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Reticulate Evolution in Diphasiastrum (Lycopodiaceae)

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Reticulate Evolution in Diphasiastrum (Lycopodiaceae) !" # # $%$&$' ( )*+&)$&%%'&*'*,& -./.0. &))%+' !"!##"#$##% % %& '( ) * ' +.,' D.!##"'-* . /' ' 0!1'12 ' '"34"42254353#40' 6 % ' ) 7 ' + % 8 9 ' : 7 % !"#$ !#%& ) % % %% % 9 ' % + % ( % , % ; ' ( ) % ) %% ' ' % ) % ' ( % 8 * % ' () % ) % ' % % : < '( % < %7 % 8 '- ) '= % 7 8 % % )) % 8 ' )( * > : 8 % # + " + , + + "-./ 01 + ( ?+ , !##" 6++0240!5 6+>"34"42254353#40 $ $$$ 4""25. $@@ '<'@ A B $ $$$ 4""25/ In memory of Morten T. Cover: Diphasiastrum alpinum from Northern Norway. Photo: B. Pettersen. List of papers This thesis is based on the following papers, which are referred to by their Roman numerals I Aagaard, S.M.D., Vogel, J.C. & N. Wikström. Resolving maternal relationships in the clubmoss genus Diphasiastrum (Lycopodiaceae). In press. Taxon. II Aagaard, S.M.D., Greilhuber, J., Zhang, X.-C. & N. Wikström. Oc- currence and evolutionary origins of polyploids in the club moss ge- nus Diphasiastrum (Lycopodiaceae). In press. Molecular Phyloge- netics and Evolution. III Aagaard, S.M.D., Greilhuber, J., Vogel, J.C. & N. Wikström. Re- ticulate phylogenetic patterns in diploid European Diphasiastrum (Lycopodiaceae). Manuscript IV Aagaard, S.M.D., Gyllenstrand, N. & N. Wikström. Homoploid hybridization in Central European Diphasiastrum (Lycopodiaceae). Manuscript. V Thulin, M., Aagaard, S.M.D., Wikström, N. & C. Jarvis. Revised lectotypification of Lycopodium complanatum L. (Lycopodiaceae). In press. Taxon. Papers I, II and V are reproduced by kind permission of the publishers. All papers included in the thesis are written by the first author, with addi- tional comments and elaborations of technical procedures provided by the co-authors. The laboratory work and analyses were conducted by S. M. D. Aagaard with the exception of the Feulgen DNA image densitometry analy- ses in paper II and III, which were conducted by Professor Johann Greil- huber. Contents Introduction.....................................................................................................9 Polyploidy ................................................................................................11 Homoploid hybridization and speciation .................................................12 Aims .........................................................................................................15 Materials & Methods ....................................................................................16 Taxa included in the phylogenetic analyses .............................................16 Investigated populations...........................................................................17 Chloroplast sequence data (Paper I).........................................................18 Chloroplast microsatellites (Paper I – IV)................................................18 Flow densitometry data (Paper II and Paper III)......................................18 Nuclear single and low copy regions (Paper II – Paper IV).....................19 Patterns expected in polyploids (Paper II)...........................................19 Patterns expected in homoploid hybrids (Paper III and Paper IV) ......19 Results & Discussion ....................................................................................20 Intrageneric relationships and diversity (Paper I and Paper V)................20 Polyploidy in Diphasiastrum (Paper II) ...................................................22 Homoploid hybridization in Diphasiastrum (Paper III and Paper IV).....24 Phylogenetic analyses..........................................................................24 Admixture analyses .............................................................................26 Concluding remarks .................................................................................28 Svensk sammanfattning (Swedish summary) ...............................................30 Acknowledgments.........................................................................................32 References.....................................................................................................33 Introduction “The taxonomist whose species concept is the result of experience princi- pally with plant taxa in which characters are more gross and differences be- tween species are of a more qualitative nature is likely to be unimpressed by the subtle and quantitative differences he finds between L. complanatum and its relatives. He may feel that no matter how constant they are (and many of the