The genus Fomitopsis (Polyporales, Basidiomycota) reconsidered.

Based on seven- and three-gene datasets, we discuss four alternative approaches for a reclassification of Fomitopsidaceae (Polyporales, Basidiomycota). After taking into account morphological diversity in the family, we argue in favour of distinguishing three genera only, viz. Anthoporia, Antrodia and Fomitopsis. Fomitopsis becomes a large genus with 128 accepted species, containing almost all former Fomitopsis spp. and most species formerly placed in Antrodia, Daedalea and Laccocephalum. Genera Buglossoporus, Cartilosoma, Daedalea, Melanoporia, Neolentiporus, alongside twenty others, are treated as synonyms of Fomitopsis. This generic scheme allows for morphologically distinct genera in Fomitopsidaceae, unlike other schemes we considered. We provide arguments for retaining Fomitopsis and suppressing earlier (Daedalea, Caloporus) or simultaneously published generic names (Piptoporus) considered here as its synonyms. Taxonomy of nine species complexes in the genus is revised based on ITS, ITS + TEF1, ITS + TEF1 + RPB1 and ITS + TEF1 + RPB2 datasets. In total, 17 species are described as new to science, 26 older species are reinstated and 26 currently accepted species names are relegated to synonymy. A condensed identification key for all accepted species in the genus is provided. Taxonomic novelties: New species: Fomitopsis algumicola Grebenc & Spirin, F. caseosa Vlasák & Spirin, F. cupressicola Vlasák, J. Vlasák Jr. & Spirin, F. derelicta Vlasák & Spirin, F. dollingeri Vlasák & Spirin, F. fissa Vlasák & Spirin, F. lapidosa Miettinen & Spirin, F. lignicolor Vlasák & Spirin, F. maculosa Miettinen & Spirin, F. pannucea Runnel & Spirin, F. perhiemata Viner & Spirin, F. purpurea Spirin & Ryvarden, F. retorrida Spirin & Kotiranta, F. solaris Rivoire, A.M. Ainsworth & Vlasák, F. tristis Miettinen & Spirin, F. tunicata Miettinen & Spirin, F. visenda Miettinen & Spirin. New combinations: Fomitopsis aculeata (Cooke) Spirin & Miettinen, F. aethalodes (Mont.) Spirin, F. alaskana (D.V. Baxter) Spirin & Vlasák, F. albidoides (A. David & Dequatre) Bernicchia & Vlasák, F. amygdalina (Berk. & Ravenel) Spirin & Vlasák, F. angusta (Spirin & Vlasák) Spirin & Vlasák, F. atypa (Lév.) Spirin & Vlasák, F. caespitosa (Murrill) Spirin & Miettinen, F. calcitrosa (Spirin & Miettinen) Spirin & Miettinen, F. circularis (B.K. Cui & Hai J. Li) Spirin, F. concentrica (G. Cunn.) M.D. Barrett, F. cyclopis (Miettinen & Spirin) Miettinen & Spirin, F. dickinsii (Berk. ex Cooke) Spirin, F. elevata (Corner) Spirin & Miettinen, F. eucalypti (Kalchbr.) Spirin, F. ferrea (Cooke) Spirin & Viner, F. flavimontis (Vlasák & Spirin) Vlasák & Spirin, F. foedata (Berk.) Spirin & Miettinen, F. gilvidula (Bres.) Spirin & Miettinen, F. glabricystidia (Ipulet & Ryvarden) Miettinen & Ryvarden, F. globispora (Ryvarden & Aime) Spirin, F. hartmannii (Cooke) M.D. Barrett & Spirin, F. hyalina (Spirin, Miettinen & Kotir.) Spirin & Miettinen, F. hypoxantha (Bres.) Spirin & Miettinen, F. incana (Lév.) Spirin & V. Malysheva, F. infirma (Renvall & Niemelä) Miettinen & Niemelä, F. juniperina (Murrill) Spirin & Vlasák, F. kuzyana (Pilát ex Pilát) Spirin & Vlasák, F. leioderma (Mont.) Spirin & Vlasak, F. leucaena (Y.C. Dai & Niemelä) Spirin & Miettinen, F. luzonensis (Murrill) Spirin & Miettinen, F. maculatissima (Lloyd) Spirin, F. madronae (Vlasák & Ryvarden) Vlasák & Ryvarden, F. malicola (Berk. & M.A. Curtis) Spirin, F. marchionica (Mont.) Spirin & Miettinen, F. marianii (Bres.) Spirin, Vlasák & Cartabia, F. mellita (Niemelä & Penttilä) Niemelä & Miettinen, F. microcarpa (B.K. Cui & Shun Liu) Spirin, F. micropora (B.K. Cui & Shun Liu) Spirin, F. modesta (Kuntze ex Fr.) Vlasák & Spirin, F. monomitica (Yuan Y. Chen) Spirin & Viner, F. morganii (Lloyd) Spirin & Vlasák, F. moritziana (Lév.) Spirin & Miettinen, F. neotropica (D.L. Lindner, Ryvarden & T.J. Baroni) Vlasák, F. nigra (Berk.) Spirin & Miettinen, F. nivosella (Murrill) Spirin & Vlasák, F. oboensis (Decock, Amalfi & Ryvarden) Spirin, F. oleracea (R.W. Davidson & Lombard) Spirin & Vlasák, F. philippinensis (Murrill) Spirin & Vlasák, F. primaeva (Renvall & Niemelä) Miettinen & Niemelä, F. psilodermea (Berk. & Mont.) Spirin & Vlasák, F. pulverulenta (Rivoire) Rivoire, F. pulvina (Pers.) Spirin & Vlasák, F. pulvinascens (Pilát ex Pilát) Niemelä & Miettinen, F. quercina (L.) Spirin & Miettinen, F. ramentacea (Berk. & Broome) Spirin & Vlasák, F. renehenticii (Rivoire, Trichies & Vlasák) Rivoire & Vlasák, F. roseofusca (Romell) Spirin & Vlasák, F. sagraeana (Mont.) Vlasák & Spirin, F. sandaliae (Bernicchia & Ryvarden) Bernicchia & Vlasák, F. sclerotina (Rodway) M.D. Barrett & Spirin, F. serialiformis (Kout & Vlasák) Vlasák, F. serialis (Fr.) Spirin & Runnel, F. serrata (Vlasák & Spirin) Vlasák & Spirin, F. squamosella (Bernicchia & Ryvarden) Bernicchia & Ryvarden, F. stereoides (Fr.) Spirin, F. subectypa (Murrill) Spirin & Vlasák, F. substratosa (Malençon) Spirin & Miettinen, F. tropica (B.K. Cui) Spirin, F. tumulosa (Cooke) M.D. Barrett & Spirin, F. tuvensis (Spirin, Vlasák & Kotir.) Spirin & Vlasák, F. uralensis (Pilát) Spirin & Miettinen, F. ussuriensis (Bondartsev & Ljub.) Spirin & Miettinen, F. variiformis (Peck) Vlasák & Spirin, F. yunnanensis (M.L. Han & Q. An) Spirin, Daedaleopsis candicans (P. Karst.) Spirin, Megasporoporia eutelea (Har. & Pat.) Spirin & Viner, Neofomitella hemitephra (Berk.) M.D. Barrett, Pseudophaeolus soloniensis (Dubois) Spirin & Rivoire, P. trichrous (Berk. & M.A. Curtis) Vlasák & Spirin. New synonyms: Antrodia bondartsevae Spirin, A. huangshanensis Y.C. Dai & B.K. Cui, A. taxa T.T. Chang & W.N. Chou, A. wangii Y.C. Dai & H.S. Yuan, Antrodiella subnigra Oba, Mossebo & Ryvarden, Antrodiopsis Audet, Boletus quercinus Schrad., Brunneoporus Audet, Buglossoporus Kotl. & Pouzar, Buglossoporus eucalypticola M.L. Han, B.K. Cui & Y.C. Dai, Caloporus P. Karst., Cartilosoma Kotlaba & Pouzar, Coriolus clemensiae Murrill, C. cuneatiformis Murrill, C. hollickii Murrill, C. parthenius Hariot & Pat., C. rubritinctus Murrill, Daedalea Pers., Daedalea allantoidea M.L. Han, B.K. Cui & Y.C. Dai, D. americana M.L. Han, Vlasák & B.K. Cui, D. radiata B.K. Cui & Hai J. Li, D. rajchenbergiana Kossmann & Drechsler-Santos, D. sinensis Lloyd, Daedalella B.K. Cui & Shun Liu, Dentiporus Audet, Flavidoporia Audet, Fomes subferreus Murrill, Fomitopsis cana B.K. Cui, Hai J. Li & M.L. Han, F. caribensis B.K. Cui & Shun Liu, F. cystidiata B.K. Cui & M.L. Han, F. ginkgonis B.K. Cui & Shun Liu, F. iberica Melo & Ryvarden, F. incarnata K.M. Kim, J.S. Lee & H.S. Jung, F. subfeei B.K. Cui & M.L. Han, F. subtropica B.K. Cui & Hai J. Li, Fragifomes B.K. Cui, M.L. Han & Y.C. Dai, Leptoporus epileucinus Pilát, Melanoporia Murrill, Neoantrodia Audet, Neolentiporus Rajchenb., Nigroporus macroporus Ryvarden & Iturr., Niveoporofomes B.K. Cui, M.L. Han & Y.C. Dai, Pilatoporus Kotl. & Pouzar, Piptoporus P. Karst., Polyporus aurora Ces., P. durescens Overh. ex J. Lowe, P. griseodurus Lloyd, Poria incarnata Pers., Pseudoantrodia B.K. Cui, Y.Y. Chen & Shun Liu, Pseudofomitopsis B.K. Cui & Shun Liu, Ranadivia Zmitr., Rhizoporia Audet, Rhodofomes Kotl. & Pouzar, Rhodofomitopsis B.K. Cui, M.L. Han & Y.C. Dai, Rhodofomitopsis pseudofeei B.K. Cui & Shun Liu, R. roseomagna Nogueira-Melo, A.M.S. Soares & Gibertoni, Rubellofomes B.K. Cui, M.L. Han & Y.C. Dai, Subantrodia Audet, Trametes fulvirubida Corner, T. lignea Murrill, T. lusor Corner, T. pseudodochmia Corner, T. subalutacea Bourdot & Galzin, T. supermodesta Ryvarden & Iturr., T. tuberculata Bres., Tyromyces multipapillatus Corner, T. ochraceivinosus Corner, T. palmarum Murrill, T. singularis Corner, T. squamosellus Núñez & Ryvarden, Ungulidaedalea B.K. Cui, M.L. Han & Y.C. Dai. Lectotypes: Hexagonia sulcata Berk., Polyporus castaneae Bourdot & Galzin, Poria incarnata Pers., Trametes subalutacea Bourdot & Galzin, Ungulina substratosa Malençon. Neotypes: Agaricus soloniensis Dubois, Boletus pulvinus Pers. Citation: Spirin V, Runnel K, Vlasák J, Viner I, Barrett MD, Ryvarden L, Bernicchia A, Rivoire B, Ainsworth AM, Grebenc T, Cartabia M, Niemelä T, Larsson K-H, Miettinen O (2024). The genus Fomitopsis (Polyporales, Basidiomycota) reconsidered. Studies in Mycology 107: 149-249. doi: 10.3114/sim.2024.107.03.

