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Lentibulariaceae) Phylogenetic relationships, systematics, and biology of carnivorous Lamiales, with special focus on the genus Genlisea (Lentibulariaceae) Dissertation zur Erlangung des Doktorsgrades der Naturwissenschaften (Dr. rer. nat.) der Fakultät für Biologie der Ludwig-Maximilians-Universität München vorgelegt von Andreas Fleischmann München, 28.11.2011 Erstgutachter: Prof. Dr. Günther Heubl Zweitgutachter: Prof. Dr. Susanne Renner Tag der mündlichen Prüfung: 29.03.2012 Eidesstattliche Versicherung und Erklärung Eidesstattliche Versicherung Ich, Andreas Fleischmann, versichere hiermit an Eides statt, dass die vorgelegte Dissertation von mir selbständig und ohne unerlaubte Beihilfe angefertigt ist. München, den 28. 11. 2011 ……..……………………….......... Andreas Fleischmann Erklärung Hiermit erkläre ich, Andreas Fleischmann, dass ich mich anderweitig einer Doktorprüfung nicht unterzogen habe, und dass die Dissertation nicht ganz oder in wesentlichen Teilen einer anderen Prüfungskommission vorgelegt worden ist. München, den 28. 11. 2011 ……..……………………….......... Andreas Fleischmann Declaration of contribution as co-author In this cumulative thesis, the results from Andreas Fleischmann’s doctoral research are presented, which was carried out under the supervision of Prof. Günther Heubl at the Ludwig- Maximilians-University of Munich. The following six articles have been published in international peer-reviewed journals, and are presented in the appendix of the thesis. All of them have resulted from collaborations with other scientists, and the author’s contributions to each of them were as follows: Contribution to Article I: Schäferhoff, B., Fleischmann, A., Fischer, E., Albach, D.C., Borsch, T., Heubl, G., Müller, K.F., 2010. Towards resolving Lamiales relationships: insights from rapidly evolving chloroplast sequences. BMC Evolutionary Biology 10: 352–374. A. Fleischmann contributed to the taxon sampling, DNA extraction, and sequence generating, wrote parts of the manuscript and provided photographs for the figures. B. Schäferhoff generated sequence data, made the data analyses and drafted the manuscript. E. Fischer improved the manuscript and contributed to the conception of the study during its initial phase. D.C. Albach provided data, contributed to the conception and improved the manuscript. T. Borsch provided plant material and contributed during manuscript preparation. G. Heubl improved the manuscript in its final phase. K.F. Müller was responsible for the conception of the study and helped writing the manuscript. Contribution to Article II: Fleischmann, A., Schäferhoff, B., Heubl, G., Rivadavia, F., Barthlott, W., Müller, K.F., 2010. Phylogenetics and character evolution in the carnivorous plant genus Genlisea A. St.- Hil. (Lentibulariaceae). Molecular Phylogenetics and Evolution 56: 768–783. A. Fleischmann planned, coordinated and conducted the sampling of plant material on joint field trips to Sierra Leone, South Africa, Zambia, Venezuela and Brazil. He conducted the laboratory work, including DNA extraction, PCR amplification, PCR product purification, subsequent sequence editing and alginment, and phylogenetic analyses. The design and preparation of the figures and diagrams, and the manuscript concept and writing was done by A. Fleischmann. B. Schäferhoff conducted the amplification and sequencing of the trnK intron, and contributed to the data analyses. G. Heubl supervised the laboratory work and phylogenetic analyses, and improved the manuscript. F. Rivadavia provided herbarium specimens for DNA extraction and morphometric analyses, and contributed to the manuscript. W. Barthlott improved the manuscript in its final stage. K.F. Müller supervised the data analysis and corrected the manuscript. Contribution to Article III: Fleischmann, A., Rivadavia, F., Gonella, P.M., Heubl, G., 2011. A revision of Genlisea subgenus Tayloria (Lentibulariaceae). Phytotaxa 33: 1-40. A. Fleischmann wrote the manuscript (including the taxonomic treatments, the Latin diagnoses, and he designed the identification key to the species), and drew all figures, maps, and botanical illustrations. He made the morphometric measurements and analyses based on examination of herbarium specimens in B, K, M, RB, SPF, UEC, and plants studied in situ in Brazil. All but four of the flower photographs illustrating the plants at their natural habitats were taken by A. Fleischmann. He conducted the microscopic seed preparations and SEM images. F. Rivadavia conducted the initial fieldwork, provided herbarium specimens, and contributed to the manuscript. P.M. Gonella helped with measurements of herbarium specimens, contributed to the manuscript, provided photographs, and helped with fieldwork in Brazil. G. Heubl improved the manuscript. Contribution to Article IV: