Mycological Progress (2018) 17:357–364 https://doi.org/10.1007/s11557-017-1362-5

ORIGINAL ARTICLE

Purpureocorticium microsporum () gen. et sp. nov. from East Asia

Sheng-Hua Wu1 & Dong-Mei Wang2 & Yu-Ping Chen1

Received: 25 July 2017 /Revised: 8 November 2017 /Accepted: 13 November 2017 /Published online: 25 November 2017 # German Mycological Society and Springer-Verlag GmbH Germany, part of Springer Nature 2017

Abstract A corticioid fungal found in mountainous Taiwan, and Yunnan Province of China, is described as a new and new species: Purpureocorticium microsporum (, Basidiomycota). Morphological study and phylogenetic analyses based on sequence data respectively derived from the large subunit nuclear ribosomal RNA gene (LSU nrDNA) alone and the combined dataset of LSU nrDNA and translation elongation factor 1-α gene (tef1-α) indicated that Purpureocorticium does not belong to any clade of the Agaricomycetes. Purpureocorticium and P. microsporum are characterized by having a resupinate basidiocarp with smooth hymenial surface which turns purple in KOH, and becomes purplish after storage, micro- scopically having compact texture of subiculum, nodose-septate thin-walled generative hyphae, absence of cystidia, basidia subclavate with a median constriction, or utriform, bearing ovate-ellipsoid small-sized basidiospores, which are smooth and thin- walled, inamyloid, and nondextrinoid. The new combination rubescens is proposed, based on morphological and phylogenetic evidences.

Keywords China . New genus . New species . Taiwan .

Yunnan Province of China. These corticioid fungal specimens were collected from dead angiosperm branches or on rotten trunks. The hymenial surface of this species is white when Introduction fresh, turning purple in KOH. On the other hand, its hymenial surface becomes purplish pink or pinkish purple after storage Corticioid fungi are members of Agaricomycotina, mainly of several years. Its microscopic features do not exactly fit the with resupinate basidiocarps. Evidence from molecular stud- concept of any known corticioid genera. The present study ies revealed that corticioid taxa are present in all major clades based on morphological characteristics and phylogenetic anal- on Agaricomycetes (Binder et al. 2005, 2010;Wuetal.2007; yses of DNA sequences inferred from the large subunit nucle- Larsson 2007). These studies also showed that it is barely ar ribosomal RNA gene (LSU nrDNA) alone and the com- possible to evaluate their order and family rank for a great bined dataset of LSU nrDNA and the translation elongation portion of the corticioid genera, based merely on studying factor 1-α gene (tef1-α) has revealed that these collections macro- and microscopic characters. represent a new genus and new species. In the survey of corticioid fungi in Taiwan and mainland China over two decades, an unfamiliar species has been col- lected several times in mountainous Taiwan, and also once in Materials and methods

