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Phylogeny and Taxonomy of Polyblastia and Allied Taxa (Verrucariaceae)

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Phylogeny and Taxonomy of Polyblastia and Allied Taxa (Verrucariaceae) Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 370 Phylogeny and Taxonomy of Polyblastia and allied taxa (Verrucariaceae) ´ SANJA SAVIC ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 UPPSALA ISBN 978-91-554-7035-7 2007 urn:nbn:se:uu:diva-8326 ! " # $%%& %'%% ( ( ( ) * + , * -. -* $%%&* ) + ( /01* ! * 2&%* 22 * * 3- 4& 54566#5&%265&* !* 7* ( 8 ( 0 /! 1* + ( (, ( , * ! 9 ( - ( : ; , 0 , ( * ! < / 3+-1 , , : ;* + * 8 * + ( * 3 (( ( ! =! * " ( ( , * + * - , , (( * ! ( > ! , , * ! < * ! ( ( 0 (( * + , /! # 1 * - ( ( , (( ( * + ( ( ( ( * + , $ /01 , $ $ $ * ! ( < ( * %&' ) 0 - ! + 0 - + ) + 3+- - ?) ( )* + * , * * - ./ * * +012345 * & @ - > -. $%%& 3-- A65A$# 3- 4& 54566#5&%265& ' ''' 5 2$A / '== *<*= B C ' ''' 5 2$A1 List of papers This thesis is based on the following papers, which are referred to in the text by their Roman numerals: I Savi Sanja, Tibell Leif, Gueidan Cécile & Lutzoni François. Molecular Phylogeny and Systematics of Polyblastia (Verrucari- aceae, Eurotiomycetes) and allied genera. Submitted to Myco- logical Research. II Savi Sanja & Tibell Leif. Taxonomy and species delimitation in Sporodictyon (Verrucariaceae) in Northern Europe and the ad- jacent Arctic – reconciling molecular and morphological data. Submitted to Taxon. III Gueidan Cécile, Savi Sanja, Thüs Holger, Roux Claude, Keller Christine, Tibell Leif, Prieto Maria, Heiðmarsson Starri, Breuss Othmar, Orange Alan, Fröberg Lars, Amtoft Wynns Anja, Navarro-Rosinés Pere, Krzewicka Beata, Pykälä Juha, Lutzoni François. The main genera of Verrucariaceae (Ascomycota) as supported by recent morphological and molecular studies. Sub- mitted to Taxon. IV Savi Sanja & Tibell Leif. Atla – a new genus in Verrucaria- ceae. Submitted to The Lichenologist. In Paper I, II and IV SS had the major responsibility for writing the text, with comments and suggestions given by the co-authors. SS was also re- sponsible for most of the lab work, and for the phylogenetic analyses in Pa- per II and IV. The studies (I, II, IV) were planned and conducted in coopera- tion between SS and LT. In Paper III SS was responsible for writing the text of sections: ‘3. Clade E’ (excluding ‘3c’), ‘12. Polyblastia’, ’14. Thelidium group’, the nomenclatural note in section ‘13a. Staurothele’, and took part in writing the discussion. Important note. In all papers new nomenclatural combinations are sug- gested, and in paper II, III and IV new taxa are described. All papers have been submitted for publication elsewhere. Thus in order to make certain that the new names are not validly published in this thesis, references to the basionyms and Latin diagnoses, necessary according to the International Code of Botanical Nomenclature, are omitted. The statement ‘here desig- nated’, needed for lectotypification to be valid, is also omitted. Contents Introduction.....................................................................................................7 Verrucariaceae............................................................................................8 Polyblastia and allied taxa .........................................................................9 Aims..............................................................................................................11 Materials and Methods..................................................................................12 Results and discussion ..................................................................................13 Paper I ......................................................................................................13 Paper II .....................................................................................................15 Paper III....................................................................................................17 Paper IV ...................................................................................................19 Concluding remarks .................................................................................20 Sammanfattning (Swedish summary) ...........................................................22 Att förstå lavar .....................................................................................22 Att namnge lavar..................................................................................23 Vi vet faktiskt inte så mycket… ..........................................................24 