201830012018298195FF9FFFF9-632A-FFA6-2004-FFD8FF86252711701720611201718012018Zi-Qiang Wu, Tai-Min Xu, Shan Shen, Xiang-Fu Liu, Kai-Yue Luo, Chang-Lin ZhaoThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
A new poroid wood-inhabiting fungal genus, Elaphroporia, typified by E.ailaoshanensissp. nov., is proposed based on a combination of morphological features and molecular evidence. The genus is characterised by an annual growth habit, resupinate basidiocarps, becoming rigid and light-weight up on drying, a monomitic hyphal system with thick-walled generative hyphae bearing both clamp connections and simple septa, slightly amyloid, CB+ and ellipsoid, hyaline, thin-walled, smooth and IKI–, CB– basidiospores. Sequences of ITS and LSU nrRNA gene regions of the studied samples were generated, and phylogenetic analyses were performed with maximum likelihood, maximum parsimony and bayesian inference methods. The phylogenetic analysis based on molecular data of ITS+nLSU sequences showed that Elaphroporia belonged to the residual polyporoid clade and was closely related to Junghuhniacrustacea. Further investigation was obtained for more representative taxa in the Meruliaceae based on ITS+nLSU sequences, in which the result demonstrated that the genus Elaphroporia formed a monophyletic lineage with a strong support (100 % BS, 100 % BP, 1.00 BPP) and then grouped with Flaviporus and Steccherinum.
Wu Z-Q, Xu T-M, Shen S, Liu X-F, Luo K-Y, Zhao C-L (2018) Elaphroporia ailaoshanensis gen. et sp. nov. in Polyporales (Basidiomycota). MycoKeys 29: 81–95. https://doi.org/10.3897/mycokeys.29.22086
Introduction
The Polyporales is a large group of Agaricomycetes and includes more than 1800 taxa at species level belonging to 216 genera and 13 families (Kirk et al. 2008). Species in Polyporales are the key players amongst the wood-rotting fungi because of their importance in the carbon cycle (Floudas et al. 2012) and the pathogenic and potential application in biomedical engineering and biodegradation (Dai et al. 2009, Levin et al. 2016).
Molecular systematics has played a powerful role in inferring phylogenies within fungal groups since the early 1990s (White et al. 1990, Hibbett et al. 2007, Larsson 2007, Miettinen et al. 2011, Binder et al. 2013, Dai et al. 2015, Choi and Kim 2017). Recently, molecular studies involving Meruliaceae P. Karst. have been carried out (Binder et al. 2005, 2013, Miettinen and Larsson 2011, Miettinen and Rajchenberg 2012, Hibbett et al. 2016, Miettinen et al. 2016).
Larsson (2007) introduced a new division for part of the Polyporales, effectively renaming the phlebioid and residual polyporoid clades as the Meruliaceae, Phanerochaetaceae Jülich, and Byssomerulius Parmasto families. A phylogenetic study of Meruliaceae employing multi-genes suggested that 1) this family included species with both poroid and hydnoid hymenophore configurations, and 2) the genera of Flabellophora G. Cunn., Flaviporus Murrill, Junghuhnia Corda, Steccherinum Gray and Xanthoporus Audet belong to this family (Miettinen et al. 2011). Moreover, further study employing a six-gene (5.8S, nrLSU, nrSSU, rpb1, rpb2, tef1) dataset has constructed a phylogenetic and phylogenomic overview of the Polyporales, which showed that the species of Meruliaceae fall into the residual polyporoid clade (Binder et al. 2013).
