Phylogenetic Relationships and Evolutionary History of the Southern Hemisphere Genus Leptinella Cass
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Phylogenetic relationships and evolutionary history of the southern hemisphere genus Leptinella Cass. (Compositae, Anthemideae) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Naturwissenschaftlichen Fakultät III - Biologie und Vorklinische Medizin der Universität Regensburg vorgelegt von Sven Himmelreich aus Regensburg Regensburg, Juli 2009 Promotionsgesuch eingereicht am: 29.07.2009 Die Arbeit wurde angeleitet von: Prof. Dr. Christoph Oberprieler Prüfungsausschuss: Prüfungsausschussvorsitzender: Prof. Dr. Reinhard Wirth 1. Prüfer: Prof. Dr. Christoph Oberprieler 2. Prüfer: Prof. Dr. Günther Rudolf Heubl 3. Prüfer: Prof. Dr. Erhard Strohm Contents I Contents List of Figures II List of Tables III Acknowledgements IV Chapter 1 General Introduction 1 Chapter 2 Phylogeny of southern hemisphere Compositae-Anthemideae based 21 on nrDNA ITS and cpDNA ndhF sequence information Chapter 3 Phylogeny of Leptinella (Anthemideae, Compositae) inferred from 46 sequence information Chapter 4 Phylogenetic relationships in Leptinella (Anthemideae, 78 Compositae) inferred from AFLP fingerprinting Chapter 5 Evolution of dimorphic sex expression and polyploidy in Leptinella 107 Chapter 6 Conclusion 117 Summary 125 Zusammenfassung 128 References 131 Appendices 147 List of Figures II List of Figures Title Leptinella featherstonii on the Chatham Islands with Northern Royal Albatross (photo by P. de Lange, New Zealand). Fig. 1-1 Distribution of Leptinella based on Lloyd (1972c). 8 Fig. 1-2 Variation of plants in Leptinella. 9 Fig. 1-3 Capitula and florets of Leptinella. 9 Fig. 1-4 Leaves from different Leptinella taxa from cultivated plants. 10 Fig. 1-5 Postulate steps of the evolution of breeding systems in Leptinella 16 (modified from Lloyd 1975b). Fig. 2-1 Strict consensus tree of 493.976 equally most parsimonious trees 32 based on cpDNA ndhF sequence information. Fig. 2-2 Phylogenetic tree from a Maximum-Likelihood (ML) analysis 33 based on cpDNA ndhF sequence information. Fig. 2-3 Strict consensus tree of 61 equally most parsimonious trees based 35 on nrDNA ITS sequence information. Fig. 2-4 Phylogenetic tree from a Maximum-Likelihood (ML) analysis 36 based on nrDNA ITS sequence information. Fig. 3-1 Basal part of the majority rule consensus tree inferred from 59 Bayesian analysis of the combined dataset (ITS, psba-trnH, trnC- petN). Fig. 3-2 Apical part of the majority rule consensus tree inferred from 60 Bayesian analysis of the combined dataset (ITS, psba-trnH, trnC- petN). Fig. 3-3 Maximum clade credibility tree from the BEAST analysis. 64 Fig. 3-4 Phylogenetic network based on ITS data of the Leptinella main 67 group. Fig. 3-5 Phylogenetic network based on the combined cpDNA data (psbA- 69 trnH, trnC-petN) of the Leptinella main group. Fig. 4-1 AFLP analysis of all 236 investigated individuals (31 taxa) of 89 Leptinella main clade. Fig. 4-2 AFLP analysis of tetraploid individuals (81, individuals, 15 taxa) 90 of Leptinella main clade. Fig. 4-3 Midpoint rooted neighbour-joining tree using Nei-Li distances of 92 taxon group A. Fig. 4-4 Midpoint rooted neighbour-joining tree using Nei-Li distances of 94 taxon group B. Fig. 4-5 Midpoint rooted neighbour-joining tree using Nei-Li distances of 96 taxon group C. Fig. 4-6 Axes 1 and 2 of the principal coordinate analysis for a) taxa 98 group A, b) taxa group B, and c) taxa group C. Fig. 5-1 Phylogenetic tree from the Bayesian analysis of the combined 115 dataset from chapter 3. The member of each group are shown alphabetically on the right side with their ploidy level and sex expression. Fig. 6-1 Results of the DNA sequencing and AFLP fingerprinting studies of 120 Leptinella (chapters 3 and 4). Fig. 6-2 Long distance dispersal events within the Cotula-group and 123 Leptinella as suggested by molecular phylogenies (chapters 2 and 3). List of Tables III List of Tables Tab. 1-1 Taxa of Leptinella and information to sex expression, chromosome 18 number and distribution. Tab. 1-2 Summary of sex expressions in Leptinella according to Lloyd 19 (1972a,b, 1975a,b, 1980a). Tab. 2-1 Species analysed in this study and their accession data. 25 Tab. 3-1 Species analysed in this study, their accession data and additional 50 information. Tab. 3-2 Comparison of phylogenetic analysis statistics for the various 57 molecular datasets analyzed in this study. Tab. 3-3 Sequence divergence in the ITS dataset. 63 Tab. 3-4 Divergence age estimates (crown age). 63 Tab. 4-1 Samples include in the AFLP analysis. 82 Tab. 5-1 Sex expression and ploidy level of Leptinella taxa. 112 Tab. 5-2 Summary of sex expressions in Leptinella. 