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Molecular Phylogeny and Taxonomic Position of Trametes Cervina and Description of a New Genus Trametopsis

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Molecular Phylogeny and Taxonomic Position of Trametes Cervina and Description of a New Genus Trametopsis CZECH MYCOL. 60(1): 1–11, 2008 Molecular phylogeny and taxonomic position of Trametes cervina and description of a new genus Trametopsis MICHAL TOMŠOVSKÝ Faculty of Forestry and Wood Technology, Mendel University of Agriculture and Forestry in Brno, Zemědělská 3, CZ-613 00, Brno, Czech Republic [email protected] Tomšovský M. (2008): Molecular phylogeny and taxonomic position of Trametes cervina and description of a new genus Trametopsis. – Czech Mycol. 60(1): 1–11. Trametes cervina (Schwein.) Bres. differs from other species of the genus by remarkable morpho- logical characters (shape of pores, hyphal system). Moreover, an earlier published comparison of the DNA sequences within the genus revealed considerable differences between this species and the re- maining European members of the genus Trametes. These results were now confirmed using se- quences of nuclear LSU and mitochondrial SSU regions of ribosomal DNA. The most related species of Trametes cervina are Ceriporiopsis aneirina and C. resinascens. According to these facts, the new genus Trametopsis Tomšovský is described and the new combination Trametopsis cervina (Schwein.) Tomšovský is proposed. Key words: Trametopsis, Trametes, ribosomal DNA, polypore, taxonomy. Tomšovský M. (2008): Molekulární fylogenetika a taxonomické zařazení outkovky jelení, Trametes cervina, a popis nového rodu Trametopsis. – Czech Mycol. 60(1): 1–11. Outkovka jelení, Trametes cervina (Schwein.) Bres., se liší od ostatních zástupců rodu nápadnými morfologickými znaky (tvar rourek, hyfový systém). Také dříve uveřejněné srovnání sekvencí DNA v rámci rodu Trametes odhalilo významné rozdíly mezi tímto druhem a ostatními evropskými zástupci rodu. Uvedené výsledky byly nyní potvrzeny za použití sekvencí jaderné LSU a mitochondriální SSU oblasti ribozomální DNA, přičemž nejpříbuznějšími druhu Trametes cervina jsou Ceriporiopsis anei- rina a C. resinascens. Na základě těchto skutečností je popsán nový rod Trametopsis Tomšovský a je navržena druhová kombinace Trametopsis cervina (Schwein.) Tomšovský. INTRODUCTION The taxonomic position of Trametes cervina (Schwein.) Bres. was not suffi- ciently resolved until recently. The species differs from other European Trametes species by large irregular pores tending to split, forming daedaleoid, nearly irpicoid hymenophores, and by a distinct hyphal system quite unique among Trametes species. The hyphal system of Trametes cervina is not easy to describe. Different authors classify it as dimitic (Kotlaba and Pouzar 1957, 1983; Ryvarden and Gilbertson 1994) or trimitic (Donk 1974, Bernicchia 2005). The most detailed 1 CZECH MYCOL. 60(1): 1–11, 2008 description of the hyphal system of this species was published by Jahn (1983), who stated it to have a “dimitic with a trimitic aspect“. The problem of the doubt- ful classification of the hyphal system is caused by the presence of numerous sclerified thick-walled intercalary segments of the contextual generative hyphae, which may be confused with genuine skeletal hyphae when the distal clamps are overlooked or detached by cutting. The second type of thick-walled hyphae is not easy to classify as it possesses characters of both skeletal and binding hyphae. These hyphae originate from a generative hypha, they are thick-walled, rather rarely branched, usually curved and flexuous. This type of skeletoid hyphae (term proposed by Jahn 1983) is quite unique and differs from true binding hyphae in Trametes s. str. According to its hyphal system, T. cervina was in the past placed in the genus Antrodia (Kotlaba and Pouzar 1983). However, the genus Antrodia encompasses species causing a brown rot, while the rot of T. cervina is white. The white rot na- ture of T. cervina was confirmed (Tomšovský and Homolka 2004), but the produc- tion of ligninolytic enzymes is substantially lower (almost undetectable) if com- pared with genuine Trametes species. Dai (1996) proposed the combination Funalia cervina (Schwein.) Y. C. Dai due to cyanophilous skeletal hyphae and a change of colour in the hairs on the pileus after KOH treatment. Molecular phylo- geny methods based on comparisons of DNA sequences of the ITS and LSU regions of ribosomal DNA were applied to T. cervina by Tomšovský et al. (2006). The re- sults did not support the theory about close relatedness between T. cervina and other European Trametes species or fungi traditionally placed in Funalia (Funalia is the synonym of Coriolopsis; see details in Ryvarden and Gilbertson 1994). Ac- cording to Ryvarden and Gilbertson (1994), pileate fungi with skeletal hyphae caus- ing white rot should in general belong to the genus Tyromyces. However, the