Hyphoderma pinicola sp. nov. of H. setigerum complex (Basidiomycota) from Yunnan, China

Abstract

Backgroud

Hyphoderma setigerum (Fr.) Donk is a white-rot wood-decaying corticoid fungal species. It occurs worldwide from tropical to temperate regions. However, taxonomic studies in recent decades showed that H. setigerum is a species complex with four separate species, before this study.

Results

Hyphoderma pinicola sp. nov. was collected on dead wood of Pinus yunnanensis Franch. in the temperate montane belt at 2200-2400 m altitudes, in Yunnan Province of China. Within the H. setigerum complex this new taxon is distinguished by having 2-sterigmate basidia, long basidiospores, and nearly naked septocystidia. A description and illustrations of this new species are provided, along with a key to five species of the H. setigerum complex. Phylogenetic reconstruction based on 5.8S-ITS2 sequences indicated that H. pinicola belongs to the H. setigerum complex and has a separate position within the clade including H. subsetigerum and H. setigerum s.s. Bayesian inference of phylogeny based on two datasets, ITS and 28S nuclear ribosomal DNA sequences, confirmed the independent status of H. pinicola.

Conclusion

Morphological and phylogenetic studies showed that H. pinicola represents a fifth species of H. setigerum complex.

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RESEARCH Open Access

Hyphoderma pinicola sp. nov. of H. setigerum complex (Basidiomycota) from Yunnan, China

Eugene Yurchenko1 and Sheng-Hua Wu2*

Abstract

Backgroud: Hyphoderma setigerum (Fr.) Donk is a white-rot wood-decaying corticoid fungal species. It occurs worldwide from tropical to temperate regions. However, taxonomic studies in recent decades showed that H. setigerum is a species complex with four separate species, before this study.

Results: Hyphoderma pinicola sp. nov. was collected on dead wood of Pinus yunnanensis Franch. in the temperate montane belt at 22002400 m altitudes, in Yunnan Province of China. Within the H. setigerum complex this new taxon is distinguished by having 2-sterigmate basidia, long basidiospores, and nearly naked septocystidia. A description and illustrations of this new species are provided, along with a key to five species of the H. setigerum complex. Phylogenetic reconstruction based on 5.8S-ITS2 sequences indicated that H. pinicola belongs to the H. setigerum complex and has a separate position within the clade including H. subsetigerum and H. setigerum s.s. Bayesian inference of phylogeny based on two datasets, ITS and 28S nuclear ribosomal DNA sequences, confirmed the independent status of H. pinicola.

Conclusion: Morphological and phylogenetic studies showed that H. pinicola represents a fifth species ofH. setigerum complex.

Keywords: Corticioid fungi; Meruliaceae; Polyporales; Taxonomy

Background

Hyphoderma Wallr. is the largest genus of Basidiomycota with resupinate non-poroid basidiomata. Currently, 103 species are recognized under Hyphoderma in Index Fungorum (Kirk, 2014). According to Dai (2011), 24 species of Hyphoderma s.l. (including Mutatoderma (Parmasto)C.E. Gmez and Peniophorella P. Karst.) were listed in the mycobiota of China. Hyphoderma setigerum (Fr.)Donk occurs worldwide from tropical to temperate regions. However, taxonomic studies in recent decades showed that H. setigerum is a species complex with four species. A new species belonging to the H. setigerum complex is described in the present paper. This new taxon is based on specimens collected in 2001 on dead branches of Pinus yunnanensis Franch., from the temperate montane belt of Yunnan Province, China.

Methods

Reference herbarium materials and study of the morphologyThe specimens studied of this new species are deposited in the herbaria TNM and MSK (herbarium acronyms follow Index Herbariorum, http://sweetgum.nybg.org/ih

Web End =http://sweetgum.nybg.org/ih ). The isolate is kept in the culture collection of TNM.

