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Hydnophanerochaete and Odontoefibula, two new genera of phanerochaetoid fungi (Polyporales, Basidiomycota) from East Asia
expand article infoChe-Chih Chen, Sheng-Hua Wu§, Chi-Yu Chen
‡ National Chung Hsing University, Taichung, Taiwan
§ National Museum of Natural Science, Taichung, Taiwan
Open Access

Abstract

Two new genera with phylogenetic affinities to Phanerochaete s.l. are presented, namely Hydnophanerochaete and Odontoefibula. The generic type of Hydnophanerochaete is Phanerochaete odontoidea. Odontoefibula is established based on a new species: O. orientalis (generic type). Both genera have effused basidiocarps with odontioid hymenial surface, simple-septate generative hyphae, cystidia lacking, clavate basidia and ellipsoid basidiospores that are smooth, thin-walled, inamyloid, non-dextrinoid and acyanophilous. Hydnophanerochaete is additionally characterised by a compact texture in the subiculum with thick-walled generative hyphae and quasi-binding hyphae. Odontoefibula has a dense texture of subiculum with thin- to slightly thick-walled hyphae and further a dark reddish reaction of basidiocarps when treated with KOH. Multi-marker phylogenetic analyses based on sequences, inferred from the ITS+nuc 28S+rpb1+rpb2+tef1 dataset, indicate that Hydnophanerochaete and Odontoefibula are placed in the Meruliaceae and Donkia clades of Phanerochaetaceae, respectively. Phanerochaete subodontoidea is a synonym of P. odontoidea, according to morphological and molecular evidence.

Keywords

Meruliaceae, multi-marker phylogeny, new species, Phanerochaetaceae, phlebioid clade

Introduction

The genus Phanerochaete P. Karst., typified by P. alnea (Fr.) P. Karst., belongs to Polyporales Gäum of the Basidiomycota R.T. Moore and is one of the largest genera of corticoid fungi, including over 150 names according to Index Fungorum (http://www.indexfungorum.org/). Basidiocarps are typically membranaceous, effused, with various hymenial surfaces (i.e. smooth, tuberculate, odontioid, hydnoid, merulioid or poroid). Microscopically, Phanerochaete has a monomitic hyphal system, ordinarily simple-septate generative hyphae (rare clamp connections can be found in the subiculum), ellipsoid to cylindrical thin-walled basidiospores and clavate basidia. Phanerochaete is widespread and grows on diverse woody substrates (i.e. twigs and branches or trunks of angiosperms or gymnosperms), causing a white rot. Phanerochaete s.l. has attracted increasing study interest due to its abundant taxonomic diversity and potential applications in the field of biodegradation and bioconversion (Sánchez 2009).

Phanerochaete was traditionally treated as a genus in the broad sense (Eriksson et al. 1978; Burdsall 1985; Wu 1990). In recent years, Phanerochaete has been shown to be a polyphyletic group with members distributed throughout the phlebioid clade of Polyporales (De Koker et al. 2003; Wu et al. 2010; Floudas and Hibbett 2015; Miettinen et al. 2016), which was recently recognised as three families: Phanerochaetaceae Jülich, Irpicaceae Spirin & Zmitr and Meruliaceae Rea (Justo et al. 2017). Based on the combined morphological and molecular approaches, many studies have been conducted to revise the generic concept of Phanerochaete s.l. Some segregated genera have been recovered or proposed, e.g. Efibula Sheng H. Wu, Hydnophlebia Parmasto, Phaeophlebiopsis Floudas & Hibbett, Phlebiopsis Jülich, Rhizochaete Gresl., Nakasone & Rajchenb. and Scopuloides (Massee) Höhn. & Litsch. (Wu 1990; Greslebin et al. 2004; Wu et al. 2010; Floudas and Hibbett 2015).

Phanerochaete odontoidea Sheng H. Wu and P. subodontoidea Sheng H. Wu were described from Taiwan (Wu 2000). Both species have ceraceous basidiocarps with odontioid to hydnoid hymenial surface, compact subiculum, but no cystidia. These species have been shown to be phylogenetically far from the core Phanerochaete clade (Wu et al. 2010; Ghobad-Nejhad et al. 2015; Wu et al. 2018) and were placed by Justo et al. (2017) in Meruliaceae. In this study, we evaluate the generic placement of P. odontoidea and P. subodontoidea, as well as morphologically similar species. To accommodate our target taxa, we found it necessary to introduce two new genera placed within Meruliaceae and Phanerochaetaceae, respectively.

When Phanerochaete odontoidea and P. subodontoidea were described, they were separated by basidiospore width (Wu 2000). After 2000, we have accumulated more collections identified as P. odontoidea and P. subodontoidea from China, Japan, Taiwan and Vietnam. To better reflect their morphological variations, this study provides updated morphological and molecular evidence for revising their species concepts.

Materials and methods

Morphological studies

The specimens used for illustrations and descriptions are deposited at the herbarium of National Museum of Natural Science of ROC (TNM, acronym according to Index Herbariorum; http://sweetgum.nybg.org/science/ih/). Free-hand thin sections of basidiocarps were mounted in three mounting media for microscopic studies: 5% (w/v) KOH with 1% (w/v) phloxine was used for observation and measurements; Melzer’s reagent (IKI) was utilised to check amyloidity and dextrinoidity; and Cotton Blue (CB, Fluka 61335) was employed to determine cyanophily. Sections were studied with a Leica DM2500 (Leica, Wetzlar) microscope. Drawings were done with the aid of a drawing tube. We followed the method for measurements of microscopic characters by Wu (1990). The abbreviations below were used when presenting statistical measurements of basidiospores: L = mean basidiospore length, W = mean basidiospore width, Q = variation in L/W ratio, n = number of measured spores. The terminology of microscopic characters follows Wu (1990).

