Molecular evolution, phylogenetics and biogeography in southern hemispheric bryophytes with special focus on Chilean taxa. Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Rolf Blöcher aus Biedenkopf/Lahn Bonn 2004 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn 1. Referent: Prof. Dr. Jan-Peter Frahm 2. Referent: Prof. Dr. Wilhelm Barthlott Tag der Promotion: 20. Dezember 2004 Diese Dissertation ist auf dem Hochschulschriftenserver der ULB Bonn http://hss.ulb.uni-bonn.de/diss_online elektronisch publiziert meinen Eltern, Doris und Horst Blöcher Contents 1 Introduction...........................................................................................................1 2 A comparison of the moss floras of Chile and New Zealand. ...............................9 2.1 Introduction ...................................................................................................9 2.2 Comparison.................................................................................................10 2.3 Results ........................................................................................................11 2.4 Discussion...................................................................................................12 3 A preliminary study on the phylogeny and molecular evolution of the Ptychomniaceae M. Fleisch. (Bryopsida) with special emphasis on Ptychomnion ptychocarpon and Dichelodontium. ...........................................................................21 3.1 Introduction .................................................................................................21 3.2 Material & Methods .....................................................................................25 3.3 Results ........................................................................................................30 3.3.1 Sequence Variation..............................................................................30 3.3.2 Genetic distances.................................................................................32 3.3.3 Phylogenetic analysis...........................................................................33 3.4 Discussion...................................................................................................37 4 The systematic affinities of selected Gondwanan bryophyte taxa based on molecular sequence data ..........................................................................................42 4.1 Introduction .................................................................................................42 4.2 Material and Methods..................................................................................45 4.3 Results ........................................................................................................49 4.3.1 Sequence variation ..............................................................................49 4.3.2 Phylogenetic analysis...........................................................................49 4.3.3 Synthesis. ............................................................................................53 4.4 Discussion...................................................................................................54 5 Molecular evolution, phylogenetics and biogeography of the genus Lepyrodon (Lepyrodontaceae, Bryopsida) ..................................................................................58 5.1 Introduction .................................................................................................58 5.1.1 The genus Lepyrodon ..........................................................................58 5.1.2 Morphological relationships within the genus.......................................58 5.1.3 The systematic position of Lepyrodontaceae .......................................59 5.2 Material & Methods .....................................................................................60 5.3 Results ........................................................................................................67 5.3.1 Sequence variation ..............................................................................67 Contents 5.3.2 Phylogenetic analysis...........................................................................70 5.4 Discussion...................................................................................................79 5.4.1 Genetic results .....................................................................................79 5.4.2 Phylogenetic and taxonomic results.....................................................82 5.4.3 Biogeographical implications................................................................86 6 Molecular circumscription and biogeography of the genus Acrocladium (Bryopsida) ................................................................................................................89 6.1 The genus Acrocladium ..............................................................................89 6.1.1 Status of Acrocladium ..........................................................................89 6.1.2 Distribution of Acrocladium ..................................................................90 6.1.3 Ecology of Acrocladium........................................................................90 6.2 Material & Methods .....................................................................................91 6.3 Results ........................................................................................................98 6.3.1 Sequence variation ..............................................................................98 6.3.2 Genetic distances...............................................................................101 6.3.3 Phylogenetic analysis.........................................................................104 6.4 Discussion.................................................................................................106 6.4.1 The status of A. auriculatum and A. chlamydophyllum.......................106 6.4.2 Possible explanations for the disjunct distribution of Acrocladium .....108 7 Molecular evolution, phylogenetics and biogeography of the genus Catagonium (Plagiotheciaceae, Bryopsida).................................................................................111 7.1 Introduction ...............................................................................................111 7.1.1 Morphological characterisation ..........................................................112 7.2 Material & Methods ...................................................................................115 7.3 Results ......................................................................................................122 7.3.1 Phylogenetic results...........................................................................122 7.3.2 Indel matrix ........................................................................................130 7.3.3 Genetic distances...............................................................................131 7.4 Discussion.................................................................................................136 7.4.1 The ‘Northern South American’ species.............................................136 7.4.2 The systematic position of C. nitens ssp. maritimum .........................137 7.4.3 The relationship within Catagonium nitens.........................................137 8 The 'Gondwana connection' and their genetic patterns in bryophytes..............140 9 Summary..........................................................................................................146 Contents 10 Acknowledgements.......................................................................................149 11 References ...................................................................................................152 Index to tables Index to figures Appendix 1 Introduction 1 1 Introduction Biologists have long been fascinated by the existence of disjunct distributions of certain plant and animal taxa. Especially the southern temperate disjunctions between southern South America and New Zealand have attracted their attention. The taxa characterized by these distribution patterns are assumed to share a common history. Generally two different hypotheses are used to explain their disjunct distribution. The first can be described by the term ‘vicariance’ which refers to disjunct distribution patterns as a result of the splitting of populations by e.g. the fragmentation of landmasses (e.g. Croizat et al., 1974). The second hypothesis explains the existing distribution patterns based on long distance dispersal events. For the first explanation based on vicariance events an understanding of the past fragmentation processes of the continental landmasses is necessary. The former connection of the recent southern continents in a large landmass, the Gondwana continent, is nowadays widely accepted. Over a period of c. 180 Myr mainly continental drift led to the recent formation of the continents (e.g. McLoughlin, 2001). During the Permian to Jurassic period the supercontinent Pangea consisted of a northern land mass, Laurasia and a southern land mass Gondwana, that were partly separated by an ocean, the Tethys. During that time Pangea extended from