Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 333 Disentangling Lecania RIKKE REESE NÆSBORG ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 UPPSALA ISBN 978-91-554-6953-5 2007 urn:nbn:se:uu:diva-8183 Dissertation presented at Uppsala University to be publicly examined in Lindahlsalen, EBC, Norbyvägen 18A, Uppsala, Friday, September 28, 2007 at 09:00 for the degree of Doctor of Philosophy. The examination will be conducted in English. Abstract Reese Næsborg, R. 2007. Disentangling Lecania. Acta Universitatis Upsaliensis. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 333. 42 pp. Uppsala. ISBN 978-91-554-6953-5. This thesis focuses on phylogenetic, taxonomic, ecological, and conservation aspects of the crustose lichen genus Lecania (Ramalinaceae, lichenized Ascomycota). Lecania has previously been defined on basis of relatively few morphological characters, and the genus had never been treated in molecular phylogenies. The molecular phylogeny of the genus is inferred from DNA sequences. Twenty-five species traditionally placed in Lecania are included in the study along with 21 species from closely related genera. Lecania is a polyphyletic genus. A well-supported monophyletic group containing 16 Lecania species, including the type species L. fuscella is discovered, i.e. Lecania s. str. Nine species formerly included in Lecania do not belong in the genus. A new species, L. belgica, is described. The relationships of a group of morphologically similar Lecania species, i.e. the L. cyrtella group are investigated using morphological and molecular methods. Haplotype network and phylogenetic analyses indicate that the included species, as conceived in the morphological examinations, all are monophyletic. Two new species, L. leprosa and L. madida,are described, L. proteiformis is resurrected from synonymy, and the known range of L. prasinoides is greatly expanded. The type species Lecania fuscella has become endangered in many countries. Twelve localities in Sweden where the species had been found historically are investigated, but L. fuscella is only recovered in one locality. The species composition in these 12 localities, 58 old and 5 new collections with L. fuscella is determined and analyzed. The vegetation community differs between the old and the new collections, and between the locality where the species is recovered and those where it is not. Lecania fuscella has not been able to adapt to environmental changes and now only appears in a specific type of vegetation community. The phylogenetic diversity of the species is calculated, but does not reflect the species’ evolutionary potential. Keywords: Lecania, Ramalinaceae, Crustose lichens, Phylogeny, Taxonomy, Morphology, New species, Phylogenetic diversity, Rare species, ITS, mt-SSU, RPB2, IGS Rikke Reese Næsborg, Department of Evolution, Genomics and Systematics, Systematic Botany, Norbyv. 18D, Uppsala University, SE-75236 Uppsala, Sweden © Rikke Reese Næsborg 2007 ISSN 1651-6214 ISBN 978-91-554-6953-5 urn:nbn:se:uu:diva-8183 (http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-8183) Distributor: Uppsala University Library, Box 510, SE-751 20 Uppsala www.uu.se, [email protected] Following our will and wind we may just go where no one’s been. We’ll ride the spiral to the end and may just go where no one’s been. Spiral out. Keep going. Maynard Keenan Cover: Lecania fuscella, asci with spores and paraphyses. Illustration by the author. Papers included in this thesis This thesis is based on the following papers, which are referred to in the text by their Roman numerals: I Reese Næsborg, R., Ekman, S., & Tibell, L. 2007. Molecular phylogeny of the genus Lecania (Ramalinaceae, lichenized As- comycota). Mycological Research 111: 581–591. II Reese Næsborg, R. & van den Boom, P. P. G. Lecania belgica van den Boom & Reese Næsborg, a new saxicolous species from Western Europe. Submitted to Lichenologist. III Reese Næsborg, R. Taxonomic revision of the Lecania cyrtella group (Ramalinaceae, lichenized Ascomycota) based on molecu- lar and morphological evidence. Submitted to Mycologia. IV Reese Næsborg, R. The phylogenetic diversity of Lecania fus- cella does not reflect its evolutionary potential. Manuscript. Paper I is reprinted with kind permission of the publisher. All papers in this thesis were written by the first author, with comments and suggestions given by the co-authors. In paper I RRN is responsible for most of the laboratory work and two of the phylogenetic analyses. Important note. In papers II and III the manuscripts contain descriptions of new species. These papers have been submitted for publication elsewhere, and in order to make it clear that the names of these new species are not validly published in this thesis the Latin diagnoses, which are necessary according to the International Code of Botanical Nomenclature, are omitted. Contents Introduction.....................................................................................................9 Lichens .....................................................................................................10 Ramalinaceae ...........................................................................................10 Lecania.....................................................................................................11 Aims .........................................................................................................15 Materials and methods ..................................................................................16 Results and discussion ..................................................................................17 Paper I and II............................................................................................17 Paper III....................................................................................................19 Paper IV ...................................................................................................22 Dansk resumé................................................................................................26 Laver ........................................................................................................26 Lavsystematik...........................................................................................27 Lecania og min forskning ........................................................................28 Svensk sammanfattning ................................................................................32 Lavar ........................................................................................................32 Lavsystematik...........................................................................................33 Lecania och min forskning.......................................................................34 Acknowledgements.......................................................................................37 References.....................................................................................................40 Abbreviations bp. base pairs BP-ML Bootstrap Proportion Maximum Likelihood BP-MP Bootstrap Proportion Maximum Par- simony DD Data Deficient IGS InterGenic Spacer region ITS Internal Transcribed Spacer region LSU Large SubUnit mt-SSU mitochondrial Small SubUnit NMS Non-metric Multidimensional Scal- ing PD Phylogenetic Diversity PP Posterior Probability RPB2 RNA polymerase II second largest subunit s. lat. sensu lato (in a broad sense) s. str. sensu stricto (in a narrow sense) SSU Small SubUnit Introduction Systematics is a field of science which addresses the diversity of life and the evolutionary relationships among living organisms. In order to understand these relationships several tools or disciplines are employed, i.e. identifica- tion, taxonomy, and phylogenetics. First, the organisms must be identified and described, since this is the foundation for deducing relationships. When organisms are properly identified and described, they can be named (taxon- omy) and arranged in groups according to their evolutionary relatedness (phylogenetics). Sometimes, however, inference of a molecular phylogeny precedes description and naming of taxa. The evolutionary relatedness can be reconstructed using both morphological and molecular characters, the latter usually in the form of DNA sequences. Phylogenetic reconstructions are often elucidated by depicturing trees where monophyletic groups share the same common ancestor and, to some extent, the same derived characters. Systematics is therefore a fundamental discipline on which many other fields of biology rely. An area where systematics has both indirect and direct influence and im- pact is conservation biology. Indirectly, since biological diversity could not be assessed without proper identification, description, and naming of taxa. Recently, systematics has also offered more direct contributions to the field of conservation biology. Owing to the limited resources available for assign- ing protection of endangered species and/or habitats, it is often necessary to prioritize or rank the species/habitats requiring protection. Systematics offers objective means of setting these priorities by evaluating the biological dis- tinctiveness of the species per se or of the area for one/more groups of or- ganisms. The biological distinctiveness is reflected in genetically distinct lineages, and one method to estimate these genetically distinct lineages is the measure