DNA extraction and sequencing

Dried specimens or mycelia grown on MEA were used for isolating genomic DNA. The material was first fragmented into a fine powder with the aid of liquid nitrogen and a TissueLyser II (Qiagen, Hilden, Germany). DNA was obtained using the Plant Genomic DNA Extraction Miniprep System (Viogene-Biotek Corp., New Taipei, Taiwan) based on the manufacturer’s instructions. Five genetic markers were amplified in this study: nuc rDNA ITS1-5.8S-ITS2 (ITS) using primer pair ITS1/ITS4 (White et al. 1990); D1-D2 domains of nuc 28S rDNA (nuc 28S) using primer pair LR0R/LR5 (http://www2.clarku.edu/faculty/dhibbett/Protocols_Folder/Primers/Primers.pdf); RNA polymerase II largest subunit (rpb1) using primer pair RPB1-Af/RPB1-Cr (Stiller and Hall 1997; Matheny et al. 2002) or alternative primers RPB1- 2f, RPB1-2.1f, RPB1-2.2f and RPB1-2.1r (Frøslev et al. 2005); RNA polymerase II second largest subunit (rpb2) using primer pair RPB2-f5F/RPB2-b7.1R (Liu et al. 1999; Matheny 2005); and translation elongation factor 1-α (tef1) using primer pair EF1-983F/EF1-2212R (Rehner and Buckley 2005). The PCR protocols for ITS and nuc 28S gene regions were as follows: initial denaturation at 95 °C for 5 min, followed by 40 cycles at 94 °C for 45 s, 53 °C for ITS and 50 °C for nuc 28S for 45 s and 72 °C for 45 s and a final extension of 72 °C for 10 min. The PCR protocols for rpb1, rpb2 and tef1 include initial denaturation at 94 °C for 2 min, followed by 35 cycles at 94 °C for 40 s, 60 °C for 40 s and 72 °C for 2 min and a final extension of 72 °C for 10 min. PCR products were purified and sequenced by the MB Mission Biotech Company (Taipei, Taiwan). Newly obtained sequences for each of the five markers were assembled and manually adjusted using BioEdit (Hall 1999) and then submitted to the DNA Data Bank of Japan (DDBJ) (http://www.ddbj.nig.ac.jp/; Table 1). We have verified the accuracy and identity of consensus sequences by comparing with sequences in GenBank (https://www.ncbi.nlm.nih.gov/genbank/).

Species and sequences used in the phylogenetic analyses. Newly generated sequences are set in bold.