Beck, S.G., Fleischmann, A., Huaylla, H., Müller, K.F., Borsch, T., 2008. Pinguicula chuquisacensis (Lentibulariaceae), a new species from the Bolivian Andes, and first insights on phylogenetic relationships among South American Pinguicula. Willdenowia 38: 201–212. Andreas Fleischmann wrote the entire taxonomic part of the article, including the species description, the Latin diagnosis, and the identification key for the Bolivian species of Pinguicula. He examined the herbarium material and made the morphological analyses for the species description, and he made the botanical line drawing based on the studied type specimens. He wrote major parts of the introduction and discussion parts. Further, he extracted DNA from herbarium specimens or plants from his private living collection for all newly sampled taxa of Pinguicula for this article. S.G. Beck introduced the new species to the knowledge of the other co-authors, and improved the manuscript. H. Huaylla contributed habitat description and observations made on living plants at the locus classicus. K.F. Müller performed the data analysis, and wrote the material and methods part. T. Borsch was responsible for DNA amplification and generated the molecular sequences, and contributed to all parts of the manuscript. Contribution to Article V: Fleischmann, A., Rivadavia, F., 2009. Utricularia rostrata (Lentibulariaceae), a new species from the Chapada Diamantina, Brazil. Kew Bulletin 64: 155–159. A. Fleischmann wrote the manuscript and taxonomical treatment of the new species (including the Latin diagnosis and the identification key), made the morphological measurments and microscope analyses, the SEM images of the seed, and drew the botanical illustration. F. Rivadavia discovered the new species, provided herbarium specimens for morphological analysis, drew the distribution map, and contributed to the manuscript. Contribution to Article VI: Fleischmann, A., Heubl, G., 2009. Overcoming DNA extraction problems from carnivorous plants. Anales del Jardín Botánico de Madrid 66: 209–215. A. Fleischmann developed the concept of the study, provided plant material, made the laboratory work, prepared the figures and wrote the manuscript. G. Heubl supervised the laboratory work and improved the manuscript. All photographs, images, and textual illustrations in this work were made by Andreas Fleischmann. „Die Resultate der eigenen Untersuchung seien im Folgenden geschildert, wobei sich zeigen wird, dass Genlisea eine der merkwürdigsten unter allen tierfangenden Pflanzen ist.“ Karl von Goebel, 1891, Pflanzenbiologische Schilderungen II, p. 121. Contents 1. Introduction ......................................................................................................................... 1 1.1. Carnivorous Plants ..................................................................................................... 1 1.2. Lentibulariaceae ......................................................................................................... 4 1.3. Genlisea ..................................................................................................................... 6 1.3.1. Distribution ....................................................................................................... 10 1.3.2. Habitats ............................................................................................................. 11 1.3.3. History of Genlisea research in Munich ........................................................... 14 2. Discussion ........................................................................................................................... 15 2. 1. Systematics, phylogeny and evolution ...................................................................... 15 2.1.1. Evolution of carnivory in Lamiales: Article I ................................................... 15 2.1.2. Phylogeny and evolution of Genlisea: Article II .............................................. 18 2.1.3. Radiation of the Andean Pinguicula: Article IV ............................................... 20 2.2. Biodiversity and taxonomy ........................................................................................ 22 2.2.1. Infrageneric classification of Genlisea: Article II ............................................. 22 2.2.2. Five new species and a revision of Genlisea subgenus Tayloria: Article III .... 23 2.2.3. Preliminary notes on Genlisea flower biology and pollination ......................... 24 2.2.4. A new Andean species of Pinguicula: Article IV ............................................. 27 2.2.5. A new Brazilian species of Utricularia: Article V ........................................... 28 2.2.6. Checklist of the genus Genlisea .......................................................................
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