Description of macromorphology is based on dry basidiomata. Microscopic measurements and drawings were made from material mounted in 3% KOH water solution. Melzers reagent was used to examine amyloidity or dextrinoidity of spore walls, but also to study crystalline incrustations on hyphae, or in the hymenium. Cyanophily of the spore wall was tested in cotton blue-lactophenol solution. To determine average spore size, 30 randomly selected spores from a squash basidioma preparation were measured.

DNA extraction, amplification, and sequencingNuclear ribosomal DNA sequences of Hyphoderma were analyzed in addition to morphological study. The material for DNA isolation was mycelium grown in pure culture

* Correspondence: mailto:[email protected]

Web End [email protected]

2Department of Biology, National Museum of Natural Science, Taichung 404, TaiwanFull list of author information is available at the end of the article

2014 Yurchenko and Wu; licensee Springer. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0

Web End =http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly credited.

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(Wu 010836), and basidioma pieces taken from herbarium specimens (TNM F13635, TNM F13637). Both kinds of material were homogenized in liquid nitrogen. DNA was extracted with Plant Genomic DNA Extraction Mini-prep Kit (Viogene, Taiwan), according to manufacturers protocol. Primer pair ITS1/ITS4 was used for amplification of internal transcribed spacer region, including ITS1,5.8S, and ITS2, under PCR conditions, described in White et al. (1990). The DNA fragment of ribosomal large subunit gene (28S), was amplified with primers LR0R/LR5 (Moncalvo et al., 2000), following PCR settings as described in Wu et al. (2007). Amplifications were run on a Mastercycler Gradient 5331 thermal cycler (Eppendorf, Germany). Amplification products were purified with a PCR-M Clean Up kit (Viogene) and sequenced with an ABI PRISM BigDye Terminator Cycle Sequencing Ready Reaction kit on ABI 3730 DNA sequencer (Applied Biosystems, USA). The resulting sequences were deposited in NCBI GenBank (Table 1).

Sequence alignment and reconstruction of phylogeny The datasets were composed of the sequences obtained in this study and taken from GenBank (Table 1), with the aim to elucidate phylogenetic distances between the new taxon and other Hyphoderma species. Two species of Mutatoderma, earlier known under Hyphoderma, were added to both 5.8S-ITS2 and 28S datasets. From about 50 ITS sequences of H. setigerum available in GenBank, we selected those representing the main clades within this species complex (Nilsson et al., 2003). The selected taxa in the ingroup belong to residual polyporoid clade of Agaricomycetes (Binder et al., 2013). Phanerochaete sordida (P. Karst.) J. Erikss. & Ryvarden, a member of phlebioid clade, was selected as an out-group in both datasets.

Sequences were aligned on-line in MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server

Web End =http:// http://mafft.cbrc.jp/alignment/server

Web End =mafft.cbrc.jp/alignment/server ), using E-INS-i strategy for ITS and G-INS-i for 28S (Katoh et al., 2009). Before alignment, ITS1 and 28S segments were cut from the sequences where needed. Sequences that were too short were removed from the datasets after preliminary alignments. Final datasets were edited manually in MEGAv. 3.1 (Kumar et al., 2004). Ready data matrices, together with resultant phylograms, were deposited in TreeBase (http://purl.org/phylo/treebase/phylows/study/TB2:S16046

Web End =http://purl.org/phylo/treebase/phylows/study/TB2:S16046). The best-fit models of nucleotide evolution were estimated by MrModeltest v. 2.3 (Nylander, 2004), with Akaike Information Criterion as a relative quality measure of the model (Posada and Buckley, 2004). The input file for MrModeltest was generated in PAUP* v. 4.0b10 (Swofford, 2002).

Bayesian analysis of phylogeny was performed in MrBayesv. 3.2.1 (Ronquist and Huelsenbeck, 2003). Both 5.8SITS2 and 28S datasets were individually analyzed in two independent runs, each with four MC3 chains running for

1 million generations, with tree and parameter sampling every 500 generations. Burn-in was as default setting (discarding 25% of samples). The datamatrix of 5.8S-ITS2 was analyzed with different sets of parameters for two partitions, 5.8S and ITS2, according to best-fit models. FigTreev. 1.3.1 was used to view and capture the resultant phylo-grams and CorelDraw v. 9 for drawing the images.