Taxon Strain/Specimen ITS nuc 28S rpb1 rpb2 tef1
Antrodia serialis KHL 12010 (GB) JX109844 JX109844 JX109870 JX109898
Aurantiporus croceus Miettinen-16483 KY948745 KY948901 KY948927
Bjerkandera adusta HHB-12826-Sp KP134983 KP135198 KP134784 KP134913 KT305938
Bjerkandera aff. centroamericana L-13104-sp KY948791 KY948855 KY948936
Byssomerulius corium FP-102382 KP135007 KP135230 KP134802 KP134921
Candelabrochaete africana FP-102987-Sp KP135294 KP135199 KP134872 KP134975
Ceraceomyces serpens HHB-15692-Sp KP135031 KP135200 KP134785 KP134914
Ceriporia alachuana FP-103881-Sp KP135341 KP135201 KP134845 KP134896
Ceriporia reticulata KHL 11981 (GB) JX109899
Ceriporia reticulata RLG-11354-Sp KP135041 KP135204 KP134794 KP134922
Ceriporiopsis aneirina HHB-15629-Sp KP135023 KP135207 KP134795
Ceriporiopsis carnegieae RLG-7277-T KY948792 KY948854 KY948935
Ceriporiopsis fimbriata Dai 11672 KJ698633 KJ698637
Ceriporiopsis gilvescens L-3519-sp KY948761 KY948919
Ceriporiopsis gilvescens Niemela-5516 HQ659222
Ceriporiopsis guidella HUBO 7659 FJ496687 FJ496722
Ceriporiopsis kunmingensis C.L. Zhao 152 KX081072 KX081074
Ceriporiopsis lagerheimii 58240 KX008365 KX081077
Ceriporiopsis pseudoplacenta Miettinen 18997 (H) KY948744 KY948902 KY948926
Cerrena unicolor FD-299 KP135304 KP135209 KP134874 KP134968
Climacodon sanguineus BR5020180728797 KX810931 KX810932 KX810934
Climacodon septentrionalis AFTOL-767 AY854082 AY684165 AY864872 AY780941 AY885151
Crustodontia chrysocreas I HHB-6333-Sp KP135358 KP135263 KP134861 KP134908
Crustodontia chrysocreas II FBCC307 LN611114 LN611114
Daedalea quercina FP-56429 KY948809 KY948883 KY948989
Datronia mollis RLG6304sp JN165002 JN164791 JN164818 JN164872 JN164901
Donkia pulcherrima I GC 1707-11 LC378994 LC379152 LC379157 LC387351 LC387371
Donkia pulcherrima II AH39127 KX810937
Donkia pulcherrima II Gothenburg-2022 KX752591 KX752591
Efibula americana FP-102165 KP135016 KP135256 KP134808 KP134916
Emmia lacerata FP-55521-T KP135024 KP135202 KP134805 KP134915
Fomitopsis pinicola AFTOL-770 AY854083 AY684164 AY864874 AY786056 AY885152
Gelatoporia subvermispora FD-354 KP135312 KP135212 KP134879
Geliporus exilisporus I GC 1702-15 LC378995 LC379153 LC379158 LC387352 LC387372
Geliporus exilisporus II Dai 2172 KU598211 KU598216
Gloeoporus pannocinctus L-15726-Sp KP135060 KP135214 KP134867 KP134973
Grammothelopsis puiggarii RP 134 KP859299 KP859308
Hapalopilus nidulans FD-512 KP135419 KP134809
Hapalopilus nidulans Josef Vlasak JV0206/2 (JV) KX752623
Hapalopilus ochraceolateritius Miettinen-16992.1 KY948741 KY948891 KY948965
Heterobasidion annosum AFTOL-ID 470 DQ206988 DQ667160 DQ028584
Heterobasidion annosum DAOM-73191 AF287866 AY544206
Hydnophanerochaete odontoidea Chen 1376 LC363485
Hydnophanerochaete odontoidea GC 1308-45 LC363486 LC363492 LC363497 LC387353 LC387373
Hydnophanerochaete odontoidea GC 1607-20 LC378996
Hydnophanerochaete odontoidea GC 1710-59 LC378997
Hydnophanerochaete odontoidea WEI 15-309 LC378998
Hydnophanerochaete odontoidea WEI 15-348 LC378999
Hydnophanerochaete odontoidea Wu 0106-35 LC379000 LC379154 LC379159 LC387354 LC387374
Hydnophanerochaete odontoidea (Phanerochaete subodontoidea) Wu 911206-38 LC379001
Hydnophanerochaete odontoidea Wu 9310-29 LC379002
Hydnophanerochaete odontoidea Wu 9310-8 MF399408 GQ470653 LC314328 LC387355 LC387375
Hydnophanerochaete odontoidea (Phanerochaete subodontoidea) CWN00776 LC363487 GQ470663 LC363498 LC387356 LC387376
Hydnophlebia chrysorhiza FD-282 KP135338 KP135217 KP134848 KP134897
Hydnophlebia omnivora I KKN-112-Sp KP135334 KP135216 KP134846
Hydnophlebia omnivora II ME-497 KP135332 KP135218 KP134847
Hydnopolyporus fimbriatus Meijer3729 (O) JN649346 JN649346 JX109875 JX109904
Hyphoderma mutatum HHB-15479-Sp KP135296 KP135221 KP134870 KP134967
Hyphoderma setigerum CHWC 1209-9 LC387357 LC270919
Hyphoderma setigerum FD-312 KP135297 KP135222 KP134871
Hyphodermella corrugata MA- 24238 FN600378 JN939586
Hyphodermella poroides Dai 10848 KX008368 KX011853
Hyphodermella rosae FP-150552 KP134978 KP135223 KP134823 KP134939
Irpex lacteus DO 421/951208 (O) JX109882 JX109911
Irpex lacteus FD-9 KP135026 KP135224 KP134806
Leptoporus mollis TJV–93–174T KY948795 EU402510 KY948957
Lilaceophlebia livida I FBCC937 LN611122 LN611122
Lilaceophlebia livida II FP-135046-sp KY948758 KY948850 KY948917
Lopharia cinerascens FP-105043-sp JN165019 JN164813 JN164840 JN164874
Luteoporia albomarginata GC 1702-1 LC379003 LC379155 LC379160 LC387358 LC387377
Meruliopsis taxicola SK 0075 (GB) JX109847 JX109847 JX109873 JX109901
Merulius tremellosus ES2008-2 (GB) JX109859 JX109916
Merulius tremellosus FD-323 KP135231 KP134856 KP134900
Mycoacia fuscoatra HHB-10782-Sp KP135365 KP135265 KP134857 KP134910
Mycoacia fuscoatra KHL 13275 (GB) JX109908
Mycoacia nothofagi HHB-4273-Sp KP135369 KP135266 KP134858 KP134911
Obba rivulosa FP-135416-Sp KP135309 KP135208 KP134878 KP134962
Odontoefibula orientalis GC 1604-130 LC363489 LC363494 LC363500 LC387359 LC387378
Odontoefibula orientalis GC 1703-76 LC379004 LC379156 LC379161 LC387360 LC387379
Odontoefibula orientalis Wu 0805-59 LC363488 LC363493 LC363499 LC387361 LC387380
Odontoefibula orientalis Wu 0910-57 LC363490 LC363495 LC363501 LC387362 LC387381
Odoria alborubescens BP106943 MG097864 MG097867 MG213724 MG213723
Oxychaete cervinogilvus Schigel-5216 KX752596 KX752596 KX752626
Phaeophlebiopsis caribbeana HHB-6990 KP135415 KP135243 KP134810 KP134931
Phaeophlebiopsis peniophoroides FP-150577 KP135417 KP135273 KP134813 KP134933
Phanerina mellea WEI 17-224 LC387333 LC387340 LC387345 LC387363 LC387382
Phanerochaete arizonica RLG-10248-Sp KP135170 KP135239 KP134830 KP134949
Phanerochaete chrysosporium HHB-6251-Sp KP135094 KP135246 KP134842 KP134954
Phanerochaete ericina HHB-2288 KP135167 KP135247 KP134834 KP134950
Phanerochaete exilis HHB-6988 KP135001 KP135236 KP134799 KP134918
Phanerochaete laevis HHB-15519-Sp KP135149 KP135249 KP134836 KP134952
Phanerochaete livescens Wu 0711-81 LC387334 MF110289 LC387346 LC387364 LC270920
Phanerochaete magnoliae HHB-9829-Sp KP135089 KP135237 KP134838 KP134955
Phanerochaete pseudosanguinea FD-244 KP135098 KP135251 KP134827 KP134942
Phanerochaete rhodella FD-18 KP135187 KP135258 KP134832 KP134948
Phanerochaete sp. HHB-11463 KP134994 KP135235 KP134797 KP134892
Phanerochaete taiwaniana Wu 0112-13 MF399412 GQ470665 LC314332 LC387365 LC387383
Phebia acerina FD-301 KP135378 KP135260 KP134862
Phlebia acanthocystis I GC 1703-30 LC387338 LC387343 LC387366 LC387384
Phlebia acanthocystis II FP150571 KY948767 KY948844 KY948914
Phlebia albida GB-1833 KY948748 KY948889 KY948960
Phlebia brevispora FBCC1463 LN611135 LN611135
Phlebia centrifuga HHB-9239-Sp KP135380 KP135262 KP134844 KP134974
Phlebia coccineofulva HHB-11466-sp KY948766 KY948851 KY948915
Phlebia deflectens FCUG 1568 AF141619 AF141619
Phlebia firma Edman K268 EU118654 EU118654 JX109890
Phlebia floridensis HHB-9905-Sp KP135383 KP135264 KP134863 KP134899
Phlebia hydnoidea HHB-1993-sp KY948778 KY948853 KY948921
Phlebia lilascens FCUG 1801 AF141621 AF141621
Phlebia ochraceofulva FBCC295 LN611116 LN611116
Phlebia radiata AFTOL-484 AY854087 AF287885 AY864881 AY218502 AY885156
Phlebia setulosa HHB-6891-Sp KP135382 KP135267 KP134864 KP134901
Phlebia sp. FD-427 KP135342 KP134849
Phlebia sp. GC 1703-31 LC387339 LC387344 LC387347 LC387367 LC387385
Phlebia sp. GC 1708-118 LC387337 LC387342 LC387349 LC387368 LC387386
Phlebia sp. GC 1710-83 LC387336 LC387341 LC387350 LC387369 LC387387
Phlebia sp. HHB-17984 KP135359 KP135261 KP134860 KP134907
Phlebia sp. HHB-18295 KP135405 KP135269 KP134814 KP134938
Phlebia subochracea I HHB-8715-sp KY948770 KY948846 KY948913
Phlebia subochracea II HHB-8494-sp KY948768 KY948845 KY948912
Phlebia subserialis FCUG 1434 AF141631 AF141631
Phlebia uda FP-101544-Sp KP135361 KP135232 KP134859 KP134909
Phlebia unica KHL 11786 (GB) EU118657 EU118657 JX109861 JX109889
Phlebiopsis crassa KKN-86-Sp KP135394 KP135215 KP134820 KP134928
Phlebiopsis gigantea FP-70857-Sp KP135390 KP135272 KP134821 KP134930
Phlebiopsis ravenelii FP-110129-Sp KP135362 KP135274 KP134850 KP134898
Phlebiporia bubalina Dai 13168 KC782526 KC782528
Pirex concentricus OSC-41587 KP134984 KP135275 KP134843 KP134940
Rhizochaete filamentosa HHB-3169-Sp KP135410 KP135278 KP134818 KP134935
Rhizochaete radicata FD-123 KP135407 KP135279 KP134816 KP134937
Rhizochaete rubescens Wu 0910-45 LC387335 MF110294 LC387348 LC387370 LC270925
Riopa metamorphosa Viacheslav Spirin 2395 (H) KX752601 KX752601 KX752628
Sarcodontia crocea OMC-1488 KY948798 KY948903 KY948928
Scopuloides rimosa I HHB-7042-Sp KP135350 KP135282 KP134853 KP134903
Scopuloides rimosa II RLG-5104 KP135351 KP135283 KP134852 KP134904
Skeletocutis nivea ES2008-1 (GB) JX109858 JX109858 JX109886 JX109915
Steccherinum ochraceum KHL 11902 (GB) JQ031130 JQ031130 JX109865 JX109893
Stereum hirsutum AFTOL-ID 492 AY854063 AY864885 AY218520 AY885159
Stereum hirsutum FPL-8805 AF393078
Terana caerulea FP-104073 KP134980 KP135276 KP134865 KP134960
Trametes versicolor FP-135156-sp JN164919 JN164809 JN164825 JN164850 DQ028603
Trametopsis cervina TJV–93–216T JN165020 JN164796 JN164839 JN164877 JN164882
Tyromyces chioneus FD-4 KP135311 KP135291 KP134891 KP134977

