint data. Molec. Phylogen. Evol. 41: 566-578. 2.3.4 Zusammenfassung Manuskript 4: Bräuchler, C., Doroszenko, A., Esser, .... 20 H.-J. & Heubl, G. 2008: Killickia (Lamiaceae) – a new genus from KwaZulu-Natal, South Africa. Bot. J. Linn. Soc. 157: 575–586. 2.3.5 Zusammenfassung Manuskript 5: Bräuchler, C., Heubl, G. & Ryding, O. .. 21 2008: The genus Micromeria Benth. (Lamiaceae) – a synoptical update. Willdenowia 38: 363-410. VI Inhaltsverzeichnis ___________________________________________________________________________ 2.3.6 Zusammenfassung Manuskript 6: Bräuchler, C., Meimberg, H. & .............. 22 Heubl, G.: Molecular phylogeny of Menthinae (Lamiaceae, Nepetoideae, Mentheae) – taxonomy, biogeography and conflicts. Molec. Phylogen. Evol., accepted for publication. 3. Ausblick ........................................................................................................................ 25 4. Literaturverzeichnis ................................................................................................. 26 5. Abbildungs- und Tabellenverzeichnis ................................................................ 32 6. Appendix 6.1 Manuskript 1 6.2 Manuskript 2 6.3 Manuskript 3 6.4 Manuskript 4 6.5 Manuskript 5 6.6 Manuskript 6 VII Einleitung ___________________________________________________________________________ 1. Einleitung Die Lamiaceae oder Labiatae (Lippenblüter) stellen mit derzeit mehr als 7000 bekannten Species in über 230 Gattungen (Harley & al., 2004) eine abgeleitete und sehr artenreiche Familie innerhalb der Angiospermae (Bedecktsamer) dar. Es handelt sich dabei überwiegend um Kräuter oder (Halb-)Sträucher, selten auch Bäume, mit häufig vierkantigem Stängel und gegenständigen, meist dekussiert angeordneten Blättern. Die typische Infloreszenz ist ein Thyrsus mit cymösen Teilblütenständen und zygomorphen, zweilippigen Blüten. Der Fruchtknoten ist meist vierteilig gespalten mit laminar oder submarginal plazentierten Samenanlagen. Allgemein bekannt sind die Lamiaceae wegen der Verwendung vieler Arten als Gewürz- und Arzneipflanzen. Salbei, Minze, Thymian, Bohnenkraut und Basilikum sind nur einige Vertreter, die hier angeführt werden sollen. Ihre große Bedeutung verdanken sie vor allem dem Vorkommen von ätherischen Ölen, die fast ausschließlich in den subkutikulären Räumen der Drüsenschuppen sowie den Köpfchen der Drüsenhaare abgelagert werden und hauptsächlich Monoterpene (z.B. Menthol, Thymol, Pulegol) oder seltener Sesquiterpene beinhalten (Hegnauer, 1989). Desweiteren kommen z.T. bittere und biologisch vielseitig aktive Diterpene, Triterpene, Iridoide, Kaffeesäurederivate (teilweise mit Bitterstoffeigenschaften) und Flavonoide vor. Trotz der Wichtigkeit vieler Arten und ihrer Inhaltsstoffe war die Familie seit Linné (1753) weit von einer natürlichen Einteilung entfernt. Vor allem in der zweiten Hälfte des 18. Jahrhunderts wurden neu endeckte Arten von einzelnen Bearbeitern so unterschiedlich auf diverse Gattungen verteilt, dass Lindley (1829) den taxonomischen Zustand der Labiatae sogar als „disgrace of botany“ bezeichnete. Die erste umfassende Bearbeitung von Bentham (1832-36) leistete einen wesentlichen Beitrag zur Klärung der chaotischen Verhältnisse. Vor allem in seiner später modifizierten Form (Bentham, 1848; 1876) hat dieses Konzept bis heute viel von seiner Bedeutung behalten. 1.1 Abgrenzung und infrafamiliäre Gliederung der Lamiaceae Nicht nur die Unterteilung der Familie sondern auch die Abgrenzung zu den nahestehenden Verbenaceae war lange Zeit unklar. Bis vor ca. 25 Jahren wurden die Lamiaceae als Gruppe verstanden, die durch einen gynobasisch inserierten Griffel charakterisiert ist. Bereits Junell (1934) hatte jedoch darauf hingewiesen, dass die Familie so umgrenzt keine natürliche Einheit darstelle und eher alle Vertreter mit laminarer Plazentation der Samenanlagen und cymösen Infloreszenzen in den Lamiaceae zusammengefasst werden sollten; in die Verbenaceae stellte er alle Taxa mit marginaler Plazentation und racemösen Infloreszenzen. Sein Konzept wurde allerdings wenig beachtet und erhielt erst durch die morphologisch-kladistischen Untersuchungen von Cantino (1992a, b) und verschiedene molekulare Untersuchungen (Wagstaff & al., 1995; Wagstaff & Olmstead, 1997; Wagstaff & al., 1998, s. Abb. 1; Lindqvist & Albert, 2002) neue Unterstützung. Mit der Veröffentlichung von Harley & al. (2004) kann es
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