Wood-rotting fungi is a cosmopolitan group and it has a rich diversity on the basis of growing on boreal, temperate, subtropical, and tropical vegetations (Gilbertson and Ryvarden 1987, Núñez and Ryvarden 2001, Dai 2012, Ryvarden and Melo 2014, Dai et al. 2015). During investigations on wood-inhabiting fungi in southern China, an additional taxon was found which could not be assigned to any described genus. It produces annual, resupinate basidiocarps, a monomitic hyphal system with generative hyphae bearing both simple septa and clamp connections, slightly amyloid, CB+ and ellipsoid, hyaline, thin-walled, smooth basidiospores. These characters make it distinguishable from all known poroid and hydnoid wood-inhabiting fungal genera (Gilbertson and Ryvarden 1987, Núñez and Ryvarden 2001, Bernicchia and Gorjón 2010, Ryvarden and Melo 2014). In this study, the authors expand samplings from previous studies to examine taxonomy and phylogeny of this new genus within the Polyporales, based on the internal transcribed spacer (ITS) regions and the large subunit nuclear ribosomal RNA gene (nLSU) sequences.
Materials and methods
Morphological studies. The specimens studied are deposited at the herbarium of Southwest Forestry University (SWFC). Macro-morphological descriptions are based on field notes. Special colour terms follow Petersen (1996). Micro-morphological data were obtained from the dried specimens and observed under a light microscope following Dai (2010). The following abbreviations were used: KOH = 5% potassium hydroxide, CB = Cotton Blue, CB– = acyanophilous, IKI = Melzer’s reagent, IKI– = both inamyloid and indextrinoid, IKI+ = amyloid, L = mean spore length (arithmetic average of all spores), W = mean spore width (arithmetic average of all spores), Q = variation in the L/W ratios between the specimens studied, n (a/b) = number of spores (a) measured from given number (b) of specimens.
DNA extraction and sequencing. CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain genomic DNA from dried specimens, according to the manufacturer’s instructions with the modification that a small piece of dried fungal specimen (about 30 mg) was ground to powder with liquid nitrogen. The powder was transferred to a 1.5 ml centrifuge tube, suspended in 0.4 ml of lysis buffer and incubated in a 65 °C water bath for 60 min. After that, 0.4 ml phenol-chloroform (24:1) was added to each tube and the suspension was shaken vigorously. After centrifugation at 13 000 rpm for 5 min, 0.3 ml supernatant was transferred to a new tube and mixed with 0.45 ml binding buffer. The mixture was then transferred to an adsorbing column (AC) for centrifugation at 13 000 rpm for 0.5 min. Then, 0.5 ml inhibitor removal fluid was added in AC for a centrifugation at 12 000 rpm for 0.5 min. After washing twice with 0.5 ml washing buffer, the AC was transferred to a clean centrifuge tube, and 100 ml elution buffer was added to the middle of the adsorbed film to elute the genome DNA. The ITS region was amplified with primer pairs ITS5 and ITS4 (White et al. 1990). The nuclear LSU region was amplified with primer pairs LR0R and LR7 (https://sites.duke.edu/vilgalyslab/rdna_primers_for_fungi/). The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 58 °C for 45 s and 72 °C for 1 min, and a final extension of 72 °C for 10 min. The PCR procedure for nLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 48 °C for 1 min and 72 °C for 1.5 min, and a final extension of 72 °C for 10 min. The PCR products were purified and directly sequenced at Kunming Tsingke Biological Technology Limited Company. All newly generated sequences were deposited at GenBank (Table 1).
A list of species, specimens and GenBank accession number of sequences used in this study.
Species name
Sample no.
GenBank accession no.
References
ITS
nLSU
Abortiporusbiennis
TFRI 274
EU232187
EU232235
Larsson (2007)
Antrodiaalbida
CBS 308.82
DQ491414
AY515348
Kim et al. (2007)
Antrodiaheteromorpha
CBS 200.91
DQ491415
AY515350
Kim et al. (2007)
Antrodiellaamericana
Gothenburg 3161
JN710509
JN710509
Miettinen et al. (2011)
Antrodiellapallasii
Renvall 89a
AF126896
–
Binder et al. (2013)
Antrodiellasemisupina
FCUG 960
EU232182
EU232266
Binder et al. (2005)
Antrodiella sp.