Släktskapsförhållanden inom familjen Verrucariaceae .......................25 Revision av Sporodictyon....................................................................26 Ett nytt släkte, Atla ..............................................................................27 Om ett och annat - annat…..................................................................28 Acknowledgements – Tack –Zahvala – Hvala .............................................29 References.....................................................................................................31 Abbreviations ITS Internal Transcribed Spacer region LSU Large SubUnit RPB1 largest subunit of the RNA Polymerase II s.lat. sensu lato (in a broad sense) s.str. sensu strict (in a narrow sense) MP Maximum Parsimony Introduction Lichens are symbiotic associations between fungi (the mycobionts) and a photosynthetic algal/cyanobacterial partner (the photobionts). In general, the symbiosis is considered obligatory for successful growth and reproduction of the heterotrophic fungal partner. However, also the autotrophic photobiont obviously benefits from the relationship. Thus the lichen is typically a highly stable association which extends the ecological range of both partners. The body (thallus) of most lichens is quite different from that of either the fungus or alga growing separately, and may to some extent resemble simple plants in form and growth (Sanders 2001). The 83rd plate from Ernst Haeckel's Kunstformen der Natur (1904), depicting or- ganisms classified as ‘Lichenes’ (inverted colour). Approximately one-fifth of all known fungal species form obligate sym- biotic associations with a green alga, a cyanobacterium or simultaneously 7 with both types of photobionts. Many of the ascomycetes (ca 42%) and a few basidiomycetes participate in lichen symbioses. Ascomycota is a phy- lum that includes more than 98% of the known lichenized fungal species. According to Lutzoni & al. (2001) the lichens in this phylum evolved earlier than previously believed, and gains of lichenization have been infrequent during the evolution of Ascomycota. In contrast multiple independent losses of lichenization have occurred. As a consequence, major Ascomycota line- ages of exclusively non-lichen-forming species are derived from lichen- forming ancestors (including well-known fungi such as Penicillium and As- pergillus). The lichen association is thus believed to be a symbiosis with a long evo- lutionary history. However, the past has left behind clues to what once hap- pened. Systematists use these clues and try to reconstruct evolutionary rela- tionships between organisms. The study of evolutionary relationships among organisms based on molecular, mostly DNA sequence data, have provided new insights about the relationships that often contradict the traditional, pri- marily morphology-based hypotheses. During the past two decades, contributions to fungal classification based on molecular data have led to major changes in our understanding of the evolution of fungi and of their phylogenetic affinities (see Hibbett & al. 2007 for a review). Verrucariaceae The family Verrucariaceae (Verrucariales, Chaetothyriomycetidae, Eurotio- mycetes, Ascomycota) is a large family of mostly lichenized species includ- ing 45 genera and about 750 species (Hawksworth & al. 1995; Eriksson 2006). The family includes lichens producing perithecia as fruiting bodies, and are often referred to as ‘pyrenocarpous lichens’, along with some other orders. The first studies to include molecular data on the family Verrucariaceae aimed at finding the phylogenetic relationships between high-rank taxa within the ascomycetes (Lutzoni & al. 2001, 2004; Liu & Hall 2004; Lumbsch & al. 2002, 2004, 2005; Geiser & al. 2006; Spatafora & al. 2006). They showed that Verrucariales is sister to an order of non-lichenized fungi, the Chaetothyriales. A few molecular studies have focused on the Verrucari- aceae at the infrageneric level (Amtoft 2006; Amtoft & al. 2008; Heiðmars- son 2003, on Dermatocarpon). Most of the genera in Verrucariaceae are crustose; these include, e.g. Polyblastia, Agonimia, Henrica, Staurothele, Thelidium and Verrucaria. The most recent phylogenetic study on Verrucariaceae was published by Gueidan & al. (2007), presenting a multigene phylogeny of 15 genera of Verrucari- aceae, including the most species-rich ones. There the current morphology- 8 based classification was compared with a molecularly based phylogeny. It was shown that spore septation is not useful as a primary morphological feature for recognizing genera, since none of the
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