Results

A key to species of Hyphoderma setigerum complex1. Conspicuous capitate cystidia with naked, bulbous apices present, up to 14 m wide apically, coarsely encrusted below the apex H. nudicephalum1 Cystidia more or less cylindrical, seldom with naked

capitate apex, but the apical bulb not exceeding 8 m in width 22. Basidia 2-sterigmate 32. Basidia 4-sterigmate 43. Basidiospores reaching 1517.5 m long, often suballantoid; basidia 2532 m long; septocystidia naked or weakly encrusted, often with thin (up to0.5 m) wall H. pinicola3. Basidiospores 1012 m long, only a little depressed adaxially; basidia 1725 m long; septocystidia heavily encrusted, typically with thick (up to 1 m) wall H. bisetigerum4. Basidiospores 68 2.83.2 m; known from Taiwan, Japan, China H. subsetigerum4. Basidiospores 710(14) 35(6) m; worldwide H. setigerum

Hyphoderma pinicolaYurchenko & Sheng H. Wu, sp. nov. Figures 1 and 2.

MycoBank 804684

HolotypeChina. Yunnan Prov., Hoching County, Sungkuei, alt. 2200 m, on dead corticated branch of Pinus yunnanensis, coll. S.H. Wu & S.Z. Chen, 1 Aug 2001, Wu 010832 (TNM F13637; isotype in MSK).

EtymologySpecific epithet refers to the host preference (Pinus).

DiagnosisBasidioma thin; hymenial surface chalky white, warted; septocystidia scattered, almost naked and often thin-

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Table 1 Taxa used in this study, along with their specimen /strain numbers, locality information and GenBank accession numbers

Species name Isolate / Specimen voucher Country of origin GenBank accession no. for nrDNA5.8S-ITS2 28S

Hyphoderma cremeoalbum / NH 11538 (GB) Turkey DQ677492 DQ677492

Hyphoderma definitum / GEL 2898 AJ406509 Hyphoderma definitum / FCUG 2426 Russia (Krasnodar krai) AJ534293 Hyphoderma definitum / NH 12266 (GB) Russia DQ677493 DQ677493 Hyphoderma granuliferum / KHL 12561 (O) Costa Rica JN710545 JN710545 Hyphoderma incrustatum KHL 6685 / AY586668 Hyphoderma litschaueri / NH 7603 (GB) Canada DQ677496 DQ677496 Hyphoderma litschaueri / CFMR:DLL2011-050 USA KJ140573 Hyphoderma macaronesicum E09/57-9 / TFC:Mic.15981 Canary Islands HE577027 Hyphoderma medioburiense / NH 10950 (GB) Spain DQ677497 DQ677497 Hyphoderma nemorale EM 2793 / AY586669 Hyphoderma nudicephalum / TMIC 33708 Japan AJ534264 Hyphoderma nudicephalum / TMIC 50048 Japan AJ534265 Hyphoderma nudicephalum / FCUG 2949 Japan AJ534266 Hyphoderma nudicephalum / TMIC 30479 Japan AJ534267 Hyphoderma nudicephalum Wu 9508225 / China AJ534268 Hyphoderma nudicephalum Wu 930729 / Taiwan AJ534269 Hyphoderma nudicephalum / TMIC 50049 Japan AJ534270 Hyphoderma nudicephalum / GEL 4727 AJ406510 Hyphoderma obtusiforme / KHL 1464 JN572909 Hyphoderma obtusiforme / KHL 11105 JN572910 Hyphoderma obtusum JS 17804 / AY586670 Hyphoderma occidentale KHL 8469G / AY586674 Hyphoderma occidentale / KHL 8477 (GB) Sweden DQ677499 DQ677499 Hyphoderma pinicola* Wu 010830 / TNM F13635 China KJ885179 KJ885180 Hyphoderma pinicola* Wu 010832 / TNM F13637 China KJ885181 KJ885182 Hyphoderma pinicola* Wu 010836 / TNM F13643 China KC928278 KC928279 Hyphoderma prosopidis E09/58-9 / ARIZ:H.H. Burdsall 8479 USA HE577029 Hyphoderma roseocremeum NH 10545 / AY586672 Hyphoderma setigerum / GEL4001 AJ406511 Hyphoderma setigerum / FCUG 1688 Finland AJ534272 Hyphoderma setigerum / FCUG 1200 Norway AJ534273 Hyphoderma setigerum KHL 8544 / AY586673 Hyphoderma setigerum FCUG 1264 / NH 8544 (GB) Sweden FN907905 FN907905 Hyphoderma setigerum FCUG 2499 / Argentina GQ409515 Hyphoderma setigerum FCUG 2530 / Argentina GQ409516 Hyphoderma setigerum FCUG 3038 / South Africa GQ409517 Hyphoderma setigerum FCUG 3037 / South Africa GQ409518 Hyphoderma setigerum CFMR FP101976 / USA GQ409519 Hyphoderma setigerum CFMR HHB9443 / USA GQ409520 Hyphoderma subtestaceum CFMR HHB11620 / USA GQ409521 Hyphoderma subtestaceum CFMR MJL1536 / USA GQ409522