pileus habitus of the two taxa is different – pilei of Tyromyces are sappy to watery while those of Trametes cervina are tough. Therefore, the putative relatedness of T. cervina and Tyromyces should be tested by molecular phylogeny methods. The results of Tomšovský et al. (2006) revealed remarkable proximity of T. cervina to Ceriporiopsis aneirina among a large DNA set encompassing vari- ous species of Aphyllophorales. Nevertheless, the DNA of C. aneirina was not se- quenced by the authors themselves. The sequence was obtained from the GenBank database and the only ITS sequence of the species was accessible at the time of manuscript preparation. Due to possible misidentification of the se- quenced fungus, the proximity of T. cervina and C. aneirina should be confirmed by further study including new sequences of the latter species. The aim of the study was to solve the taxonomic position of Trametes cervina us- ing widely adopted methods of molecular taxonomy applying DNA sequences of nu- clear (large subunit – LSU) and mitochondrial (small subunit – mitSSU) ribosomal DNA with regard to macro– and micromorphological characters of the species. 2 TOMŠOVSKÝ M.: MOLECULAR PHYLOGENY AND TAXONOMIC POSITION OF TRAMETES CERVINA MATERIAL AND METHODS The earlier obtained DNA sequences of Trametes cervina LSU region (Tab. 1) were completed by mitSSU sequences of Trametes cervina and LSU and mitSSU sequences of Ceriporiopsis aneirina and C. resinascens. C. resinascens was added to the study due to its close proximity to C. aneirina, whereas neither species seems to be closely related to the type of the genus – C. gilvescens (Tomšovský, in prep.). The DNA was isolated from the dried tissue of herbarium specimens or from cultures grown on Petri dishes with 2 % malt extract agar according to Tomšovský et al. (2006). For numbers of source herbarium specimens and cultures, see Tab. 1. Tab. 1. Fungal material newly sequenced in this study and analysed sequences from the GenBank. Species Geographic Herbarium Culture num- GenBank Reference origin specimen ber* Acc. No.** Trametopsis cervina Czech Republic PRM CCBAS 018N AY855907 Tomšovský (as Trametes cervina) 900574 EU368500 et al. 2006 / this study Trametopsis cervina Czech Republic PRM – AY855917 Tomšovský (as Trametes cervina) 891515 – et al. 2006 Trametopsis cervina ? – DAOM 52884 – unpublished (as Trametes cervina) AY986497 Ceriporiopsis aneirina Czech Republic BRNM MUAF 888 EU368503 this study 706970 EU368504 Ceriporiopsis resinascens Czech Republic BRNM MUAF 886 EU368501 this study 706968 EU368502 Trametes suaveolens Czech Republic – CCBAS 026N AY855909 Tomšovský – et al. 2006 Trametes suaveolens ? – DAOM 196328 AF518656 Hibbett and – Binder 2002 Trametes suaveolens ? – DAOM 19632 – Hibbett and U27079 Donoghue 1995 Antrodia albida USA – CBS 308.82 AY515348 Kim et al. 2005 / DQ491441 unpublished Antrodia albida Belgium CBS CBS 458.86 AY515349 Kim et al. 2005 H-9334 – Antrodia albida Norway – FCUG 1100 AY333846 unpublished – Tyromyces chioneus ? KEW – AF393080 Binder and 141 AF334896 Hibbett 2002 * Culture collection abbreviations are listed according to Anonymus 2, except for MUAF (= Culture Collection of Dept. of Forest Protection and Wildlife Management, Mendel University, Brno). ** The first number refers to the LSU sequence, the second one to the mitSSU sequence. 3 CZECH MYCOL. 60(1): 1–11, 2008 For amplification of LSU and mitSSU the primer pairs LR0R/LR6 and MS1/MS2 were used (Anonymus 1; White et al. 1990). The DNA was amplified with PCR, using the Mastercycler® ep thermocycler (Eppendorf). PCR amplifications were performed according to Tomšovský et al. (2006). The PCR products were purified with NucleoSpin Extract II (Macherey-Nagel) prior to sequencing. Sequences were determined with an ABI PRISM 3100 Avant DNA sequencer (Applied Biosystems) at the Department of Animal Morphology, Physiology and Genetics, Faculty of Agriculture, Mendel Uni- versity, Brno using the ABI PRISM BigDye terminator v1.1 cycle sequencing kit (Applied Biosystems). All samples were sequenced with the primers used in the PCR. The sequences were deposited in the EMBL Nucleotide Sequence Database and their GenBank accession numbers are given in Tab. 1. Additional LSU and mitSSU sequences were searched for in the GenBank. The type species se- quences of the genera Antrodia, Trametes and Tyromyces (Antrodia albida, Trametes suaveolens and Tyromyces chioneus, respectively) were added to the alignment to elucidate the taxonomic position of Trametes cervina in relation to these genera (for GenBank accession numbers, see Tab. 1). In the case of mitSSU also the sequence of Trametes cervina deposited in the GenBank was added to the dataset. All sequences were edited manually using BioEdit version 4.7.1. (Hall 1999). The LSU alignment with introduced gaps consisted of 573 including 442 constant, 127 variable and 86 parsimony informa-
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