Section Editor: Yu-Cheng Dai Morphological and cultural studies

* Sheng-Hua Wu Macro- and microscopic studies were based on dried speci- [email protected] mens. Thin free-hand sections of basidiocarps were prepared 1 Department of Biology, National Museum of Natural Science, for microscopic study. For observations and measurements of Taichung 404, Taiwan, Republic of China microscopic characters, the sections were mounted in 5% ’ 2 Department of Pharmacology, University of Illinois, KOH to ensure rehydration. Melzer sreagentwasappliedto Chicago, IL 60612, USA detect amyloidity and dextrinoidity. Cotton blue was used as a 358 Mycol Progress (2018) 17:357–364 mounting medium to determine cyanophily. The following heuristic search with 1000 random taxa stepwise addition se- abbreviations were used for basidiospore measurements: L = quences, TBR branch swapping, and MAXTREES set to mean spore length with standard deviation, W = mean spore autoincrease. All transformations were considered unordered width with standard deviation, Q = variation in L/W ratio, n = and equally weighted, with gaps treated as missing data. number of spores measured from each specimen. Living Calocera cornea was used as the outgroup for rooting pur- mycelia were isolated from the woody substratum beneath pose. The relative robustness of the clades was assessed by the the basidiocarps and grown on 1.5% malt extract agar. All bootstrap method (Hillis and Bull 1993) using 1000 heuristic studied fungal specimens and living cultures are deposited in search replicates with random taxa stepwise addition se- the herbarium of the National Museum of Natural Science of quences and TBR branch swapping with MAXTREES set to the Republic of China (TNM). autoincrease. Cultural descriptions and the species code follow Nobles For the Bayesian analysis, MrModeltest 2.3 (Nylander (1965), with amendments by Boidin and Lanquetin (1983). 2004) was used to determine the best-fit evolution model for Nobles’ code as detailed by Nakasone (1990) was adopted each dataset for Bayesian inference (BI). BI was calculated in this study. General methods of cultural studies have been with MrBayes3.1.2 with the K2 + I + G model (Ronquist and previously described by Wu (1996). Huelsenbeck 2003). The Markov chain Monte Carlo search was run with four chains for 500,000 generations, with trees Phylogenetic analyses sampled every 100 generations, and the average standard de- viation of split frequencies was below 0.05. According to Hall The sampling of representative orders and clades of (2004), the option of burnin was set to discard 10% trees. Agaricomycetes for the analyses of LSU nrDNA alone and a Branches that received bootstrap support for maximum parsi- combined dataset of LSU nrDNA and tef1-α sequences has mony (MP) and Bayesian posterior probabilities (BPPs) great- consulted some phylogenetic studies (Binder et al. 2005, er than or equal to 75% (MP) and 0.95 (BPP) were considered 2010, 2013; Hibbett 2006; Larsson 2007; Wu et al. 2007, as significantly supported, respectively. 2010; Li and Cui 2013). In addition, resemblance of the chief morphological characteristics of the presented new taxon was also considered for choosing the genera and species included Results in the analyses. The analyzed species and strains in both anal- yses, though not totally the same, nevertheless represented Phylogenetic analyses closely related taxa. The taxa and strains sampled are listed in Table 1. The LSU nrDNA amplification delimited by the primer pair The material for DNA isolation was taken from mycelia LR0R/LR5 yields PCR products of ca. 920 bp long. The final grown on malt extract agar (MEA). The samples were alignment of 38 sequences included 2171 positions. After ex- ground into fine powder with a pestle and mortar. DNA cluding ambiguous sites at both ends, 902 alignment sites was extracted from the fine powder using the Plant were used for the phylogenetic analysis. The MP analysis Genomic DNA Extraction Miniprep System (Viogene, revealed five most parsimonious trees [1330 steps, consisten- Taiwan), according to the manufacturer’s instructions. cy index (CI) = 0.4, retention index (RI) = 0.425]. Of the 902 The primer pair LR0R/LR5 was used to amplify the LSU included sites, 549 were constant, 106 were variable but par- nrDNA region. Part of tef1-α was amplified with primer simoniously uninformative, and 247 sites were parsimony in- pair EF1-1953R and Efdf (http://www2.clarku.edu/faculty/ formative. The best model for the combined LSU nrDNA dhibbett/Protocols_Folder/Primers/Primers.pdf). The dataset estimated and applied in the Bayesian analysis is: protocol for polymerase chain reaction (PCR) amplifica- K2+I+G,lsetnst=2,rates=invgamma;prsetstatefreqpr= tion followed Wu et al. (2007). PCR products were purified dirichlet (1,1,1,1). Bayesian analysis resulted in a similar to- with the PCR-M Clean Up system (Viogene). Nucleotide pology as MP analysis, with an average standard deviation sequences were determined from both strands using the (SD) of split frequencies = 0.023842 (BI). ABI PRISM BigDye Terminator Cycle Sequencing One of the most parsimonious trees is shown in Fig. 2.In Ready Reaction Kit on an ABI 3730 DNA sequencer total, 22 clades were recognized from the ingroup taxa: (Perkin Elmer, Applied Biosystems, Foster City, CA). Agaricales, , antrodia clade, Atheliales, All sequences included in the analyses were aligned and Boletales, Cantharellales, Corticiales, grifola clade, adjusted manually in BioEdit 7.0.4.1 (Hall 1999). The opti- gelatoporia clade, Gloeophyllales, Gomphales, mized sequence dataset (deposited in TreeBASE: http://purl. Hymenochaetales, Jaapiales, core polyporoid clade, phlebioid org/phylo/treebase/phylows/study/TB2:S21019?x-access- clade, phlebiella clade, residual polyporoid clade, Russulales, code=b08363e7ba49950c7d3d98d90465b73f&format=html) , Trechisporales, tyromyces clade, and was analyzed in PAUP* 4.0b10 (Swofford 2002)using Purpureocorticium. In the phylogenetic tree (Fig. 2), the two Mycol Progress (2018) 17:357–364 359