A revised genus-level classification for Cerrenaceae (Polyporales, Agaricomycetes).

Cerrenaceae is a small family of polypores and hydnoid fungi in the order Polyporales (Basidiomycota). The family consists of white-rot fungi, some of which are serious tree pathogens. Combining morphological evidence with a phylogenetic dataset of six genetic markers, we revise generic concepts in the family and propose a seven-genus classification system for the family. Two genera are introduced as new: the monotypic Acanthodontia for Radulodon cirrhatinus, and Lividopora for the Rigidoporus vinctus complex. We re-introduce the name Somion for the Spongipellis delectans complex. Other recognized genera in the family are Cerrena, Irpiciporus, Pseudolagarobasidium, and Radulodon. New species introduced are Irpiciporus branchiformis from Tanzania, Lividopora armeniaca, and L. facilis from Southeast Asia, and Somion strenuum from East Asia. We provide nomenclatural comments on all the names combined to the above Cerrenaceae genera and typify Cerrena unicolor, C. zonata, Polyporus carneopallens (= L. vincta), Somion occarium, and S. unicolor. The genus Hyphoradulum belongs to Cystostereaceae (Agaricales), and we transfer the type species H. conspicuum to Crustomyces. Our study highlights the importance of integrating different basidiocarp types in analyses when revising genus classification in macrofungi. Citation: Miettinen O, Vlasák J, Larsson E, Vlasák J Jr., Seelan JSS, Hernawati, Levicky Q, Larsson K-H, Spirin V (2023). A revised genus-level classification for Cerrenaceae (Polyporales, Agaricomycetes). Fungal Systematics and Evolution 12: 271-322. doi: 10.3114/fuse.2023.12.14.

Four new species of Hydnellum (Thelephorales, Basidiomycota) with a note on Sarcodon illudens.

Four new Hydnellum species are described. Hydnellum roseoviolaceum sp. nov. grows in dry pine heaths on acidic, sandy soil. It is close to H. fuligineoviolaceum, another pine-associated species, but differs by smaller spores, an initially rose-coloured instead of violet flesh in fresh basidiomata and a mild taste. Hydnellum scabrosellum sp. nov. grows in coniferous forests on calcareous soil. It shares a general morphology with H. scabrosum, which also is its closest relative. It differs by having smaller and slenderer basidiomata and by the yellowish ochraceous colour of flesh and spines in dried specimens compared to the whitish or reddish brown colour seen in H. scabrosum. Hydnellum fagiscabrosum sp. nov. is another species with morphological and phylogenetic affinities to H. scabrosum. However, it is associated with trees from Fagales whereas H. scabrosum is associated with Pinaceae. Hydnellum nemorosum sp. nov. is yet another species that associates with broadleaved trees. It seems to be a rare species, morphologically reminiscent of H. fuligineoviolaceum, H. ioeides and H. scabrosum, but it is phylogenetically close to H. fennicum. Sequences from the type specimens of H. glaucopus, H. lepidum, H. scabrosum, Sarcodon illudens and S. regalis are included in the analyses. Specimens given the provisional name "Sarcodon pseudoglaucopus" in Sweden are now shown to be referable to S. illudens. The analyses further showed that S. illudens is close to H. lepidum. The new combination Hydnellum illudens is proposed. Sarcodon regalis and H. lepidum are shown to be conspecific and, although their basionyms were simultaneously published, the name S. regalis was only validated in a later publication. Hydnellum lepidum therefore takes priority and S. regalis becomes a synonym. Citation: Nitare J, Ainsworth AM, Larsson E, Parfitt D, Suz LM, Svantesson S, Larsson K-H (2021). Four new species of Hydnellum (Thelephorales, Basidiomycota) with a note on Sarcodon illudens. Fungal Systematics and Evolution 7: 233-254. doi: 10.3114/fuse.2021.07.12.

Polyozellus vs. Pseudotomentella: generic delimitation with a multi-gene dataset.

Polyozellus and Pseudotomentella are two genera of closely related, ectomycorrhizal fungi in the order Thelephorales; the former stipitate and the latter corticioid. Both are widespread in the Northern Hemisphere and many species from both genera seem to be restricted to old growth forest. This study aimed to: a) identify genetic regions useful in inferring the phylogenetic relationship between Polyozellus and Pseudotomentella, b) infer this relationship with the regions identified and c) make any taxonomic changes warranted by the result. RPB2, mtSSU and nearly full-length portions of nrLSU and nrSSU were found to be comparatively easy to sequence and provide a strong phylogenetic signal. A STACEY species tree of these three regions revealed that Polyozellus makes Pseudotomentella paraphyletic. As a result, nearly all species currently placed in Pseudotomentella were recombined to Polyozellus. Pseudotomentella larsenii was found to be closer to Tomentellopsis than Polyozellus, but its placement needs further study and it was hence not recombined.

