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Dissertationskonzept Michael Miller TECHNISCHE UNIVERSITÄT MÜNCHEN Lehrstuhl für Tierökologie DNA-Taxonomie in der Entomologie Eine Fallstudie der DNA-basierten Systematik bei Insekten Michael Andreas Miller Vollständiger Abdruck der von der Fakultät Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt der Technischen Universität München zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften genehmigten Dissertation. Vorsitzender: Univ.-Prof. Dr. Reinhard Schopf Prüfer der Dissertation: 1. Univ.-Prof. Dr. W. Back (i. R.) 2. Univ.-Prof. Dr. D. Metzler, Ludwig-Maximilians-Universität München 3. Univ.-Prof. Dr. J. P. Geist Die Dissertation wurde am 04.05.2011 bei der Technischen Universität München eingereicht und durch die Fakultät Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt am 12.07.2012 angenommen. F:\Diss1\FinalVersion\Titelblatt Promotion-Miller-2012.doc 12.10.2012 Meinen Eltern und meinen Söhnen Sebastian, Korbinian und Johannes und Ulla. DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller Triumphe halten keine Lehren bereit, Mißerfolge dagegen befördern die Erkenntnis auf mannigfaltige Art Hans Magnus Enzensberger - 1 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller - 2 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller DNA-Taxonomie in der Entomologie Eine Fallstudie der DNA-basierten Systematik bei Insekten Michael A. Miller Freising - 3 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller - 4 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller Danksagung Mein großer Dank gebührt Prof. Dr. Werner Back, der die Betreuung der Arbeit übernommen hat, und Prof. Dr. Roland Gerstmeier für die wissenschaftliche Unterstützung und Aufnahme in seiner Arbeitsgruppe am Lehrstuhl für Tierökologie des Wissenschaftszentrums Weihenstephan. Besonders danken möchte ich Dipl.-Biol. Ulla Oliveira Biener, Dipl.-Biol. Stefan Friedrich und Prof. Dr. Roland R. Melzer, die mich freundschaftlich und ideell anleiteten und ermutigten. Herzlich gedankt sei an dieser Stelle allen Mitautoren. Es war eine anspruchsvolle Herausforderung, mit Euch zusammenzuarbeiten, ich konnte viel von Euch lernen und es hat mir Spaß gemacht, mich in die jeweiligen Taxa und Fragestellungen einzuarbeiten. Namentlich erwähnen möchte ich (alphabetisch sortiert): Werner Back, Michael Balke, Ulf Buchsbaum, Verena Etscher, Günter Müller, Axel Hausmann, Axel Hille, Thomas Knebelsberger, Torsten Laamanen, Rudolf Meier, Günter Müller, Paul Opler, Ignacio Ribera, Wolfgang Speidel, Wolfgang ten Hagen und Alfried Vogler. Dankend erwähnen möchte ich nochmals Prof. Dr. Diethard Tautz (MPI f. Evolutionsforschung, Plön), der mir die DNA-Analyse beibrachte und die DNA- Taxonomie erfand, sowie die Leute aus seinem damaligen Labor des Zoologischen Instituts der LMU in München, die das Tagesgeschäft der Lehre übernahmen, besonders (alphabetisch sortiert): Andreas Berghammer, Christoph Eckert, Claudia Englbrecht, Melanie Markmann, Nico Prpic, Kornelia Rassmann, Sebastian Steinfartz, Barbara Wimmer. Erst im Lauf der Karriere wird einem die Bedeutung der Schule, aus der man kommt, bewusst. - 5 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller - 6 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller Inhaltsverzeichnis Abkürzungsverzeichnis ............................................................................................... 8 Abbildungsverzeichnis ................................................................................................ 8 1. Einleitung und Problemstellung ............................................................................ 10 1.1 Neue Herausforderungen ................................................................................ 10 1.2 Lösungsansätze .............................................................................................. 11 2. Theoretische und methodologische Grundlagen .................................................. 12 2.1 Die Historie der Molekularen Analysen ........................................................... 12 2.2 Die DNA-Taxonomie ....................................................................................... 13 2.3 Der Artbegriff ................................................................................................... 16 2.4 Vorstellung der untersuchten Themenkomplexe ............................................. 17 3. Material und Methoden ......................................................................................... 25 3.1 DNA-Extraktion ............................................................................................... 25 3.2 Analyseverfahren ............................................................................................ 