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Table 1 Taxa used in this study, along with their specimen /strain numbers, locality information and GenBank accession numbers (Continued)

Hyphoderma subsetigerum Wu 9508155 / China AJ534275 Hyphoderma subsetigerum / TMIC 33552 Japan AJ534276 Hyphoderma subsetigerum Wu 930418 / Taiwan AJ534277 Hyphoderma subsetigerum Wu 920215 / Taiwan AJ534278 Hyphoderma transiens / NH 12304 (GB) Sweden DQ677504 DQ677504 Mutatoderma heterocystidium** / NH 7574 (GB) Canada DQ677495 DQ677495 Mutatoderma mutatum / NH 12026 (GB) Russia DQ677498 DQ677498 Phanerochaete sordida / KHL 12054 (GB) Norway EU118653 EU118653 *Data in bold indicate the sequences obtained in this study.

**In GenBank under the name Hyphoderma heterocystidiatum (Burt) Donk.

walled; basidia with two large sterigmata; basidiospores (10)1316(17.5) m long, predominantly suballantoid.

DescriptionBasidiomata effused, membranaceous, 60150 m thick. Hymenial surface chalky white, minutely warted (34 warts/mm), between warts from the beginning minutely porulose, then continuous, minutely cracking with age. Margin zone up to 1.5 mm broad, concolorous with the main hymenial surface, abrupt or usually diffuse or slightly fibrillose. Hyphal system monomitic, all hyphae colorless, thin-walled, clamped at all primary septa. Subiculum thin, consisting of thin, fairly compact layer of more or less horizontal hyphae next to the substratum, and loose intermediate layer composed of variously oriented hyphae. Subicular hyphae moderately branched, 34.5 m diam, naked to richly encrusted. Subhymenium not distinctly differentiated from subiculum. Subhymenial hyphae moderately to richly branched, 2.5 3 m diam, moderately to richly encrusted (most incrustation dissolving in KOH solution). Cystidia of two kinds: (1) septocystidia scattered, projecting, irregularly cylindrical, straight to strongly twisted, simple or branched, with (2)46(8) and more predominantly clamped septa, apically near cylindrical or subcapitate, 65180 m long, up to 711 m broad in swellings and

the least 3.5 m broad in constrictions, colorless, moderately thin-walled (walls up to 0.5 m thick) to seldom thick-walled (up to 2 m thick), naked or slightly encrusted, especially in basal part; (2) aseptate cystidia may be found in younger portions, slightly protruding, subcylindrical, 3545 (4.5)67 m, thin-walled, naked. Basidia subcylindrical or narrowly utriform, 25 28(32) 56.5 m, colorless, thin-walled, naked, with two sterigmata measuring 6.57.5(9) 11.5(1.8) m. Spores cylindrical to allantoid, rarely slightly sigmoid, (10)1316(17.5) (3.5)44.5(6) m (rarely up to19.5 m long), colorless, thin-walled, smooth, negative in Melzers reagent, acyanophilous, with a short rounded apiculus.