X 418
JN710523
JN710523
Miettinen et al. (2011)
Atraporiellaneotropica
Ryvarden 44447
HQ659221
HQ659221
Miettinen and Rajchenberg (2012)
Ceriporiaviridans
Dai 7759
KC182777
–
Jia et al. (2014)
Ceriporiopsisbalaenae
H7002389
FJ496669
FJ496717
Tomšovský et al. (2010)
Ceriporiopsisconsobrina
Rivoire 977
FJ496667
FJ496716
Tomšovský et al. (2010)
Ceriporiopsisgilvescens
BRNM 667882
FJ496685
FJ496719
Tomšovský et al. (2010)
Ceriporiopsisgilvescens
BRNM 710166
FJ496684
FJ496720
Tomšovský et al. (2010)
Ceriporiopsisgilvescens
Yuan 2752
KF845946
KF845953
Zhao and Cui (2014)
Ceriporiopsisguidella
HUBO 7659
FJ496687
FJ496722
Tomšovský et al. (2010)
Cinereomyceslindbladii
FBCC 177
HQ659223
HQ659223
Miettinen and Rajchenberg (2012)
Climacocystisborealis
KH 13318
JQ031126
JQ031126
Binder et al. (2013)
Coriolopsiscaperata
LE(BIN)-0677
AB158316
AB158316
Tomšovský et al. (2010)
Dacryoboluskarstenii
KHL 11162
EU118624
EU118624
Binder et al. (2005)
Daedaleaquercina
DSM 4953
DQ491425
DQ491425
Kim et al. (2007)
Diplomitoporusflavescens
X 84
FN907908
–
Miettinen et al. (2011)
Earliellascabrosa
PR1209
JN165009
JN164793
Justo and Hibbett (2011)
Etheirodonfimbriatum
Larsson 11905
JN710530
JN710530
Miettinen et al. (2011)
Flabellophora sp.1
X 1357
JN710533
JN710533
Miettinen et al. (2011)
Flabellophora sp.2
X 340
JN710534
JN710534
Miettinen et al. (2011)
Flabellophora sp.3
X 1277
JN710535
JN710535
Miettinen et al. (2011)
Flabellophora sp.4
X 439
JN710536
JN710536
Miettinen et al. (2011)
Flaviporusbrownii
X 1216
JN710537
JN710537
Miettinen et al. (2011)
Flaviporusliebmannii
X 251
JN710541
JN710541
Miettinen et al. (2011)
Flaviporusliebmannii
X 249
JN710539
JN710539
Miettinen et al. (2011)
Flaviporusliebmannii
X 666
JN710540
JN710540
Miettinen et al. (2011)
Fomitopsispinicola
CBS 221.39
DQ491405
DQ491405
Kim et al. (2007)
Fomitopsisrosea
ATCC 76767
DQ491410
DQ491410
Kim et al. (2007)
Fragiliporiafragilis
Dai 13080
KJ734260
KJ734264
Zhao et al. (2015)
Fragiliporiafragilis
Dai 13559
KJ734261
KJ734265
Zhao et al. (2015)
Fragiliporiafragilis
Dai 13561
KJ734262
KJ734266
Zhao et al. (2015)
Frantisekiamentschulensis
BRNM 710170
FJ496728
–
Tomšovský et al. (2010)
Frantisekiamentschulensis
1377
JN710544
JN710544
Miettinen et al. (2011)
Ganodermalingzhi
Wu 1006-38
JQ781858
–
Zhao et al. (2015)
Gelatoporiasubvermispora
BRNU 592909
FJ496694
FJ496706
Tomšovský et al. (2010)
Gloeoporusdichrous
KHL 11173
EU118627
EU118627
Binder et al. (2005)
Grammothelopsissubtropica
Cui 9035
JQ845094
JQ845097
Zhao et al. (2015)
Heterobasidionannosum
PFC 5252
KC492906
KC492906
Binder et al. (2013)
Hornodermoporusmartius
MUCL 41677
FJ411092
FJ393859
Zhao et al. (2015)
Hypochniciumbombycinum
MA 15305
FN552537
–
Binder et al. (2013)
