Phylogenie, Evolution und Biogeographie der GattungenSempervivum und Jovibarba
Dissertation Zur Erlangung des Grades Doktor der Naturwissenschaften
Am Fachbereich Biologie Der Johannes Gutenberg-Universität Mainz
Johannes Thomas Klein geboren am 07.10.1983 in Bruchsal
Mainz，Januar 2015 Inhaltsverzeichnis Allgemeine Einleitung...... 1
1. Artikel:，，Phylogeny, Biogeography，and Evolution of Edaphic Association in the European OreophytesSempervivum andJovibarba (Crassulaceae)55...... 5
،ﺀﻷ^،ةﻷ^٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠_٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠٠أ Introduction...... 5 Materials and Methods...... 6 Taxon Sampling...... 6 DNA Extraction，Amplification，and Sequencing...... 6 Sequence Alignment and Phylogenetic Analysis...... 7 Molecular Dating...... 7 Ancestral Edaphic Association Reconstruction...... 8 Ancestral Area Reconstruction...... 8 و ...... Results Variability of Markers and Exclusion of ...... Taxa 9 Molecular PhylogenySempervivum of andJovibarba ...... 9 Molecular Dating Estimation...... 13 Reconstruction of Edaphic Association...... 16 Ancestral Arca Reconstruction...... 16 Discussion...... 16 Phylogeny and BiogeographySempervivum of andJovibarba ...... 16 Evolution of Edaphic Association...... 21 Acknowledgments...... 22 Appendix A ...... 23 Accessions Sequenced for This Study...... 23 Sequences Retrieved from GenBank...... 27 Literature Cited...... 29
2. Artikel: “AHopatric hybrids as evidence for past range dynamicsSempervivum in andJovibarba^ ...... 33 Introduction...... 33 Materials and methods...... 35 Taxon sampling...... 35 DNA extraction，amplification and sequencing...... 36 Cut-gel separation of PCR fragments...... 36 Assessment of parental lineages...... 37 Results...... 38 Jovibarba globifera ssp. hirta...... 38 Sempervivum arachnoideum...... 40 Sempervivum cantabricum...... 42 Sempervivum xfunckii...... 43 Sempervivum macedonicum...... 44 Sempervivum marmoreum...... 46 Sempervivum thompsonianum...... 50 Parental lineages in sw Asia — phylogeny and nuclear sequence divergence.············ 51 Sempervivum altum...... 54 Sempervivum annae...... 54 Sempervivum armenum...... 56 Sempervivum artvinense...... 56 Sempervivum atropatanum...... 57 Sempervivum borissovae ands. ingwersenii...... 58 Sempervivum davisii...... 60 Sempervivum glabrifolium...... 61 Sempervivum ossetiense...... 64 Sempervivum pisidicum...... 65
Sempervivum〇 ร ท 〇พ ร ¥...... 68 Sempervivum transcaucasicum...... 69 Discussion...... 73 Allopatric hybrids as evidence for past migration and past ranges...... 74 Ecological novelty...... 82 Acknowledgments...... 83 Literature Cited...... 151 Appendix A l...... 162
3. Artikel: “Evidence for homoploid and polyploid hybrid speciation involving the same parent species(Sempervivum tectorum X s. calcareum^ Crassulaceae) and implications for the geography of hybrid speciation”...... 167 Abstract...... 167 Introduction...... 168 Materials and methods...... 172 Taxon sampling...... 172 DNA extraction, amplificationand sequencing...... 172 Cloning of PCR fragments...... 173 Sequence alignment and phylogenetic analysis...... 173 Chromosome numbers...... 174 Results...... 175 Sequence comparison...... 175 Phylogenetic analysis of sequences...... 176 Chromosome numbers...... 177 Discussion...... 180 Sempervivum plants in the Rhine Gorge and the Massif Central are homoploid and polyploid hybrid species，respectively，originating from the same pair of parental species...... 180 The geographical setting of hybrid speciationSempervivum in and the role of Quaternary climatic oscillations for hybrid species formation...... 183 Acknowledgments...... 185 References...... 185 Biosketches...... 190 Author contributions...... 190 Supporting information...... 193
Zusammenfassung...... 196 Literatur zur Einleitung...... 198 Danksagung...... 201 Lebenslauf...... 202