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A Phylogenetic Study of the Lecanora Rupicola Group (Lecanoraceae, Ascomycota)

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A Phylogenetic Study of the Lecanora Rupicola Group (Lecanoraceae, Ascomycota) A phylogenetic study of the Lecanora rupicola group (Lecanoraceae, Ascomycota) Grube, M; Baloch, E; Arup, Ulf Published in: Mycological Research DOI: 10.1017/S0953756204009888 2004 Link to publication Citation for published version (APA): Grube, M., Baloch, E., & Arup, U. (2004). A phylogenetic study of the Lecanora rupicola group (Lecanoraceae, Ascomycota). Mycological Research, 108(5), 506-514. https://doi.org/10.1017/S0953756204009888 Total number of authors: 3 General rights Unless other specific re-use rights are stated the following general rights apply: Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal Read more about Creative commons licenses: https://creativecommons.org/licenses/ Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. LUND UNIVERSITY PO Box 117 221 00 Lund +46 46-222 00 00 Download date: 02. Oct. 2021 Mycol. Res. 108 (5): 506–514 (May 2004). f The British Mycological Society 506 DOI: 10.1017/S0953756204009888 Printed in the United Kingdom. A phylogenetic study of the Lecanora rupicola group (Lecanoraceae, Ascomycota) Martin GRUBE1*, Elisabeth BALOCH1 and Ulf ARUP2 1 Institut fu¨r Botanik, Karl-Franzens-University Graz, Holteigasse 6, A-8010 Graz, Austria. 2 Department of Plant Ecology and Systematics, University of Lund, O¨stra Vallgatan 18, SE-223 61 Lund, Sweden. E-mail : [email protected] Received 16 September 2003; accepted 17 December 2003. A molecular phylogeny of the Lecanora rupicola group is presented, based on ITS sequence analyses. The study includes saxicolous and corticolous members of the Lecanora rupicola group as well as other Lecanora species with pruinose apothecia. A phylogenetic hypothesis for species in Lecanora s. lat. and various other genera in Lecanoraceae, based on an alignment-free distance estimation technique, shows that the Lecanora rupicola group forms a monophyletic clade within Lecanoraceae. Affinities to the core group of Lecanora are not well supported, likewise the monophyly of Lecanora s. str. with other species groups in Lecanora, such as the lobate taxa (and Rhizoplaca) is not supported. A more detailed analysis involving Lecanora species with pruinose apothecial discs was carried out with model-based Bayesian Markov chain Monte Carlo (B/MCMC) tree sampling. The results suggest the monophyly of the Lecanora species that are characterized by the presence of chromones. Corticolous as well as saxicolous species are included. Lepraria flavescens is closely related to the Lecanora swartzii subgroup, and the new name Lecanora rouxii nom. nov. is introduced for that species. Other Lecanora species with pruinose discs are not closely related to the Lecanora rupicola group. INTRODUCTION and heterogeneous crustose genus. Due to the size of the genus (ca 300 spp.; Kirk et al. 2001) it is under- The classification of lichenized genera, as it evolved standable that there exists no recent monograph of during the past two centuries, was guided primarily the whole genus, except for the posthumous publi- by morphological and chemical characters. This was cation of Motyka (1995, 1996a, b, c), which has been straightforward since, in contrast to the mycelia of most rejected as a nomenclatural work. Certain morpho- non-lichenized fungi, lichen thalli provide a number of logically defined groups, have, however, been studied morphological characters that are easily observable. in detail, for example subgenus Placodium (Poelt 1958), A higher number of thallus characters is usually found the Lecanora rupicola group (Leuckert & Poelt 1989), in lichens with a more complicated organization, i.e. the Lecanora dispersa group (Poelt, Leuckert & Roux with foliose or fruticose growth. This contributed to 1995), corticolous species with pruinose discs a present-day situation with numerous well-delimited (Lumbsch et al. 1997), species with a dark hypothecium foliose and fruticose genera, whereas some large crus- (Lumbsch, Guderley & Elix 1996), or regional mono- tose genera (e.g. Arthonia, Buellia, Caloplaca, Leca- graphs of the Lecanora subfusca group (Brodo 1984, nora), are insufficiently circumscribed and understood Miyawaki 1988, Lumbsch 1994, Ju¨ riado 1998, Guderley using anachronistic concepts. Most lichenologists 1999). The delimitation and relationships of these agree that such large crustose genera are hetero- groups to others in the