Additional specimens examinedCHINA. Yunnan Prov., Hoching County, Sungkuei, alt. 2200 m, on dead twig of Pinus yunnanensis, coll. S.H. Wu & S.Z. Chen, 1 Aug 2001, Wu 010830 (TNM F13635); Chuhsiung, Tzuhsishan, alt. 2400 m, on dead twig of Pinus sp., coll. S.H. Wu & S.Z. Chen, 2 Aug 2001, Wu 010836 (TNM F13643).

DistributionThe species is so far known only from the temperate belt in northwest part of Yunnan Province, China.

Figure 1 Macromorphology of Hyphoderma pinicola (TNM F13635). A, Basidioma in central part (holotype); B, Basidioma in marginal part

(holotype); C, Thin, minutely porulose basidioma. Scale bars = 1 mm.

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Figure 2 Micromorphology of Hyphoderma pinicola (TNM F13637). A, B, Vertical sections through basidioma; C, Naked and encrusted

subicular hyphae; D, Two straight and one distorted septocystidia; E, Basidia; F, Normal basidiospores; G, Basidiospore with a false septum.

Scale bars: for A = 100 m; for BG = 10 m.

RemarksAnother bi-sterigmate species from the H. setigerum complex, H. bisetigerum Boidin & Gilles, was described from Madagascar (Boidin and Gilles, 2003). However, this species differs from H. pinicola in bearing shorter basidia, shorter basidiospores, and heavily encrusted septocystidia (see key). In the global survey of the H. setigerum complex (Nilsson et al., 2003), the largest spores (1214.5 4.5 5 m) were found in the material from Greenland; these however are shorter than in H. pinicola. According to a detailed morphological study of H. setigerum in Belarus and northwest Russia (Yurchenko and Zmitrovich, 2001), the largest spores do not exceed 14 m long, with averages from 7.3 to 11.2 m. The average spore size inH. pinicola specimens was 15.1 4.3 m (holotype),14.4 4.3 m (TNM F13635), and 14.1 4.5 m (TNM F13643). The preference of growing on coniferous wood is rare for H. setigerum s.l., and it is a distinctive ecological feature of H. pinicola.

Molecular phylogenyThe 5.8S-ITS2 dataset analyzed by Bayesian analysis included 356 positions together with introduced gaps (99 positions in partial 5.8S, 86 of which were constant; 257 positions in complete ITS2, 66 of which were constant). MrModeltest suggested GTR + I + G as the best-fit model of nucleotide evolution for 5.8S + ITS2, K80 for partial 5.8S, and GTR + I + G for the whole dataset. The aligned datamatrix of partial 28S sequences included 882 positions, of which 820 were constant. The best-fit model of nucleotide evolution suggested for it by MrModeltest was GTR + I + G.

Both phylograms generated using Bayesian approach (Figures 3 and 4) confirmed that H. pinicola belongs to the genus Hyphoderma. According to the phylogram based on 5.8S-ITS2 (Figure 3), three specimens of H. pinicola constitute a well-supported clade (with Bayesian posterior probability value, PP = 1.00) within the H. setigerum complex. The phylogram based on partial 28S (Figure 4)

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Figure 3 Phylogram obtained to reveal the phylogenetic position of Hyphoderma pinicola via Bayesian analysis (5.8S-ITS2 dataset).

Numbers above branches denote Bayesian posterior probability (PP) value (if PP 0.50). Thick branches have PP 0.99. Scale bar for branch

length indicates the number of nucleotide substitutions per site.