26 3.3 Neubeschreibungen ........................................................................................ 28 3.4 Artabgrenzungen/Phylogenie .......................................................................... 30 3.5 Identifikationen ................................................................................................ 33 3.6 Ökologische Anwendungen ............................................................................ 35 4. Ergebnisse und Diskussion .................................................................................. 36 4.1 DNA-Extraktion ............................................................................................... 36 4.2 Analyseverfahren ............................................................................................ 37 4.3 Neubeschreibungen ........................................................................................ 42 4.4 Artabgrenzungen/Phylogenie .......................................................................... 47 4.5 Identifikationen ................................................................................................ 64 4.6 Ökologische Anwendungen ............................................................................ 69 5. Zusammenfassung ............................................................................................... 74 6. Literaturverzeichnis............................................................................................... 77 7. Weitere eigene Veröffentlichungen ..................................................................... 104 - 7 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller Abkürzungsverzeichnis bp Basenpaar(e) CI Consistency index COI Cytochromoxidase Untereinheit I ILD Inkongruenz-Längendifferenz ML Maximum Likelihood MP Maximum Parsimony mtDNA mitochondriale Desoxribonukleinsäure nDNA nukleäre Desoxribonukleinsäure NJ Neighbor Joining PBS Partitionierter Bremer-Support PP Posterior Probability RI Retention index RILD bewertete Inkongruenz-Längendifferenz ti Transition tv Transversion Abbildungsverzeichnis Abbildung 1: Abhängigkeit des PCR-Erfolgs vor der Probenhistorie............................................. 37 Abbildung 2: Phylogenie der Gattung Themira .............................................................................. 41 Abbildung 3: Holotypen der neubeschriebenen Leucoblepsis taiwanensis sp. nov. ..................... 42 Abbildung 4: Tabelle der variablen Basenpositionen bei Leucoblepsis taiwanensis sp.nov. ........ 42 Abbildung 5: Tabelle der variablen Basenpositionen bei Paracymoriza naumanniella sp. nov. ... 43 Abbildung 6: Typen von Paracymoriza naumanniella sp. nov.. ..................................................... 44 Abbildung 7: Idaea nigra Hausmann & Bläsius sp. nov. ................................................................ 45 Abbildung 8: Holotypen der neubeschriebenen Colias-Arten ........................................................ 46 Abbildung 9: Netzwerkanalyse der Unterarten von Anthocharis cardamines ............................... 48 Abbildung 10: Phylogenetische Verwandtschaftsverhältnisse der Untergattung Elphinstonia ..... 50 Abbildung 11: Distanzanalyse-Baum der Gattungen Euchloe und Iberochloe gen. nov............... 52 Abbildung 12: Boxplots der genetischen Distanzen in der Artengruppe II von Euchloe ............... 56 Abbildung 13: Boxplots der genetischen Distanzen in der Artengruppe V von Euchloe ............... 57 Abbildung 14: NJ-Baum der Gattung Callophrys nebst Außengruppenvertretern ........................ 59 Abbildung 15: Network-Analyse der untersuchten Callophrys und Außengruppen ...................... 60 Abbildung 16: Phylogenie der Gattung Rhantus ............................................................................ 62 Abbildung 17: NJ – Analyse der genetischen Distanzen von D. tutti und D. chrysitis ................... 65 Abbildung 18: Tabelle der variablen Sequenzpositionen bei Polythore spaeteri. ......................... 66 Abbildung 19: Strikter Konsensusbaum der Phyllodromica subaptera-Artengruppe .................... 68 Abbildung 20: Distanzbaum (UPGMA) der untersuchten Raupen und der Referenztaxa ............ 70 Abbildung 21: Abhängigkeit der Detektionserfolges von der Ingestionsdauer. ............................. 72 - 8 - DNA-Taxonomie in der Entomologie Dissertation Michael A. Miller Hinweis: Aus dieser Dissertation wurden bereits einige Teilaspekte publiziert. Die einzelnen Fragestellungen der vorliegenden Arbeit werden deshalb im Folgenden teilweise etwas kursorisch dargestellt und übergreifend diskutiert. Für eine detaillierte und umfangreiche Darstellung der Literaturdiskussion, des Material- und Methodenteils sowie der Ergebnisse
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