Table 1 Taxa used in this study, along with their strain/specimen Species Strain or specimen no. GenBank accession no. numbers and GenBank accession numbers. Newly generated LSU nrDNA tef1-α sequences are shown in bold Athelia arachnoidea CBS:418.72 GU187557 GU187672 Botryobasidium sp. Wu 1207-59 MF110286 LC270916 Calocera cornea AFTOL-ID 438 AY701526 AY881019 eludens JS27202 AF090878 – Ceraceomyces tessulatus KHL8474 AY586642 – Columnocystis abietina KHL12474 EU118619 – Coprinus comatus AFTOL-ID 626 AY635772 AY881026 Daedalea quercina Miettinen 12662 JX109855 JX109912 Dentocorticium sulphurellum FPL11801 AF393055 JN164903 Diplomitoporus crustulinus FP101824 AY333816 LC270917 Fomitopsis pinicola AFTOL-ID 770 AY684164 AY885152 Ganoderma applanatum Wei 5787a KF495011 KF494978 Gelatoporia sp. CHWC 1506-11 MF110287 LC270918 Gloeophyllum carbonarium FP97972 HM536054 HM536105 Grifola sordulenta TENN55054 AY645050 AY885154 Heliocybe sulcata IBUG 9930 HM536069 HM536115 geogenium PBM2382 AY631900 DQ059053 Hyphoderma setigerum CHWC 1209-9 MF110288 LC270919 Inonotus linteus MUCL 47139 GU462002 GU461936 Jaapia argillacea CBS:252.74 GU187581 GU187711 Lactarius lignyotus AFTOL-ID 681 AY631898 DQ435787 Leucogyrophana romellii CFMR:T-547 GU187586 GU187720 Lopharia cinerascens FP105043sp JN164813 JN164900 sordida Wu 0711-81 MF110289 LC270920 Phlebia radiata GEL5258 AJ406541 – Phlebiella californica Wu 9506-69 MF110290 LC270921 Punctularia strigosozonata Wu 0309-197 MF110291 LC270922 Purpureocorticium microsporum Wu 0806-27 MF110292 LC270923 Purpureocorticium microsporum Wu 9711-16 MF110293 LC270924 Ramaria rubella AFTOL-ID 724 AY645057 AY883435 Rhizochaete filamentosa FP105240 AY219393 – Rhizochaete rubescens Wu 0910-45 MF110294 LC270925 Rhizochaete violascens KHL11169 EU118612 – Steccherinum tenue 5356 JN710598 JN710733 Trametes suaveolens FP102529sp JN164807 JN164890 Trechispora alnicola CBS577.83 AY635768 DQ059052 Tyromyces sp. X1150 JN710603 JN710734 Veluticeps fimbriata L-10628-Sp HM536083 HM536127

sequences derived from this study were grouped together, uninformative, and 640 sites were parsimony informative. forming a distinct group (BS = 100%, BPP = 1). The best model for the combined LSU nrDNA and tef1-α The final alignment of 32 sequences from the combined partition was a GTR + I + G model. The BI analysis resulted dataset of LSU nrDNA and tef1-α included 1820 positions. in a similar topology, with an average SD of split frequen- The MP analysis revealed a most parsimonious tree (4214 cies = 0.028641 (BI). steps, CI = 0.358, RI = 0.334). Of the 1820 included sites, In the phylogeny (Fig. 3) inferred from the combined LSU 991 were constant, 189 were variable but parsimoniously nrDNA and tef1-α sequences, the two sequences derived from 360 Mycol Progress (2018) 17:357–364