Figure 4 Phylogram obtained to reveal the phylogenetic position of Hyphoderma pinicola via Bayesian analysis (28S dataset). Numbers

above branches denote Bayesian posterior probability (PP) value (if PP 0.50). Thick branches have PP 0.99. Scale bar for branch length

indicates the number of nucleotide substitutions per site.

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demonstrates that three specimens of H. pinicola also constitute a separate clade (PP = 0.88). They belong to a highly supported clade (PP = 1.00) together withH. nudicephalum Gilb. & M. Blackw. A high degree of molecular divergence in ITS and 28S sequences supports specific status of H. pinicola.

Discussion

A test of the application of molecular phylogeny in taxonomy is whether there is consistency between classification schemes based on molecular characters and morphological traits. Our results demonstrate certain congruence between morphological study and ribosomal gene sequence analyses. The phylogram inferred from5.8S-ITS2 shows H. pinicola as a member of H. setigerum complex. All specimens of the H. setigerum complex constitute a separate, strongly supported clade. However, the H. setigerum complex appeared to be not monophyletic according to 28S-based phylogram. This phylo-gram shows that H. pinicola and H. nudicephalum are separate from H. setigerum. Basal branching order in 28S-based phylogram is not strongly supported by Bayesian posterior probability values, indicating that phylogenetic differentiation within the genus is not deep.

The previous comprehensive phylogenetic study of theH. setigerum complex (Nilson et al., 2003) provided molecular grounds for recognition of H. nudicephalum and H. subsetigerum Sheng H. Wu. These species were defined morphologically: by characteristic cystidia inH. nudicephalum (Gilbertson and Blackwell, 1988) and small spores in H. subsetigerum (Wu, 1997). However, more than ten clades and subgroups were recognized and considered at species rank within this complex. As Nilsson et al. (2003: 651) noted, H. subsetigerum known from Asia (subgroup 7B) and H. setigerum s.s. from Northern Europe (subgroup 7B, AJ534272, AJ534273) constitute two different species because of incompatibility, geographic isolation and differences in spore size. Hyphoderma pinicola constitutes the third species in this small clade, possessing distinctive features in morphology, host preference, and distribution. The clade composed of long-spored specimens from Greenland is not only geographically, but also phylogenetically far from H. subsetigerum clade (see Nilsson et al., 2003, Figure 1). This analysis revealed that the sister group to the H. subsetigerum clade(i.e. the assemblage of H. pinicola, H. setigerum s.s., andH. subsetigerum) includes specimens of H. setigerum s.l. from Argentina (GQ409515, GQ409516) and South Africa (GQ409517, GQ409518). Both of these clades can be recognized as two independent species. Thus, after phylo-genetic reconstructions, presumably new species from geographically separated areas appeared to be discriminated in highly supported clades on the basis of the 5.8SITS2 dataset only.

Two specimens of H. subtestaceum (Litsch.) Donk (GQ409521, GQ409522) were included in our phylogenetic study. According to Mycobank, H. subtestaceum is a synonym of H. setigerum. However, in the phylogram based on 5.8S-ITS2 sequences (Figure 3), these two specimens constitute a highly supported clade (PP = 1.00) separate from the clade that includes the H. setigerum complex. These results suggest that specimens from North America, called H. subtestaceum, belong to an independent species.

Conclusion

Hyphoderma pinicola, known from Yunnan Province, China, and collected on dead wood of Pinus yunnanensis, represents the fifth named species of the H. setigerum complex.

Competing interestsThe authors declare that they have no competing interests.

Authors contributionsEY prepared description and illustration for the new taxon, performed phylogenetic analysis, and prepared the manuscript. SHW collected the specimens and provided preliminary identification, suggested the study topic, guided molecular lab work, examined the taxa used for phylogenetic study, and revised and prepared the manuscript for submission. Both authors read and approved the final manuscript.

AcknowledgementsThis study was supported financially by Ministry of Science and Technology of ROC (grants NSC 98-2621-B-178-002-MY3 and NSC 101-2621-B-178-001-MY3). We are grateful to S.Z. Chen (TNM) for the help in managing herbarium specimens for this study and to Y.P. Chen for performing DNA extraction and amplification.