Hypochniciumlyndoniae
NL 041031
JX124704
JX124704
Binder et al. (2005)
Junghuhniacrustacea
X 1127
JN710554
JN710554
Miettinen et al. (2011)
Junghuhniacrustacea
X 262
JN710553
JN710553
Miettinen et al. (2011)
Junghuhniamicropora
Spirin 2652
JN710559
JN710559
Miettinen et al. (2011)
Junghuhnianitida
KHL 11903
EU118638
EU118638
Binder et al. (2005)
Loweomycesfractipes
X 1149
JN710570
JN710570
Miettinen et al. (2011)
Loweomycesfractipes
X 1253
JN710569
JN710569
Miettinen et al. (2011)
Loweomycesfractipes
X 1250
JN710568
JN710568
Miettinen et al. (2011)
Mycoaciafuscoatra
KHL 13275
JN649352
JN649352
Tomšovský et al. (2010)
Mycoacianothofagi
KHL 13750
GU480000
GU480000
Tomšovský et al. (2010)
Nigroporusvinosus
X 839
N710576
N710576
Miettinen et al. (2011)
Nigroporusvinosus
8182
JN710728
JN710728
Miettinen et al. (2011)
Obbarivulosa
KCTC 6892
FJ496693
FJ496710
Miettinen and Rajchenberg (2012)
Obbavaldiviana
FF 503
HQ659235
HQ659235
Miettinen and Rajchenberg (2012)
Panusconchatus
X 1234
JN710579
JN710579
Miettinen et al. (2011)
Panusstrigellus
INPA 243940
JQ955725
JQ955732
Binder et al. (2013)
Perenniporiamedulla-panis
MUCL 49581
FJ411088
FJ393876
Robledo et al. (2009)
Perenniporiellaneofulva
MUCL 45091
FJ411080
FJ393852
Robledo et al. (2009)
Phlebiaunica
KHL 11786
EU118657
EU118657
Binder et al. (2013)
Phlebiaradiata
UBCF 19726
HQ604797
HQ604797
Binder et al. (2013)
Physisporinussanguinolentus
BRNM 699576
FJ496671
FJ496725
Tomšovský et al. (2010)
Physisporinusvitreus
3163
JN710580
JN710580
Miettinen et al. (2011)
Piloporiasajanensis
Mannine 2733a
HQ659239
HQ659239
Miettinen and Rajchenberg (2012)
Podoscyphavenustula
CBS 65684
JN649367
JN649367
Binder et al. (2013)
Polyporustuberaster
CulTENN 8976
AF516598
AJ488116
Binder et al. (2005)
Postiaguttulata
KHL 11739
EU11865
EU11865
Kim et al. (2007)
Pseudolagarobasidiumacaciicola
CBS 115543
DQ517883
–
Miettinen and Rajchenberg (2012)
Pseudolagarobasidiumacaciicola
CBS 115544
DQ517882
–
Miettinen and Rajchenberg (2012)
Pseudolagarobasidiumbelizense
CFMR 04-31
JQ070173
–
Miettinen and Rajchenberg (2012)
Skeletocutisamorpha
Miettinen 11038
FN907913
FN907913
Tomšovský et al. (2010)
Skeletocutisportcrosensis
LY 3493
FJ496689
FJ496689
Tomšovský et al. (2010)
Skeletocutisjelicii
H 6002113
FJ496690
FJ496727
Tomšovský et al. (2010)
Skeletocutisnovae-zelandiae
Ryvarden 38641
JN710582
JN710582
Miettinen et al. (2011)
Spongipellisspumeus
PRM 891931
HQ728287
HQ729021
Tomšovský et al. (2010)
Spongipellisspumeus
BRNM 712630
HQ728288
HQ728288
Tomšovský et al. (2010)
Spongipellisspumeus
BRNM 734877
HQ728283
HQ728283
Tomšovský et al. (2010)
Steccherinumfimbriatum
KHL 11905
EU118668
EU118668
Tomšovský et al. (2010)
Steccherinumochraceum
Ryberg s.n.