Phylogenetic analyses

Two datasets were compiled for phylogenetic analyses: the ITS+nuc 28S+rpb1+rpb2+tef1 dataset was analysed to confirm the generic placement of target species within the phlebioid clade of Polyporales. The ITS dataset was used to get better resolutions on species level within the Hydnophanerochaete clade of Meruliaceae. The selection of strains and species for the 5-marker dataset was based on Binder et al. (2013), Floudas and Hibbett (2015), Kuuskeri et al. (2015), Justo et al. (2017), Miettinen et al. (2016), Moreno et al. (2017), Papp and Dima (2017), Yuan et al. (2017) and Zhao et al. (2017). Alignment was done with MAFFT v. 7 using two strategies: Q-INS-I for ITS and FFT-NS-I for nuc 28S, rpb1, rpb2 and tef1 (Katoh and Standley 2013). The resulting alignments were manually adjusted in Mega 7 (Kumar et al. 2016). Heterobasidion annosum (Fr.) Bref. and Stereum hirsutum (Willd.) Pers., belonging to Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C. David, were chosen as the outgroup in the 5-marker dataset. Phlebia coccineofulva Schwein., belonging to Meruliaceae, was assigned as the outgroup in the ITS dataset. Optimised datasets were deposited at TreeBASE (submission ID 22932).

The Bayesian Inference (BI) method was carried out for both datasets using MrBayes v. 3.2.6 (Ronquist et al. 2012). The Maximum Likelihood (ML) method was carried out for the 5-marker dataset using RAxML BlackBox (Stamatakis 2014). For the BI analyses, jModeltest 2.1.10 (Darriba et al. 2012) was first used to estimate separate models for each of the markers in both datasets, based on Akaike information criterion (AIC). The Markov chain Monte Carlo (MCMC) search was run for ten million generations, with four chains and trees sampled every 100 generations. The first twenty-five percent of trees were discarded as burn-in while the remaining trees were used to construct the fifty percent majority-rule consensus phylogram with posterior probabilities (PP). For the ML analysis, the best-scoring tree with proportional values of bootstrap (BS) was computed under a GTRGAMMA model with one thousand bootstrap replicates, followed by a thorough ML search. Gaps were treated as missing data. Branches were regarded as having statistical support if values of PP and/or BS were equal to or over 0.9 and 70%, respectively. Both BI and ML analyses were performed at the CIPRES Science Gateway (Miller et al. 2010; http://www.phylo.org/). Phylograms were visualised and edited in TreeGraph 2 (Stöver and Müller 2010) and Adobe Illustrator (Adobe Systems, Inc).

Phylogeny results

The final ITS+nuc 28S+rpb1+rpb2+tef1 dataset consisted of 126 sequences and 7253 characters (of which 43.7% were parsimony-informative) including gaps and the ITS dataset comprised 12 sequences and 887 characters (of which 7.7% were parsimony-informative) including gaps. In the BI analyses, since the GTR+G+I model was selected as the best model of nucleotide substitution for each of the five markers in the 5-marker dataset, it was used for the entire alignment with five partitions. The HKY+I+G model was selected as the best model of nucleotide substitution for the ITS dataset. The fifty percent majority-rule consensus phylogram with PP support values was reconstructed after the average standard deviation of split frequencies fell below 0.001. The best-scoring ML tree with BS support values was built. Phylogenetic trees of the 5-marker dataset, inferred from BI and ML algorithms, shared similar topologies and thus only the ML tree was shown (Fig. 1).

Figure 1. 

Phylogenetic tree inferred from Maximum Likelihood analysis of the combined ITS, nuc 28S, rpb1, rpb2 and tef1 sequences of taxa in Polyporales. Nodes are labelled with Maximum Likelihood bootstrap proportional values (BS) ≥ 70% and Bayesian Posterior Probabilities (PP) ≥ 0.9. Thickened branches obtained supports by both BS ≥ 80% and PP ≥ 0.95. The taxa studied in this study are shown in bold. The pale blue boxes indicate lineages of phanerochaetoid within the phlebioid clade. Asterisks (*) represent for strains of generic type species. Scale bars = substitutions per site.