individual characters do overlap), such differences hardly constitute sub- specific, let alone specific characters.” Joan H. Wilce (1961) A hybrid is the product of a cross between two distinct species. Individuals of hybrid origin are assumed to be sterile, as the parental genomes in most cases are incompatible. But sometimes the merger of genomes between closely related species can give rise to a new distinct, evolutionary lineage. Species of hybrid origin are initially divided into two categories; those having the same number of chromosomes as their parent taxa, i.e. homoploid hybrids, and those established through chromosome doubling, i.e. allopoly- ploids (Mallet, 2007; Rieseberg and Willis, 2007). Both processes are now acknowledged to enable quick adaptation, and consequently ecological spe- cialisation (Lexer et al., 2003; Jiggins et al., 2008). Reticulate evolutionary events also constitute a major challenge in evolutionary as well as ecological studies of the club moss genus Diphasiastrum (Holub). The genus Lycopodium L. comprises about 40-50 species attributed to nine sections (Øllgaard, 1987). With 20-30 species Diphasiastrum, or Lyco- podium section Complanata Victorin, is by far the largest group. Most spe- cies in Diphasiastrum are widely distributed across the northern hemisphere (Wilce, 1965), but some are reported from tropical highlands with temperate climate. A number of endemics have also been described from islands in both the Atlantic and the Pacific Ocean (Wilce, 1961; Wilce, 1965). The most unique feature contributing to the species count in Diphasiastrum is that at least five diploid hybrid species are commonly recognized in the ge- nus (Figure 1; Wilce, 1965; Holub, 1975; Stoor et al., 1996). Polyploids are less frequently reported, but at least two polyploid species have been consid- ered in the literature (Wilce, 1961, Wilce, 1965). Clubmosses are characterized by having two independent generations; diploid sporophytes, producing wind born spores with a potentially unlim- ited dispersal range (Page, 2002), and haploid gametophytes. As the gameto- phytes are bisexual, three different breeding systems might be observed; 9 outbreeding, intergametophytic selfing or sib-mating, and intragametophytic selfing (Soltis and Soltis, 1988). Intragametophytic selfing, where two gam- etes from the same bisexual gametophyte fuse and produce a completely homozygous individual, could potentially aid the establishment and genetic ‘fixation’ of a new species of hybrid origin (Rieseberg and Willis, 2007). A number of factors make it problematic to investigate hybridization in Diphasiastrum. They have a simple morphology with few discrete morpho- logical features that can be evaluated (Wilce, 1961; Wilce, 1965, Vogel and Rumsey 1999). Also, little is known about their reproductive biology. The mycorrhizal dependence of the gametophyte makes their spores difficult to germinate in controlled laboratory environments (Whittier, 1977), and cross- ing experiments are virtually impossible to accomplish. Counting chromosomes in Diphasiastrum is difficult, but in most cases a base chromosome number n=23-24 has been reported (Wagner, 1992). Even though diploid homoploid hybrid species are assumed to predominate in Diphasiastrum, two taxa, D. wightianum (Grev. et Hook.) and D. zanclo- phyllum (J. H. Wilce) Holub, have been recognized to be of polyploid hybrid origin (2n=4x=96). Diphasiastrum wightianum has been confirmed to be tetraploid by chromosome counts (Ninan, 1958). Diphasiastrum zanclophyl- lum has been considered to be a polyploid based on the observation that its spores are significantly larger than in known diploids (Wilce, 1961; Wilce 1965). Observing the morphological diversity among accessions identified as D. wightianum and D. zanclophyllum, Wilce (1965) discussed the possibility that the number of polyploids in Diphasiastrum has been underestimated. Nonetheless, reports on ploidal level of accessions identified to be putative homoploid hybrid species confirm chromosome numbers equal to that of the suggested parent taxa (Wagner, 1992). 10 Polyploidy Polyploidization is a relatively common mode of sympatric speciation among vascular plants, and is considered more frequent in homosporous ferns and lycophytes than other plant groups (Otto and Whitton, 2000). The occurrence of polyploids has been well accounted for in two other genera in Lycopodiaceae, Huperzia Bernh. and Lycopodiella Holub (Wagner et al., 1985), but reports from Diphasiastrum and Lycopodium sens. str. are less frequent (Wagner, 1992). Depending on the mode of origin and degree of divergence between
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