EU118669
EU118670
Larsson (2007)
Steccherinumochraceum
KHL 11902
JQ031130
JQ031130
Miettinen et al. (2011)
Stereumhirsutum
NBRC 6520
AB733150
AB733325
Binder et al. (2013)
Truncosporaochroleuca
MUCL 39726
FJ411098
FJ393865
Robledo et al. (2009)
Tyromyceschioneus
Cui 10225
KF698745
KF698756
Zhao et al. (2015)
Xanthoporussyringae
X 339
JN710606
JN710606
Miettinen et al. (2011)
Xanthoporussyringae
Cui 2177
DQ789395
–
Miettinen et al. (2011)
Xanthoporussyringae
Gothenburg 1488
JN710607
JN710607
Miettinen et al. (2011)
Elaphroporiaailaoshanensis
CLZhao 595
MG231568
MG748854
Present study
Elaphroporiaailaoshanensis
CLZhao 596
MG231572
MG748855
Present study
Elaphroporiaailaoshanensis
CLZhao 597
MG231847
MG748856
Present study
Elaphroporiaailaoshanensis
CLZhao 598
MG231823
MG748857
Present study
Phylogenetic analysis. Sequencher 4.6 (GeneCodes, Ann Arbor, MI, USA) was used to edit the DNA sequence. Sequences were aligned in MAFFT 6 (Katoh and Toh 2008, http://mafft.cbrc.jp/alignment/server/) using the “G-INS-I” strategy and manually adjusted in BioEdit (Hall 1999). The sequence alignment was deposited in TreeBase (submission ID 21778). Sequences of Heterobasidionannosum (Fr.) Bref. and Stereumhirsutum (Willd.) Pers. obtained from GenBank were used as outgroups to root trees following Binder et al. (2013) in Figure 1 and Xanthoporussyringae (Parmasto) Audet. obtained from GenBank was used as an outgroup to root trees following Miettinen et al. (2011) in the ITS+nLSU analyses (Fig. 2).
Maximum parsimony strict consensus tree illustrating the phylogeny of Elaphroporiaailaoshanensis and related species in Polyporales based on ITS+nLSU sequences. Branches are labelled with parsimony bootstrap values (before slash) higher than 50 % and Bayesian posterior probabilities (after slash) equal to and more than 0.95. Clade names follow Binder et al. (2013).
Maximum parsimony strict consensus tree illustrating the phylogeny of Elaphroporiaailaoshanensis and related species in the residual polyporoid clade based on ITS+nLSU sequences. Branches are labelled with parsimony bootstrap values (before slash) higher than 50% and Bayesian posterior probabilities (after slash) equal to and more than 0.95. Clade names follow Miettinen et al. (2011).
https://binary.pensoft.net/fig/182395
Maximum parsimony analysis was applied to the ITS+nLSU dataset sequences. Approaches to phylogenetic analysis followed Li and Cui (2013) and the tree construction procedure was performed in PAUP* version 4.0b10 (Swofford 2002). All characters were equally weighted and gaps were treated as missing data. Trees were inferred using the heuristic search option with TBR branch swapping and 1000 random sequence additions. Max-trees were set to 5000, branches of zero length were collapsed and all parsimonious trees were saved. Clade robustness was assessed using a bootstrap (BT) analysis with 1,000 replicates (Felsenstein 1985). Descriptive tree statistics tree length (TL), consistency index (CI), retention index (RI), rescaled consistency index (RC) and homoplasy index (HI) were calculated for each Maximum Parsimonious Tree (MPT) generated. Sequences were also analysed using Maximum Likelihood (ML) with RAxML-HPC2 through the Cipres Science Gateway (www.phylo.org; Miller et al. 2009). Branch support for ML analysis was determined by 1000 bootstrap replicates.