Fig. 1 Purpureocorticium microsporum (holotype). a Basidiocarp section. b Subicular hyphae. c Basidioles. d Basidia. e Basidiospores

this study are very similar in base composition to the represen- Fruit body effused, adnate, membranaceous. Hymenial sur- tative sequences from Gloeophyllales and Jaapiales. The newly face white when fresh, turning purple in KOH, becoming pur- suggested clade served as a sister clade of Gloeophyllales and plish pink or pinkish purple after storage of several years, Jaapiales, which were grouped together with a weak support smooth. Hyphal system monomitic; hyphae nodose-septate. (BS = 98%, BPP = 1). The other most parsimonious tree is gen- Subiculum fairly uniform, composed of medullary layer, with erally identical to Fig. 3 in topology, but in some phylogenetic compact texture; hyphae colorless, interwoven, and with irreg- trees, Purpureocorticium was grouped together with grifola ular orientation, tortuous, indistinct, ± agglutinated, occasional- clade, gelatoporia clade, or tyromyces clade. ly irregularly swollen. Hymenial layer not clearly differentiated from subiculum, possibly thickening. Cystidia lacking. Basidia Taxonomy subclavate with a median constriction, or utriform, 4-sterigmate. Basidiospores ovate-ellipsoid, smooth, thin-walled, often with Purpureocorticium Sheng H. Wu, gen. nov. one oily drop, inamyloid, nondextrinoid, acyanophilous. MycoBank MB809432. Etymology: From purpureus (= purple) + Corticium,refer- Purpureocorticium microsporum Sheng H. Wu, sp. nov. ring to the hymenial surface color. (Fig. 1) Type species. Purpureocorticium microsporum. MycoBank MB809434. Diagnosis. Purpureocorticium is characterized by resupi- Etymology: From micro + spore, referring to the small nate basidiocarps with a smooth hymenial surface which turns basidiospores. purple in KOH, and becomes purplish after storage; micro- Holotype. TAIWAN. Taichung, Hoping Hsiang, between scopically having compact texture of subiculum, nodose- 24.5–25 km of Dasyueshan Forestry Road, alt. 1700 m, on septate thin-walled generative hyphae, absence of cystidia, angiosperm branch, leg. H.J. Chen & Y.T. Wang, 12 Jun 2008, basidia subclavate with a median constriction, or utriform, Wu 0806-27 (TNM F0022571). bearing ovate-ellipsoid small-sized basidiospores, which are Fruit body effused, adnate, membranaceous, 100–250 μm smooth and thin-walled, inamyloid, and nondextrinoid. thick in section. Hymenial surface white when fresh, turning Mycol Progress (2018) 17:357–364 361

Fig. 2 Maximum parsimony strict consensus tree illustrating the proportions higher than 50% and Bayesian posterior probabilities more phylogeny of Purpureocorticium and related species based on partial than 0.95, which are shown before and after the forward slash, LSU nrDNA sequences. Branches are labeled with parsimony bootstrap respectively purple in KOH, becoming purplish pink or pinkish purple hyphae nodose-septate. Subiculum fairly uniform, composed after storage of several years, smooth, cracked when dry; mar- of medullary layer, with compact texture; hyphae colorless, gin thinning, byssoid, glossy. Hyphal system monomitic; interwoven, and with irregular orientation, tortuous, indistinct, 362 Mycol Progress (2018) 17:357–364

Fig. 3 Maximum parsimony strict consensus tree illustrating the with parsimony bootstrap proportions higher than 50% and Bayesian phylogeny of Purpureocorticium and related species based on the posterior probabilities more than 0.95, which are shown before and combined LSU nrDNA + tef1-α sequence dataset. Branches are labeled after the forward slash, respectively