Elaphomyces section Elaphomyces (Eurotiales, Ascomycota) - taxonomy and phylogeny of North European taxa, with the introduction of three new species.

The North European species of Elaphomyces section Elaphomyces (Eurotiales, Pezizomycotina) are studied. Three new species, E. citrinopapillatus, E. pusillus, and E. roseoviolaceus are introduced and verified by morphology and sequence data from ITS, nuclear LSU, mitochondrial SSU, and ß-tubulin. A lectotype for Elaphomyces granulatus is selected. Elaphomyces granulatus and E. muricatus are epitypified with sequenced material from the Femsjö region in South Sweden. Elaphomyces striatosporus is epitypified with sequenced material from the vicinity of the type locality in Norway. A key to all species of Elaphomyces occurring in Denmark, Norway, and Sweden is provided.

Hydnoporia, an older name for Pseudochaete and Hymenochaetopsis, and typification of the genus Hymenochaete (Hymenochaetales, Basidiomycota).

The genus name Hydnoporia is reinstated to encompass the Hymenochaete tabacina group currently addressed to Pseudochaete (illegitimate genus) or Hymenochaetopsis. Identity of the type species of Hydnoporia, Sistotrema fuscescens (= Hydnoporia olivacea in current sense), is clarified, and a lectotype is selected. In total, 12 species are combined in Hydnoporia: H. corrugata, H. gigasetosa, H. lamellata, H. laricicola, H. latesetosa, H. lenta, H. rhododendri, H. rimosa, H. subrigidula, H. tabacina, H. tabacinoides, and H. yasudai. Hydnoporia diffissa is described as new. Analyses of all available ITS (94) and newly produced tef1 sequences (20) indicate that there are at least 20-27 species in the genus. Identity of the type species of Hymenochaete, H. rubiginosa, is clarified; the name is retained for the species so named in Europe while other species are present in North America and East Asia. Additionally, three new combinations in Hymenochaete are proposed: H. campylopora (= Cyclomyces fuscus), H. microcycla (= Cyclomyces tabacinus), and H. saepiaria.

A preliminary overview of the corticioid Atractiellomycetes (Pucciniomycotina, Basidiomycetes).

The taxonomy of the corticioid fungi from the class Atractiellomycetes (Pucciniomycotina, Basidiomycetes) currently addressed to the genus Helicogloea, is revised based on morphological and nuclear ribosomal DNA (ITS and LSU) data. The genus is restricted to 25 species with semitranslucent, gelatinous basidiocarps lacking differentiated cystidia and clamps on hyphae, of which 11 are described as new to science. The asexual genus Leucogloea is placed as a synonym of Helicogloea s. str. Since the type species of Saccoblastia, S. ovispora, is combined to Helicogloea, a new genus, Saccosoma, is introduced to encompass Saccoblastia farinacea and six related species, one of which is described as new. In contrast to Helicogloea in the strict sense, the basidiocarps of Saccosoma are arid, not gelatinized, and hyphae are clamped. The third lineage of the corticioid Atractiellomycetes is represented by the Bourdotigloea vestita complex. Species of Bourdotigloea are devoid of clamps but often possess well-differentiated cystidia, as well as long, cylindrical-fusiform basidiospores. Bourdotigloea encompasses nine species, of which six are described here as new.

Vertical distribution of ectomycorrhizal fungal taxa in a podzol soil profile.