In the 5-marker analyses (Fig. 1), six main clades with high statistic supports (BS = 96–100%, PP = 1) could be recognised in the ingroup: the antrodia clade, the core polyporoid clade, the gelatoporia clade, the phlebioid clade, a residual clade and the skeletocutis-tyromyces clade. The phlebioid clade, which is the focus of this study, included three main subclades recognised as three families (BS = 100%, PP = 1): Irpicaceae, Meruliaceae and Phanerochaetaceae. Hydnophanerochaete odontoidea formed a well-supported monophyletic lineage (BS = 100%, PP = 1) within Meruliaceae and was found to be closely related to a lineage consisting of strains of Ceriporia alachuana (Murrill) Hallenb, Ceriporiopsis spp., Grammothelopsis puiggarii (Speg.) Rajchenb. & J.E. Wright, Hynophlebia spp. and Phlebia spp. (BS = 86%, PP = 1). Sequences of Odontoefibula orientalis grouped together and formed a well-supported monophyletic lineage (BS = 98%, PP = 1) within the Donkia clade of Phanerochaetaceae (BS = 97%, PP = 1) and were most closely related to a lineage made up of strains of Geliporus exilisporus (Y.C. Dai & Niemelä) Yuan Yuan, Jia J. Chen & S.H. He and Hyphodermella spp. (BS = 98%, PP = 1).

The tree inferred from the ITS dataset (Fig. 2) showed that sequences of holotype (CWN00776) and paratype (Wu 911206-38) of Phanerochaete subodontoidea were clustered with sequences of P. odontoidea within a monophyletic lineage (PP = 1).

Figure 2. 

The majority-rule consensus phylograms of the Bayesian Inference analysis of the ITS sequences of Hydnophanerochaete odontoidea. Nodes are labelled with Bayesian Posterior Probabilities ≥ 0.9. Scale bars = substitutions per site.

Taxonomy

Hydnophanerochaete Sheng H. Wu & C.C. Chen, gen. nov.

MycoBank No: MB824077

Type species

Hydnophanerochaete odontoidea (≡ Phanerochaete odontoidea).

Etymology

From hydnoid + Phanerochaete, referring to the hydnoid hymenial surface and a close affinity to Phanerochaete.

Description

Basidiocarps effused, adnate, ceraceous. Hymenial surface at first buff, with age turning ochraceous to pale brown, slightly tuberculate to grandinioid when young, becoming odontioid to hydnoid with age, without colour changes in KOH. Aculei conical to cylindrical, ca. 1–4 per mm, up to 700 μm long.

Hyphal system essentially monomitic; generative hyphae simple-septate. Subiculum fairly uniform, composed of a basal layer, with compact texture; generative hyphae somewhat horizontal, colourless, thick-walled; quasi-binding hyphae present near substratum, colourless. Hymenial layer thickening. Trama of aculei of compact texture; generative hyphae somewhat vertical, colourless, thick-walled. Cystidia lacking, but projecting hyphal ends in the hymenium may be present. Basidia clavate, 4-sterigmate. Basidiospores ellipsoid to cylindrical, smooth, thin-walled, inamyloid, non-dextrinoid, acyanophilous.

Remarks

Hydnophanerochaete is morphologically similar to the genus Hydnophlebia (Telleria et al. 2017). Both genera have resupinate basidiocarps with odontioid to hydnoid hymenial surface, a monomitic hyphal system, ordinarily simple-septate hyphae and similar basidiospore shape. However, we note three distinguishing differences. First, Hydnophlebia has membranaceous basidiocarps usually with rhizomorphic margin, while Hydnophanerochaete has ceraceous basidiocarps with fairly determinate margin. Second, occasional single or multiple clamp connections are present in subicular or aculei hyphae of Hydnophlebia, whereas they are lacking in hyphae of Hydnophanerochaete. Third, Hydnophlebia occasionally bears tubular to ventricose leptocystidia, which are lacking in Hydnophanerochaete.

Little morphological differences exist between Hydnophanerochaete and Odontoefibula: both genera have monomitic hyphal system with simple-septate hyphae and are lacking cystidia. However, Hydnophanerochaete is distinguished from Odontoefibula by its basidiocarps without colour change in KOH; additionally, its subiculum is compact, not dense.

Phanerodontia Hjortstam & Ryvarden, a recently proposed genus typified by P. dentata Hjortstam & Ryvarden (Hjortstam and Ryvarden 2010), is also morphologically similar to Hydnophanerochaete. However, the latter has a compact subiculum and quasi-binding hyphae near the substratum. Phanerodontia accommodates four species [P. chrysosporium (Burds.) Hjortstam & Ryvarden, P. dentata, P. irpicoides (Hjortstam) Hjortstam & Ryvarden and P. magnoliae (Berk. & M.A. Curtis) Hjortstam & Ryvarden], all of them possessing long leptocystidia (Hjortstam and Ryvarden 2010), whereas this structure is lacking in Hydnophanerochaete. Moreover, phylogenetically, strains of two species (P. chrysosporium and P. magnoliae) were recovered in Phanerochaetaceae which is only distantly related to Hydnophanerochaete (Fig. 1). However, the generic type has not been sequenced so far.

Hydnophanerochaete odontoidea (Sheng H. Wu) Sheng H. Wu & C.C. Chen, comb. nov.

MycoBank No: MB824078
Figs 3a and 4

Basionym

Phanerochaete odontoidea Sheng H. Wu, Botanical Bulletin of the Academia Sinica 41: 169, 2000.

Synonym

Phanerochaete subodontoidea Sheng H. Wu, Botanical Bulletin of the Academia Sinica 41: 172, 2000.

Holotype

TAIWAN. Ilan: Fushan Botanical Garden, 24°46’N, 121°35’E, 600 m alt., on fallen branch of angiosperm, leg. S.H. Wu et al., 7 Aug 1991, Wu 910807-11 (TNM F14816).

Description

Basidiocarps annual, effused, adnate, ceraceous, somewhat brittle, 50–200 μm thick in section (aculei excluded). Hymenial surface initially buff, with age turning ochraceous to pale brown, no colour changes in KOH, tuberculate to grandinioid when young, becoming odontioid to hydnoid with age, extensively cracked; margin paler to whitish, fairly determinate. Aculei conical to cylindrical, usually separate, with obtuse to acute apex, 1–4 per mm, up to 100–700 × 100–250 μm.