± agglutinated, occasionally irregularly swollen, variable in (2.7–)2.8–3.2(−3.3) × 2–2.2(−2.3) μm[Wu 0806-27:L= diam., 1–6 μm, thin-walled. Fairly big crystals crowded at 2.97 ± 0.10 μm, W = 2.08 ± 0.07 μm, Q = 1.43 (n = 30)], basal subiculum. Hymenial layer not clearly differentiated inamyloid, nondextrinoid, acyanophilous. from subiculum, may be thickening. Cystidia lacking. Additional specimens examined: CHINA. YUNNAN Basidia subclavate with a median constriction, or utriform, PROVINCE, Hsishuangbanna, Daduguang, 100°58′ E, 13–23 × 3.3–4.3 μm, 4-sterigmate. Basidiospores small, 22°24′ N, alt. 1300 m, on angiosperm branch, leg. S.H. Wu ovate-ellipsoid, smooth, thin-walled, often with one oily drop, & S.Z. Chen, 15 Aug 1997, Wu 9708-146 (TNM F0007817). Mycol Progress (2018) 17:357–364 363

TAIWAN. Chiayi County, Alishan Hsiang, Nanhsi Forest At an earlier stage of this study, Rhizochaete Gresl. et al. Road, 120°54′ E, 23°28′ N, alt. 1950 m, on angiosperm branch, was suspected as the possible genus to accommodate leg. S.H. Wu & J.Y. Tseng, 8 Dec, 1994, Wu 9412-26 (TNM P. microsporum. Rhizochaete was established by Greslebin F0020523). Taichung City, Hoping District, Anmashan, leg. et al. (2004) to accommodate some species belonging to the S.H. Wu & J.Y. Tseng, 121°00′ E, 24°16′ N, alt. 2250 m, on phlebioid clade (Binder et al. 2005, 2013;Wuetal.2010). rotten trunk, leg. S.H. Wu & H.J. Chan, 8 Nov 1977, Wu 9711- According to the protologue of Rhizochaete, its morphologies 16 (TNM F0009370). Taichung City, Hoping District, between are most similar to Phanerochaete, diagnostic by having a 24.5–25 km of Dasyueshan Forestry Road, alt. 1700 m, on smooth to tuberculate, pellicular hymenophore and hyphal angiosperm branch, leg. S.H. Wu, H.J. Chen & Y.T. Wang, cords that turn red or violet in KOH, monomitic hyphal sys- 19 Aug 2008, Wu 0808-45 (TNM F0027632); between 27 tem with simple or nodose-septate hyphae, cystidia, and small and 27.5 km of Dasyueshan Forestry Road, 120°57′ E, basidiospores. Purpureocorticium microsporum resembles 24°14′ N, alt. 1800 m, on angiosperm branch, leg. S.H. Wu, Rhizochaete because hymenophores of both turn violet in 17 Oct 2015, Wu 1510-10 (TNM F0029491). Nantou County, KOH, while Rhizochaete differs from the former in having Jenai Township, Entrance of Southern Tungyenshan, 121°06′ clavate to subcylindrical basidia, and its cystidia contain two E, 24°02′ N, alt. 1550 m, on fallen angiosperm trunk, leg. S.Z. types of encrustation. The phylogenetic analysis (Fig. 2)(also Chen, C.C. Chen & C.L. Wei, 28 Mar 2016, Chen 3167 (TNM Wu et al. 2010)showedthatRhizochaete belongs to the F0029996). phlebioid clade. Morphological features of Phanerochaete Distribution. China (Yunnan) and Taiwan. rubescens fit the generic concept of Rhizochaete, and its phy- Cultural description (Wu 0806-27). 1 wk. growth: Colony logenetic placement in Rhizochaete is supported in this study radius 19–25 mm. Matt white. Advancing zone even. Aerial (Figs. 2 and 3). A new combination of this species is thus mycelium absent. 2 wk. growth: Colony radius 39–47 mm. proposed herein. Matt white. Advancing zone even. Aerial mycelium absent or slightly pellicular, slightly zonate. 3 wk. growth: Plates partly Rhizochaete rubescens (Sheng H. Wu) Sheng H. Wu, covered. 