genus, could not be resolved geneous, but due to the low number of synapomor- by these studies, and requires molecular phylogenetic phic characters only a few, sometimes monotypic, approaches. genera have been split from the large complexes. Phylogenetic analyses in Lecanora were previously Lecanora, a representing the largest order of lichen- carried out by Arup & Grube (1998, 2000). These ized fungi, Lecanorales, is a perfect example of a large studies indicated that certain genera that were segre- gated from the huge core genus Lecanora due to their * Corresponding author. deviating growth forms, form monophyletic groups M. Grube, E. Baloch and U. Arup 507 within Lecanora. For example, Arctopeltis and Rhizo- DNA extraction, amplification, and sequencing placa were split from Lecanora only due to their foliose Total DNA was extracted from individual thalli growth form, yet the phylogenetic position of these according to a modified CTAB method (Cubero et al. genera within Lecanora is supported by secondary 1999) or using the DNeasy Plant Mini Kit (Quiagen, compounds: Arctopeltis containing chlorinated lichex- Vienna). DNA-extracts were used for PCR-amplifi- anthones is placed within the Lecanora dispersa group, cation of the ITS regions including the 5.8S gene of the which is usually rich in xanthones; and Rhizoplaca, nuclear rDNA. Alternatively to DNA-extraction, we possessing usnic acid, is related to lobate Lecanora also used algal-free sections from the lichen medulla for species which share this compound. While these two direct amplification in some cases. Primers for amplifi- genera are traditionally accepted as well-circumscribed cation were ITS1F (Gardes & Bruns 1993) and ITS4 segregates, other species groups are maintained in (White et al. 1990). 50 ml PCR mix (10 mM Tris pH 8.3/ Lecanora. 50 mM KCl/1.5 mM MgCl /50 mg gelatine) contained Beside the xanthone-rich Lecanora dispersa group, 2 1.25 U polymerase (Taq DNA polymerase, Amer- or various usnic acid containing groups, the L. rupicola sham), 0.2 mM of each of the four dNTPs, 0.5 mM of group is chemically distinct in Lecanora. All species each primer and ca 10–50 ng genomic DNA. Products assigned to this group are characterized by apothecial were cleaned using QIAquick PCR Purification Kit discs covered by a conspicuous whitish to yellowish (Qiagen). Both complementary strands were sequenced pruina, which is composed of sordidone (Huneck & using the BigDye Terminator Ready Reaction Kit Santesson 1969, Devlin et al. 1971). Members of this (Applied Biosystems, Vienna) according to the manu- group comprise saxicolous crustose lichens, and an facturers instructions. Sequences were run on an ABI earlier monographic study accepted four species 310 automated sequencer (Applied Biosystems) and (Leuckert & Poelt 1989). Only two of these were in- assembled with AutoAssembler (Applied Biosystems). cluded in the previous molecular studies of Lecanora Table 1 presents a list of specimens for which we by Arup & Grube (1998, 2000). sequenced the ITS rDNA. For the following species Sordidone is a unique chromone, which usually we retrieved sequences from GenBank: Biatora helvola occurs together with trace amounts of the dechlorinated AJ247557, Biatora subduplex AJ247540, Lecanora accessory compound eugenitol. Chromones are only albescens AF070033, L. allophana AF159939, L. carpi- known from a few other lichens which are not closely nea AF070020, L. garovaglii AF189718, L. muralis related to Lecanora, for example Siphula and Haema- AF070015, L. pruinosa AF070018, Lepraria flavescens tomma of the Lecanorales, and the Roccellaceae of AF517889, Protoparmelia badia AF070023, Pyrrho- the Arthoniales. Certain chromone derivatives are un- spora quernea AF517930, Rhizoplaca chrysoleuca common products also in few non-lichenized fungi AF159940, and R. peltata AF159925. (Turner & Aldridge 1983, Fujimoto et al. 2003, Lin The alignment for the B/MCMC analysis was et al. 2003), but so far, sordidone is only known from produced using a linear Hidden Markov Model Lecanora and one species assigned to Lepraria. Apart (HMM) implemented in the software SAM (Hughey & from the Lecanora rupicola group (sensu Leuckert & Krogh 1996; http://www.cse.ucsc.edu/reseqrch/comp- Poelt 1989), certain bark-inhabiting Lecanora species bio/sam.html). also contain sordidone. The corticolous species with pruinose discs were recently revised by Lumbsch et al. (1997), who accepted four species which contain Phylogenetic analyses sordidone. One of these, L. carpinea, was included by Arup
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