Hyphal system basically monomitic, some specimens with quasi-binding hyphae near substratum; generative hyphae simple-septate. Subiculum fairly uniform, composed of a basal layer of compact texture; generative hyphae mainly horizontal, colourless, 4–6 μm diam., with 0.8–1 µm thick walls; quasi-binding hyphae sometimes present near substratum, colourless, 1–3 µm diam. Hymenial layer thickening, with compact texture, generative hyphae somewhat vertical, colourless, 3–6 μm diam., slightly thick-walled. Trama of aculei of compact texture; generative hyphae mainly vertical, other features similar to those in subiculum; crystal masses present near apex. Cystidia lacking, but projecting hyphal ends in the hymenium may be present. Basidia clavate, 14–18 × 4.5–5.5 μm, 4-sterigmate. Basidiospores narrowly ellipsoid to cylindrical, adaxially slightly concave, smooth, thin-walled, homogeneous, inamyloid, non-dextrinoid, acyanophilous, 6–8.1 × 2.5–3.3 μm (Table 2). See also Wu (2000) for descriptions and illustrations.

Figure 3. 

Basidiocarp surfaces a Hydnophanerochaete odontoidea (holotype of Phanerochaete subodontoidea, CWN 00776) b Odontoefibula orientalis (holotype, Wu 0910-57). Scale bar: 1 mm.

Figure 4. 

Hydnophanerochaete odontoidea (holotype of Phanerochaete subodontoidea, CWN 00776) a Part of the vertical section of subiculum near substratum b Quasi-binding hyphae. Scale bar: 5 μm (a–b).

Aculei and basidiospore measurements of basidiocarps.

Species Specimens Aculei (per mm) Range (μm) L (μm) W (μm) Q n
Hydnophanerochaete odontoidea Chen 1376 1–3 (6–) 6.3–7.3 (–7.5) ´ (2.5–) 2.8–3.3 (–3.5) 6.8 3 2.2 30
CWN 00776 ‡, | 1–3 (6–) 6.8–8 (–8.5) ´ (2.5–) 2.7–3.2 (–3.5) 7.4 2.9 2.5 30
GC 1308-45 | 2–3 (6.5–) 6.7–7.6 (–8) ´ (2.8–) 2.8–3.3 (–3.8) 7.2 3.1 2.3 30
GC 1607-20 2–3 (7–) 7.4–9 (–10) ´ (2.8–) 2.9–3.5 (–4) 8.2 3.2 2.6 30
WEI 15-309 2–3 (6–) 6.1–7 (–7.5) ´ (2.5–) 2.7–3 (–3.3) 6.5 2.9 2.3 30
WEI 15-348 2–3 6–6.9 (–7.5) ´ (2.5–) 2.8–3.3 (–3.5) 6.5 3 2.1 30
Wu 0106-35 | 2–3 (6–) 6.4–7.8 (–8) ´ (2.5–) 2.8–3.1 (–3.3) 7.1 2.9 2.4 30
Wu 910807-11 3–4 (6–) 6.1–7 (–8) ´ (2.5–) 2.5–2.9 (–3.3) 6.5 2.7 2.5 30
Wu 911206-38 2–3 (6–) 6.3–7.7 (–8) ´ (2.8–) 2.9–3.2 (–3.5) 7 3 2.3 30
Wu 9310-8 †, | 2–4 (6–) 6.5–8 (–8.5) ´ (2.5–) 2.8–3.2 (–3.5) 7.2 3 2.4 30
Wu 9310-29 2–4 (6–) 6.9–8.1 (–9) ´ (2.5–) 2.7–3.3 (–3.7) 7.4 3 2.5 30
Odontoefibula orientalis GC 1604-130 | 4–5 (5–) 5.4–6.6 (–7) ´ (2.5–) 2.8–3.3 (–3.6) 6 3.1 1.96 30
GC 1703-76 | 4–5 (5.5–) 5.8–7.4 (–8) ´ (3–) 3.2–3.9 (–4) 6.6 3.5 1.85 30
Wu 0805-59 | 3–5 (5–) 5.1–6.2 (–7) ´ (2.5–) 2.9–3.4 (–3.6) 5.6 3.2 1.79 30
Wu 0807-53 3–6 (5–) 5.4–6.4 (–7) ´ (3–) 3.1–3.7 (–4) 5.9 3.4 1.71 30
Wu 0910-57 §, | 3–6 (5–) 5.4–6.1 (–6.5) ´ (2.8–) 2.9–3.4 (–3.6) 5.7 3.2 1.81 30

Habitat

On fallen branches of angiosperms or gymnosperms.

Distribution

Hitherto known from subtropical to temperate regions of China (Yunnan), Japan, Taiwan and Vietnam.

Additional specimens examined

CHINA. Yunnan: Diqing Tibetan Autonomous Prefecture, Deqin County, Xiayubeng Village, Shenhu Trail, 3500 m alt., on fallen branch of gymnosperm, leg. C.C. Chen, 14 Aug 2013, GC 1308-45 (TNM F27660). JAPAN. Honshu: Nagano Prefecture, Nagano City, Myoko-Togakushi Renzan National Park, 36°45’35”N, 138°04’20”E, 1235 m alt., on branch of Quercus sp., leg. C.C. Chen & C. L. Chen, 29 July 2016, GC 1607-20 (TNM F30785). TAIWAN. Chiayi: Yushan National Park, Nanhsi Forest Road, 23°28’N, 120°54’E, 1850 m alt., on fallen branch of angiosperm, leg. S.H. Wu & S.Z. Chen, 13 Oct 1993, Wu 9310-8 (paratype of P. odontoidea, TNM F14824); Wu 9310-29 (TNM F14826); 1800 m alt., on fallen branch of angiosperm, leg. S.H. Wu & S.Z. Chen, 13 Jun 1996, Wu 9606-55 (TNM F5085). Ilan: Fushan Botanical Garden, 24°46’N, 121°35’E, 650 m alt., on fallen branch of angiosperm, leg. S.H. Wu et al., 28 Jun 2002, Wu 0106-35 (TNM F13460). Nantou: Tungpu Township, Leleku, 1450 m alt., on fallen rotten wood, leg. W.N. Chou, 13 Apr 1994, CWN 00776 (holotype of P. subodontoidea, TNM F14836). Kaohsiung: Maolin District, Tona Nursery, 22°54’N, 120°44’E, 850 m alt., on fallen branch of angiosperm, leg. S.Z. Chen, 31 Mar 2005, Chen 1376 (TNM F18764). New Taipei: Chinshan District, Yangmingshan National Park, Yulu Historical Trail, 25°10’N, 121°35’E, 516 m alt., on fallen branch of angiosperm, leg. C.C. Chen, C.L. Wei, W.C. Chen & S. Li, 26 Aug 2015, WEI 15-309 (TNM F29370); WEI 15-348 (TNM F29384). Taichung: Chiapaotai, 850 m alt., on fallen branch of angiosperm, leg. S.H. Wu, 6 Dec 1991, Wu 911206-38 (paratype of P. subodontoidea, TNM F14818). VIETNAM. Lam Dong: Bi Doup Nui Ba National Park, 12°10’45”N, 108°40’48”E, 1447 m alt., on fallen branch of angiosperm, leg. C.C. Chen, 15 Oct 2017, GC 1710-59 (TNM F31365).