• Studies of ectomycorrhizal fungal communities in forest soils are usually restricted to the uppermost organic horizons. Boreal forest podzols are highly stratified and little is known about the vertical distribution of ectomycorrhizal communities in the underlying mineral horizons. • Ectomycorrhizal root tips were sampled from seven horizons in three continuous columns of a 52-cm deep podzol profile. Root tips were sorted into morphological groups and the colonising fungi identified by sequencing of the rDNA ITS region. The vertical distribution of mycorrhizal taxa was examined. • A relationship between ectomycorrhizal species composition and soil horizon was found. Tomentellopsis submollis, three Piloderma species and Dermocybe spp. were found predominantly in the upper horizons while Suillus luteus, Lactarius utilis and three undescribed Piloderma species were associated with the mineral horizons. • Two thirds of the root tips were found in the mineral soil and half of the taxa were restricted to the mineral horizons. The results highlight the need to include the mineral soil in order to gain a more accurate representation of the ectomycorrhizal community.

Diversity and abundance of resupinate thelephoroid fungi as ectomycorrhizal symbionts in Swedish boreal forests.

Resupinate thelephoroid fungi (hereafter called tomentelloid fungi) have a world-wide distribution and comprise approximately 70 basidiomycete species with inconspicuous, resupinate sporocarps. It is only recently that their ability to form ectomycorrhizas (EM) has been realized, so their distribution, abundance and significance as mycobionts in forest ecosystems is still largely unexplored. In order to provide baseline data for future ecological studies of tomentelloid fungi, we explored their presence and abundance in nine Swedish boreal forests in which the EM communities had been analysed. Phylogenetic analyses were used to compare the internal transcribed spacer of nuclear ribosomal DNA (ITS rDNA) sequence data obtained from mycobionts on single ectomycorrhizal tips with that obtained from sporocarps of identified tomentelloid fungi. Five species of Tomentella and one species of Pseudotomentella were identified as ectomycorrhizal fungi. The symbiotic nature of Tomentella bryophila, T. stuposa, T. badia and T. atramentaria is demonstrated for the first time. T. stuposa and Pseudotomentella tristis were the most commonly encountered tomentelloid fungi, with the other species, including T. sublilacina, only being recorded from single stands. Overall, tomentelloid fungi were found in five of the studies, colonizing between 1 and 8% of the mycorrhizal root tips. Two of the five sites supported several tomentelloid species. Tomentelloid fungi appear to be relatively common ectomycorrhizal symbionts with a wide distribution in Swedish coniferous forests. The results are in accordance with accumulating data that fungal species which lack conspicuous sporocarps may be of considerable importance in EM communities.

Import and processing of the precursor of the delta subunit of tobacco chloroplast ATP synthase.

A cDNA clone encoding the complete precursor of the delta subunit of chloroplast ATP synthase has been isolated from a tobacco (Nicotiana tabacum) leaf cDNA library in lambda gt11. The 880 bp insert encodes a polypeptide of 248 amino acid residues, of which 61 residues constitute an N-terminal presequence and 187 residues make up the mature delta subunit. Transcription and translation of the cDNA in vitro produced a protein of 29 kDa which was imported by isolated pea chloroplasts and processed to the mature 20 kDa subunit. The delta subunit precursor was processed to the mature size by a processing peptidase present in pea stromal extracts. Hybridisation of the cDNA to Southern blots of tobacco genomic DNA suggests the presence of two genes in the haploid genome.

Import and processing of the precursor form of the gamma subunit of the chloroplast ATP synthase from tobacco.

A cDNA clone encoding the complete precursor of the gamma subunit of chloroplast ATP synthase has been isolated from a tobacco (Nicotiana tabacum) leaf cDNA library in lambda gt11. The 1.4 kb insert encodes a polypeptide of 377 amino acid residues, of which 55 residues constitute an N-terminal presequence and 322 residues make up the mature gamma subunit. Hybridisation of the cDNA to Southern blots of tobacco genomic DNA indicates the presence of two genes in the haploid genome. Transcription and translation of the cDNA in vitro produced a protein of 41 kDa which was imported by isolated pea chloroplasts and processed to the mature 36 kDa subunit. The gamma subunit precursor was processed to the mature size by a processing peptidase of 180 kDa present in pea stromal extracts.