6 wk. growth: Matt white. Aerial mycelium almost comb. nov. absent, slightly granular near plate margin. Odor slightly fra- Basionym: Phanerochaete rubescens Sheng H. Wu, grant. Agar unchanged. Fruiting with basidia and basidio- Mycological Research 102: 1131 (1998). spores. Hyphal system monomitic. Advancing hyphae color- less, (1–)1.5–4(−5) μm diam., thin-walled, sparsely branched, Purpureocorticium microsporum was also suspected to be nodose-septate. Aerial generative hyphae colorless, (1–)1.5– congeneric with Ceraceomyces violascens,asthe 4(−5) μm diam., thin-to slightly thick-walled, nodose-septate, hymenophore of the latter becomes violet-red in KOH, addi- moderately or sparsely branched. Submerged hyphae color- tionally sharing nodose-septate hyphae and absence of less, 1.5–5 μm diam., generally thin-walled, occasionally cystidia of both species. However, Ceraceomyces spp. bear slightly thick-walled, nodose-septate, moderately or sparsely fairly loose texture of subiculum and clavate basidia. The branched. Cuboid crystals present in the agar. phylogenetic analyses (Figs. 2 and 3) showed that three ana- Oxidase reactions.TAA:+++,6–11; +++, 18–30. GAA: lyzed Ceraceomyces species belong to different clades. +++, tr.; ++++, tr. TYA: − (slightly yellowish brown), 18–24; Ceraceomyces violascens and C. eludens belong to the − (slightly yellowish brown), 47–54. phlebioid clades, and the former is close to Rhizochaete in Species code. 2a, 3c, 7, 36, 40, 43, 48, 50, 54. affinity (Figs. 2 and 3; see also Wu et al. 2010), correspond- ing to the same color reaction of hymenophore in KOH. However, C. tessulatus (the generic type in these analyses) Discussion belongs to the Amylocorticiales (Figs. 2 and 3; see also Binder et al. 2010). Some genera bearing more or less similar morphological fea- Basidia of Purpureocorticium microsporum are small- to tures with Purpureocorticium are compared and their phylo- medium-sized, subclavate with a median constriction, or genetic placements are discussed below. utriform (Fig. 1), somewhat resemble those of Hyphodontia Purpureocorticium microsporum canbeconsideredassim- J. Erikss s.l., while the compact texture of the subiculum, as ilar to Phlebia Fr., due to the presence of dense texture of well as the more or less glued and indistinct subicular hyphae subiculum, nodose-septate hyphae, and small basidiospores. of Purpureocorticium, differ extremely from those of However, basidia of Phlebia are strictly clavate or subclavate, Hyphodontia. Hyphodontia belongs to the Hymenochaetales while those of Purpureocorticium are subclavate with a me- (Binder et al. 2005; Larsson 2007). The subclavate median- dian constriction or utriform. The phylogenetic analyses (Figs. constricted or utriform basidia of P. microsporum are similar 2 and 3)showedthatP. microsporum does not belong to the to those of Hyphoderma Wallr., but in the latter, basidia are phlebioid clade. larger. Moreover, Hyphoderma has larger basidiospores. The 364 Mycol Progress (2018) 17:357–364 phylogenetic analyses (Figs. 2 and 3) (also Binder et al. 2005) Boidin J, Lanquetin P (1983) Basidiomycètes Aphyllophoralés – showed that Hyphoderma (generic type: H. setigerum in this épithéloïdes étalés. Mycotaxon 16:461 499 Greslebin A, Nakasone KK, Rajchenberg M (2004) Rhizochaete, a new analyses) belongs to the residual polyporoid clade. genus of phanerochaetoid fungi. Mycologia 96:260–271 The present morphological study and analyses of LSU Hall TA (1999) BioEdit: a user-friendly biological sequence alignment