Author details

1Department of Biotechnology, Paleski State University, Dnyaprouskai flatylii str. 23, BY-225710 Pinsk, Belarus. 2Department of Biology, National Museum of Natural Science, Taichung 404, Taiwan.

Received: 20 July 2014 Accepted: 20 September 2014

ReferencesBinder M, Justo A, Riley R, Salamov A, Lopez-Giraldez F, Sjkvist E, Copeland A,

Foster B, Sun H, Larsson E, Larsson K-H, Townsend J, Grigoriev IV, Hibbett DS (2013) Phylogenetic and phylogenomic overview of the Polyporales. Mycologia 105:13501373, doi:10.3852/13-003Boidin J, Gilles G (2003) Homobasidiomyctes Aphyllophorales non pors basides dominantes 2 (3) strigmates. Bull Soc Mycol Fr 119:117Dai Y-C (2011) A revised checklist of corticioid and hydnoid fungi in China for

2010. Mycoscience 52:6979, doi:10.1007/s10267-010-0068-1Gilbertson RL, Blackwell M (1988) Some new or unusual corticioid fungi from the

Gulf Coast Region. Mycotaxon 33:375386Katoh K, Asimenos G, Toh H (2009) Multiple alignment of DNA sequences with MAFFT. Methods Mol Biol 537:3964, doi:10.1007/978-1-59745-251-9_3Kirk P (ed) (2014) Index Fungorum. http://www.indexfungorum.org/Names/Names.asp

Web End =http://www.indexfungorum.org/Names/

http://www.indexfungorum.org/Names/Names.asp

Web End =Names.asp . Retrieved 29 Jun 2014Kumar S, Tamura K, Nei M (2004) MEGA3: integrated software for Molecular

Evolutionary Genetics Analysis and sequence alignment. Brief Bioinform 5:150163, doi:10.1093/bib/5.2.150Moncalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R (2000) Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences. Syst Biol 49:278305, doi:10.1093/sysbio/49.2.278

Yurchenko and Wu Botanical Studies 2014, 55:71 Page 8 of 8

http://www.as-botanicalstudies.com/content/55/1/71

Nilsson RH, Hallenberg N, Nordn B, Maekawa N, Wu S-H (2003) Phylogeography of Hyphoderma setigerum (Basidiomycota) in the Northern Hemisphere. Mycol Res 107:645652, doi:10.1017/S0953756203007925

Nylander JAA (2004) MrModeltest v2. Program distributed by the author.

Evolutionary Biology Center, Uppsala University, UppsalaPosada D, Buckley TR (2004) Model selection and model averaging in phylogenetics: advantages of the Akaike Information Criterion and Bayesian approaches over likelihood ratio tests. Syst Biol 53:793808, doi:10.1080/ 10635150490522304Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19:15721574, doi:10.1093/ bioinformatics/btg180Swofford DL (2002) PAUP*: phylogenetic analysis using parsimony (*and other methods), vol 4. Sinauer Associates, SunderlandWhite TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Innis MA, Gelfand DH, Sninsky JJ, White TJ (eds) PCR protocols: A guide to methods and applications. Academic Press, New York, pp 315322Wu SH (1997) New species of Hyphoderma from Taiwan. Mycologia 89:132140, doi:10.2307/3761182Wu SH, Wang DM, Tschen E (2007) Brunneocorticium pyriforme, a new corticioid fungal genus and species belonging to the euagarics clade. Mycologia 99:302309, doi:10.3852/mycologia.99.2.302Yurchenko EO, Zmitrovich IV (2001) Variability of Hyphoderma setigerum

(Corticiaceae s.l., Basidiomycetes) in Belarus and northwest Russia. Mycotaxon 78:423434

doi:10.1186/s40529-014-0071-5Cite this article as: Yurchenko and Wu: Hyphoderma pinicola sp. nov. ofH. setigerum complex (Basidiomycota) from Yunnan, China. Botanical Studies 2014 55:71.

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