Remarks

Phanerochaete subodontoidea morphologically resembles Phanerochaete odontoidea, whereas they were distinguished merely based on the width of basidiospores [P. odontoidea: 2.6–3 µm vs. P. subodontoidea: 3–3.7 µm, Wu (2000)]. However, after carefully measuring the basidiospore size of available specimens of these two species, we found basidiospore ranges are highly overlapping (Table 2). Additionally, the ITS sequences of the holotype of P. subodontoidea (CWN 00776) is almost identical to the ITS sequences of the paratype of P. odontoidea (Wu 9310-8). We failed to obtain sequences from the holotype of P. odontoidea (Wu 910807-11), but Wu 9310-8 was confirmed as conspecific with the holotype by morphological comparison. Thus, based on morphological and molecular evidence (Fig. 2), P. subodontoidea is treated as a synonym of P. odontoidea. A paratype specimen named P. odontoidea (Wu 9311-46) probably belongs to the genus Flavodon Ryvarden based on preliminary BLAST results of nuc 28S sequences. However, this specimen was not included in this study.

Odontoefibula C.C. Chen & Sheng H. Wu, gen. nov.

MycoBank No: MB824075

Type species

Odontoefibula orientalis.

Etymology

From odonto (= tooth-like) + efibula (= without clamp connection), referring to the odontioid hymenial surface and simple-septate hyphae of the genus.

Description

Basidiocarps annual, resupinate, effused, adnate, membranaceous to ceraceous. Hymenial surface at first honey yellow, becoming ochraceous to pale brown with age, turning dark reddish in KOH, initially smooth to slightly tuberculate, becoming grandinioid to odontioid with age. Aculei conical to cylindrical, separate or fused, up to 0.3 mm long.

Hyphal system monomitic; hyphae normally simple-septate. Subiculum uniform, with dense texture; basal hyphae interwoven, somewhat horizontal or with irregular orientation, colourless, thin- to slightly thick-walled; subicular hyphae somewhat vertical, colourless, thin- to slightly thick-walled. Subhymenium not clearly differentiated from subiculum. Central trama of fairly dense texture; hyphae vertical, colourless, thin- to slightly thick-walled. Cystidia lacking, but projecting hyphal ends in the hymenium may be present. Basidia clavate to narrowly clavate, 4-sterigmate. Basidiospores ellipsoid, smooth, thin-walled, inamyloid, non-dextrinoid, acyanophilous.

Remarks

Phaneroites Hjortstam & Ryvarden, a monotypic genus introduced to accommodate P. subquercinus (Henn.) Hjortstam & Ryvarden, resembles Odontoefibula in having odontioid hymenial surface and a monomitic hyphal system with ordinarily simple-septate hyphae. However, Phaneroites is distinguished from Odontoefibula by having thin-walled subicular hyphae, a few clamped septa on hyphae next to the substratum and subcapitate cystidia (Hjortstam and Ryvarden 2010). Moreover, basidiocarps of Odontoefibula turn dark reddish in KOH, while this reaction was not reported from Phaneroites.

Odontoefibula orientalis C.C. Chen & Sheng H. Wu, sp. nov.

MycoBank No: MB824076
Figs 3b and 5

Holotype

CHINA. Beijing: Xiangshan Park, 39°59’N, 116°11’E, 70 m alt., on fallen trunk of Amygdalus davidiana (Carrière) de Vos ex Henry, leg. S.H. Wu, 14 Oct 2009, Wu 0910-57 (TNM F23847).

Etymology

From orientalis (= Eastern world), where the specimens were collected.

Description

Basidiocarps annual, effused, adnate, membranaceous to subceraceous, somewhat brittle, 200–400 μm thick in section (aculei excluded). The hymenial surface at first honey yellow, darkening to ochraceous to pale brown with age, turning dark reddish in KOH, slightly tuberculate when young, becoming odontioid with age, extensively cracked; margin paler, thinning out, slightly filamentous. Aculei conical to cylindrical, usually fused at the base, with rounded to obtuse apex, 3–6 per mm, ca. 0.1–0.3 × 0.1–0.2 mm.

Hyphal system monomitic; hyphae simple-septate. Subiculum uniform, with dense texture, 200–300 μm thick; subicular hyphae somewhat vertical, colourless, 2.5–4 μm diam., 0.5–0.8 μm thick walls; hyphae near substratum interwoven, with irregular orientation, tortuous, colourless, irregularly swollen, 4–8 μm diam., 0.5–1 μm thick walls. Subhymenium not clearly differentiated from subiculum, with fairly dense texture, hyphae somewhat vertical, colourless, 3–4 μm diam., thin- to slightly thick-walled. Trama of aculei of dense texture; hyphae mainly vertical, other aspects similar to those in subiculum. Large crystal masses scattered throughout the section. Cystidia lacking, but projecting hyphal ends in the hymenium may be present. Basidia clavate to narrowly clavate, 25–40 × 6–7 μm, 4-sterigmate, often with small oily drops. Basidiospores ellipsoid, adaxially slightly concave, smooth, thin-walled, sometimes with small oily drops, inamyloid, non-dextrinoid, acyanophilous, 5.1–6.6 × 2.8–3.4 μm (Table 2).

