Systematics and evolution of the genus Deuterocohnia Mez (Bromeliaceae) Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) Vorgelegt im Fachbereich Naturwissenschaften der Universität Kassel von Dipl.-Biol. Nicole Schütz Kassel, 2011 Vom Fachbereich Naturwissenschaften der Universität Kassel als Dissertation angenommen. Dekan: Prof. Dr. Friedrich W. Herberg Gutachter: Prof. Dr. Kurt Weising und Prof. Dr. Georg Zizka Prüfungskommission: Prof. Dr. Pierre Ibisch, Prof. Dr. Rüdiger Wagner, Prof. Dr. Kurt Weising, Prof. Dr. Georg Zizka Datum der Disputation: 21.02. 2012 Content I Content 1 INTRODUCTION .....................................................................................................1 1.1 Biology and systematics of the Bromeliaceae Juss......................................................1 1.2 The genus Deuterocohnia Mez................................................................................... 5 1.3 Scope of the present study .......................................................................................... 9 2 MATERIAL AND METHODS................................................................................. 11 2.1 Sources of plant material............................................................................................ 11 2.1.1 Living plant material ....................................................................................................... 11 2.1.2 Herbarium specimens..................................................................................................... 12 2.2 Taxonomic treatment.................................................................................................12 2.2.1 Abbreviations and terminology..................................................................................... 12 2.2.2 Assessment of morphological and anatomical characters......................................... 13 2.2.3 Biogeographical and ecological data............................................................................. 14 2.3 Molecular methods.....................................................................................................15 2.3.1 Equipment........................................................................................................................ 15 2.3.2 Chemicals and solutions................................................................................................. 15 2.3.3 DNA isolation................................................................................................................. 17 2.3.4 Gel electrophoresis......................................................................................................... 18 2.3.5 PCR amplification and sequencing of chloroplast DNA fragments ....................... 18 2.3.5.1 Selection and amplification of target fragments......................................... 19 2.3.5.2 Sequencing procedure.................................................................................... 22 2.3.5.3 Data processing............................................................................................... 22 2.3.6 PCR amplification and sequence analysis of nuclear DNA fragments ................... 23 2.3.6.1 Selection and amplification of suitable ncDNA loci ................................. 24 2.3.6.2 DNA sequencing and cloning of PCR products........................................ 25 2.3.6.3 Data processing............................................................................................... 26 2.4 Phylogenetic analyses................................................................................................26 2.4.1 Samples included in the molecular phylogenetic analyses......................................... 26 2.4.2 Alignment......................................................................................................................... 31 2.4.3 Tree and network reconstruction ................................................................................. 31 2.4.3.1 Maximum Parsimony analysis....................................................................... 31 2.4.3.2 Distance analysis............................................................................................. 32 2.4.3.3 Maximum Likelihood analysis....................................................................... 33 2.4.3.4 Bayesian analysis.............................................................................................. 33 2.4.3.5 Networks.......................................................................................................... 34 II Content 3 RESULTS.................................................................................................................. 37 3.1 Taxonomic history.................................................................................................... 37 3.2 Morphology ..............................................................................................................42 3.2.1 Life form, duration and habit.........................................................................................42 3.2.2 Foliage ...............................................................................................................................46 3.2.3 Peduncle and inflorescence ............................................................................................49 3.2.4 Flowers ..............................................................................................................................53 3.2.5 Fruits and seeds................................................................................................................56 3.3 Anatomy ....................................................................................................................58 3.3.1 Leaf ....................................................................................................................................58 3.3.2 Peduncle............................................................................................................................59 3.4 Karyology ..................................................................................................................62 3.5 Distribution and ecology........................................................................................... 63 3.5.1 Distribution, habitat and ecology ..................................................................................63 3.5.2 Phenology and pollination..............................................................................................71 3.5.3 Seed dispersal and germination......................................................................................72 3.5.4 Conservation, use and common names........................................................................73 3.6 Molecular phylogenetic analyses .............................................................................. 75 3.6.1 Statistics of sequence data ..............................................................................................75 3.6.1.1 Intergenic spacer rpl32-trnL............................................................................75 3.6.1.2 Intergenic spacer rps16-trnK...........................................................................75 3.6.1.3 Intergenic spacer trnS-ycf3...............................................................................76 3.6.1.4 Nuclear low copy marker: PRK exon2-5 .....................................................77 3.6.1.5 Nuclear low copy marker: PHYC exon1......................................................78 3.6.2 Phylogenetic reconstructions.........................................................................................80 3.6.2.1 Individual chloroplast loci..............................................................................80 3.6.2.2 Combined chloroplast loci .............................................................................80 3.6.2.3 Individual nuclear loci.....................................................................................95 3.6.2.4 Combined nuclear loci ....................................................................................95 4 DISCUSSION ........................................................................................................... 99 4.1 Methodical aspects.................................................................................................... 99 4.1.1 Herbarium work...............................................................................................................99 4.1.2 Molecular analyses........................................................................................................ 100 4.2 Systematics of Deuterocohnia .................................................................................105 4.2.1 Synonymization of Deuterocohnia and Abromeitiella.................................................... 105 4.2.2 Species classification..................................................................................................... 108 4.2.3 Generic relationships and the monophyletic status of Deuterocohnia ..................... 115 4.2.4 Conclusion on the evolution....................................................................................... 120 Content III 5 TAXONOMIC TREATMENT............................................................................... 129 5.1 Generic description.................................................................................................