113 Acknowledgment IV Acknowledgment First of all I would thank Prof. Dr. Christoph Oberprieler who considerable contributed to the whole concept of this work. During the long time that I spent for my PhD, his many ideas and all the discussions on different topics related to the PhD thesis were always very inspiring. I am also grateful to Prof. Dr. Günther Rudolf Heubl for accepting to be referee of this thesis. I would thank all members of the working group for a very good co-operation. Especially, many thanks to my colleagues Dr. Rosa Maria Lo Presti, Roland Greiner, and Dr. Jörg Meister for helpful discussions on several topics, critical comments and the very nice atmosphere. Thanks also to Peter Hummel for his assistance in the laboratory work. I would like to acknowledge Dr. Kathrin Bylebyl, Susanne Gewolf, PD Dr. Christoph Reisch, and Dr. Christine Römermann for many helpful discussions and all colleagues of the Institute of Botany of the University of Regensburg for a nice working atmosphere. I would like to thank all colleagues in Australia, Bolivia, Chile, and New Zealand for their help in the field trip or for collecting Leptinella material. Especially, many thanks to Dr. Ilse Breitwieser, Frank Rupprecht and Dr. Ines Schönberger for their grateful help during the collecting trip in New Zealand. I would like to acknowledge Mari Källersjö and Pia Eldenäs for their cooperation and the nice time in Stockholm. Thanks a lot to Malte Andrasch, Harald Guldan, Laura Klingseisen, Philipp Kolmar, Michael Saugspier and Andrea Spitzner for their help with the sequencing and cloning of the almost complicate Leptinella during their student training in the lab. Thanks also go to Dr. Maik Bartelheimer, Dr. Burckard Braig, Lena Dietz, Margit Gratzl, and Roland Greiner for their help in putting this work in its current form. Additionally, people who supported different parts of this work are separately mentioned at the end of the respective chapter. Thanks to all of them. Last but not least, special thanks to my parents, who encouraged and supported me whenever it was necessary. Chapter 1 General Introduction 1 Chapter 1 General Introduction Chapter 1 General Introduction 2 Evolution of New Zealand plant groups New Zealand has been isolated by a distance of c. 1500 km from its closet landmass Australia after the break-up of Gondwana 80 million years ago (Cooper and Miller 1993, McLoughlin 2001). After the break-up, New Zealand had undergone several dramatic geologic and climatic events that formed a very diverse topography with a high diversity of biomes (Winkworth et al 2005, Linder 2008). Large parts (or the entire archipelago) of New Zealand were inundated during the Oligocene (Cooper and Millener 1993, Winkworth et al. 2002, Trewick and Morgan-Richards 2005). The uplift of the Southern Alps is dated to c. 12 Ma, but the alpine habitat was established only during the last 5 Ma (Chamberlain and Poage 2000, Winkworth et al. 2005). In the Pleistocene, the glacial cycles and volcanism played an important role in the evolution of the environment of New Zealand (Winkworth et al. 2005). In the past, the biogeography of the southern hemisphere plant groups has received much attention by biologists and the origin of its flora and fauna was extensively discussed. Two contradictory concepts exit about the origin of the southern hemisphere plant groups - vicariance or long distance dispersal (see reviews by Pole 1994, McGlone 2005, Trewick et al. 2007, Goldberg et al. 2008). Recent studies using molecular data suggest that long distance dispersal is more prevalent than vicariance, at least as far as the New Zealand plant and animal lineages are concerned (e.g. Pole 1994, Sanmartin and Ronquist 2004, Winkworth et al. 2005, Sanmartin et al. 2007, Goldberg et al. 2008). Several molecular phylogenies show that the divergence times of many groups are too recent to explain the observed geographic patterns by vicariance (e.g. von Hagen and Kadereit 2001, Swenson et al. 2001, Knapp et al. 2005, Wagstaff et al. 2006, Mitchell et. 2009). However, there is evidence that some New Zealand plant groups originated from before the Gondwana break-up (e.g. Agathis; Stöckler et al. 2002, Knapp et al. 2007). Long distance dispersal events were suggested from New Zealand to Australia, New Guinea, South America, southern Africa, the sub-Antarctic islands, the northern hemisphere, and vise versa (e.g. Winkworth et al. 2005, Sanmartin and Ronquist 2004, Sanmartin et al. 2007, Goldberg et al. 2008, Bergh and Linder 2009). For instance, several proven dispersal events from Australia to New Zealand are thought to be connected to the predominant West Wind Drift and the westerly sea current between these landmasses. Likewise, several cases for long distance dispersal in the reverse direction have been proven as well (reviewed in Winkworth et al. 2002, Sanmartin and Ronquist 2004, Chapter 1 General Introduction 3 Sanmartin et al. 2007, Goldberg et al. 2008). The mechanisms involved in such transoceanic long distance dispersal events are discussed in the recent literature (e.g. Wagstaff et al. 2006, Ford et al. 2007, Goldberg et al. 2008, Bergh and Linder 2009).