Figure 5. 

Odontoefibula orientalis (holotype, Wu 0910-57) a Profile of basidiocarp section b Part of the vertical section of trama c Basal hyphae d Subicular hyphae e Basidia f Basidiospores. Scale bars: 200 μm (a); 10 μm (c–d); 5 μm (e–f).

Habitat

On fallen trunk of angiosperm (e.g. Amygdalus).

Distribution

Hitherto known from China (Beijing), Japan and Taiwan.

Additional specimens examined (paratypes)

JAPAN. Honshu: Ibaraki Prefecture, Joso City, Mt. Ju-ichimen-yama, along Kinu-gawa River, on branch of Prunus sp., leg. S.H. Wu, 12 July 2008, Wu 0807-53 (TNM F22091). TAIWAN. Pingtung: Laiyi Township, Pengjishan Trail, 22°30’52”N, 120°38’07”E, 248 m alt., on fallen trunk of angiosperm, leg. C.C. Chen, 25 Mar 2017, GC 1703-76 (TNM F31460). Taichung: Hoping District, between 27–27.5 km of Dasyueshan Forestry Road, Yuanzueishan Trail, 1800 m alt., on fallen rotten trunk of angiosperm, leg. S.H. Wu, S.Z. Chen & Y.T. Wang, 22 May 2008, Wu 0805-59 (TNM F22495). Hualien: Sioulin Township, Taroko National Park, Lushui Hiking Trail, 24°10’51”N, 121°30’10”E, 578 m alt., on fallen trunk of angiosperm, leg. C.C. Chen, 24 Apr 2016, GC 1604-130 (TNM F31364).

Discussion

Our 5-marker phylogenetic analyses (Fig. 1) provided an updated taxonomic framework for evaluating generic placements of the target taxa of the phlebioid clade. The tree topologies are consistent with previous results (Wu et al. 2010; Floudas and Hibbett 2015; Justo et al. 2017; Papp and Dima 2017). Within the phlebioid clade, we recovered two monophyletic lineages of phanerochaetoid fungi (Fig. 1), which supports the status of the two genera erected here: Hydnophanerochaete, typified by P. odontoidea, is accommodated in Meruliaceae; Odontoefibula, typified by O. orientalis, is placed in Donkia clade of Phanerochaetaceae.

Phylogenetically, Hydnophanerochaete and Odontoefibula are independent from the nine lineages of phanerochaetoid fungi recognised by Floudas and Hibbett (2015) within the phlebioid clade: Efibula, Hydnophlebia, Phaeophlebiopsis, “Phanerochaeteallantospora Burds. & Gilb., Phanerochaete s.l., Phanerochaete s.s., Phlebiopsis, Rhizochaete and Scopuloides. P. allantospora was not sampled in this study; it was placed in Irpicaceae, according to the study of Justo et al. (2017). Additionally, “Phanerochaeteginnsii Sheng H. Wu represents another lineage of phanerochaetoid fungi that was not analysed in this study, nor in the study of Floudas and Hibbett (2015). This species was shown to be closely related to Phlebia centrifuga P. Karst (Wu et al. 2010).

The 5-marker phylogenetic analyses (Fig. 1) suggest a close relationship amongst Hydnophanerochaete odontoidea and the following taxa, which all have a monomitic hyphal system with simple-septate hyphae: Hydnophlebia, Ceriporia alachuana, Climacodon septentrionalis (Fr.) P. Karst. and Scopuloides rimosa (Cooke) Jülich. Like Hydnophanerochaete, Hydnophlebia and Scopuloides have an odontioid to hydnoid hymenial surface. However, Hydnophlebia differs by its membraneous basidiocarps with rhizomophic margin, occasional clamped subicular hyphae and the presence of tubular to ventricose leptocystidia (Telleria et al. 2017). Scopuloides differs by thick-walled encrusted cystidia and rather short, clavate basidia (Wu 1990). C. alachuana resembles H. odontoidea in lacking cystidia, but has a poroid hymenial surface (Ryvarden and Gilbertson 1993). C. septentrionalis has a hydnoid hymenial surface, but is clearly distinguished by its pileate basidiocarps and thick-walled encrusted cystidia (Maas Geesteranus 1971).

Quasi-binding hyphae, one of the diagnostic characters of H. odontoidea (Fig. 4), were first introduced by Wu (1990) to refer to narrow and much branched subicular hyphae with thin- to thick walls, found near the substrate. Wu (2000) omitted describing and illustrating the quasi-binding hyphae of P. odontoidea and P. subodontoidea. Quasi-binding hyphae have been reported from many species of diverse genera: Amethicium leoninum (Burds. & Nakasone) Sheng H. Wu, Crustodontia chrysocreas (Berk. & M.A. Curtis) Hjortstam & Ryvarden, Phlebiporia bubalina Jia J. Chen, B.K. Cui & Y.C. Dai, Phanerochaete ericina (Bourdot) J. Erikss. & Ryvarden, Pseudolagarobasidium calcareum (Cooke & Massee) Sheng H. Wu and Radulodon americanus Ryvarden (Wu 1990; Stalpers 1998; Chen and Cui 2014). In other words, this feature has a polyphyletic origin and does not seem to be very phylogenetically informative.

Within the Donkia clade (Fig. 1), systematic positions of two recently proposed taxa, Geliporus exilisporus and Hyphodermella poroides Y.C. Dai & C.L. Zhao, are confirmed in this study. Odontoefibula shares some ubiquitous features with the genera Donkia, Hyphodermella J. Erikss. & Ryvarden and Pirex Hjortstam & Ryvarden, many of which have ochraceous basidiocarps with odontioid to hydnoid hymenial surfaces. However, to better illustrate the correspondence between molecular data and morphology, denser taxon sampling of this clade is necessary in the future.

Acknowledgments

This study was financed by Ministry of Science and Technology of R.O.C (Taiwan) (Grant no 104-2621-B-178-001-MY3). The authors are indebted to Dr. Dimitrios Floudas (Department of Biology, Lund University, Sweden) and reviewers for their valuable comments and suggestions on improving the manuscript. We are also grateful to Ms. Siou-Zhen Chen (TNM) for managing studied specimens and to Shin-Yi Ke (TNM) for the help in DNA sequencing work.

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