nrDNA alone and the combined dataset of LSU nrDNA editor and analysis program for Windows 95/98/NT. Nucl Acids and tef1-α respectively support the generic status of Symp Ser 41:95–98 Purpureocorticium. However, morphological studies are not Hall BG (2004) Phylogenetic trees made easy: a how-to manual, 2nd edn. Sinauer Associates, Sunderland capable of evaluating the taxonomic placement of Hibbett DS (2006) A phylogenetic overview of the Agaricomycotina. Purpureocorticium, as its main diagnostic features are neither Mycologia 98:917–925. https://doi.org/10.3852/mycologia.98.6.917 peculiar nor suggestive of a possible relationship with known Hillis DM, Bull JJ (1993) An empirical test of bootstrapping as a method for orders or clades of the Agaricomycetes. The present phyloge- assessing confidence in phylogenetic analysis. Syst Biol 42:182–192 netic analyses (Figs. 2 and 3) showed that Purpureocorticium is Larsson KH (2007) Re-thinking the classification of corticioid fungi. Mycol Res 111:1040–1063 close to Jaapiales and Gloeophyllales and the core polyporoid Li HJ, Cui BK (2013) Taxonomy and phylogeny of the genus clade. Further study is needed to clarify the phylogenetic place- Megasporoporia and its related genera. Mycologia 105:368–383. ment of Purpureocorticium within Agaricomycetes. https://doi.org/10.3852/12-114 Nakasone KK (1990) Cultural studies and identification of wood- Acknowledgements This study was supported by the Ministry of Science inhabiting Corticiaceae and selected Hymenomycetes from North – and Technology of the ROC (grant no. 104-2621-B-178-001-MY3). America. Mycol Mem 15:1 412 Nobles MK (1965) Identification of cultures of wood-inhabiting Hymenomycetes. Can J Bot 43:1097–1139 Nylander JAA (2004) MrModeltest v2. Program distributed by the au- References thor. Evolutionary Biology Centre, Uppsala University Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic – Binder M, Hibbett DS, Larsson KH, Larsson E, Langer E, Langer G inference under mixed models. Bioinformatics 19:1572 1574. (2005) The phylogenetic distribution of resupinate forms across https://doi.org/10.1093/bioinformatics/btg180 the major clades of mushroom-forming fungi Swofford DL (2002) PAUP*. Phylogenetic analysis using parsimony (Homobasidiomycetes). Syst Biodivers 3:113–157. https://doi.org/ (*and other methods) version 4. Sinauer Associates, Sunderland 10.1017/S1477200005001623 Wu SH (1996) Studies on Gloeocystidiellum sensu lato Binder M, Larsson KH, Matheny PB, Hibbett DS (2010) (Basidiomycotina) in Taiwan. Mycotaxon 58:1–68 Amylocorticiales ord. nov. and Jaapiales ord. nov.: early diverging Wu SH, Wang DM, Tschen E (2007) Brunneocorticium pyriforme, a new clades of Agaricomycetidae dominated by corticioid forms. corticioid fungal genus and species belonging to the euagarics clade. Mycologia 102:865–880. https://doi.org/10.3852/09-288 Mycologia 99:302–309 Binder M, Justo A, Riley R, Salamov A, Lopez-Giraldez F, Sjökvist E, Wu SH, Nilsson HR, Chen CT, Yu SY, Hallenberg N (2010) The white- Copeland A, Foster B, Sun H, Larsson E, Larsson KH, Townsend J, rotting genus Phanerochaete is polyphyletic and distributed Grigoriev IV, Hibbett DS (2013) Phylogenetic and phylogenomic throughout the phleboid clade of the (Basidiomycota). overview of the Polyporales. Mycologia 105:1350–1373. https:// Fung Divers 42:107–118. https://doi.org/10.1007/s13225-010- doi.org/10.3852/13-003 0031-7