MrModeltest 2.3 (Posada and Crandall 1998, Nylander 2004) was used to determine the best-fit evolution model for each data set for Bayesian Inference (BI). Bayesian Inference was calculated with MrBayes 3.1.2 with a general time reversible (GTR) model of DNA substitution and a gamma distribution rate variation across sites (Ronquist and Huelsenbeck 2003). Four Markov chains were run for 2 runs from random starting trees for 5 million generations (Fig. 1), for 3 million generations (Fig. 2) and trees were sampled every 100 generations. The first one-fourth generations were discarded as burn-in. A majority rule consensus tree of all remaining trees was calculated. Branches that received bootstrap support for maximum likelihood (BS), maximum parsimony (BP) and Bayesian posterior probabilities (BPP) greater than or equal to 75 % (BP) and 0.95 (BPP) respectively, were considered as significantly supported.
Phylogeny results
The ITS+nLSU dataset (Fig. 1) included sequences from 60 fungal specimens representing 52 taxa. The dataset had an aligned length of 2143 characters, of which 1251 characters were constant, 206 parsimony-uninformative and 686 parsimony-informative. MP analysis yielded 6 equally parsimonious trees (TL = 4744, CI = 0.322, HI = 0.678, RI = 0.578, RC = 0.186). The best-fit model for ITS+nLSU alignment estimated and applied in the BI was GTR+I+G, lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). BI resulted in a similar topology with an average standard deviation of split frequencies = 0.001755.
The phylogenetic tree (Fig. 1), inferred from ITS+nLSU sequences, demonstrated seven major clades for 60 sampled species of the Polyporales. The new genus Elaphroporia fell into the Meruliaceae within the residual polyporoid clade. It was closely related to Junghuhniacrustacea (Jungh.) Ryvarden with a good support (95% BS, 89% BP, 0.97 BPP).
The ITS+nLSU (Fig. 2) dataset included sequences from 48 fungal specimens representing 31 taxa. The dataset had an aligned length of 2163 characters, of which 1429 characters were constant, 169 parsimony-uninformative and 565 parsimony-informative. MP analysis yielded 8 equally parsimonious trees (TL = 2806, CI = 0.423, HI = 0.576, RI = 0.673, RC = 0.285). The best-fit model for ITS+nLSU alignment estimated and applied in the BI was GTR+I+G, lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). BI resulted in a similar topology with an average standard deviation of split frequencies equal to 0.005758.
A further phylogeny (Fig. 2) inferred from the combined ITS+nLSU sequences was obtained for 48 fungal specimens representing 31 taxa within the residual polyporoid clade and demonstrated that the new genus formed a monophyletic entity with a high 100 % BS, 100 % BP and 1.00 BPP and sisters to Junghuhniacrustacea and then grouped with Flaviporus and Steccherinum.
Differs from other genera in Polyporales by resupinate basidiocarps becoming rigid and light-weight upon drying, a monomitic hyphal system, thick-walled generative hyphae bearing both clamp connections and simple septa and hyaline, thin-walled, smooth, IKI–, CB– basidiospores.
Type species.
Elaphroporiaailaoshanensis Z.Q. Wu & C.L. Zhao.
Etymology.
Elaphroporia (Lat.): referring to the basidiocarps light-weight upon drying.
Basidiocarps annual, resupinate, becoming rigid and light-weight up on drying. Pore surface cream to pale yellow when fresh, turning to yellow upon drying. Hyphal system monomitic; generative hyphae thick-walled bearing both clamp connections and simple septa, slightly amyloid, CB+. Basidiospores ellipsoid, hyaline, thin-walled, smooth, IKI–, CB–.
This species is distinguished by the cream to yellow pore surface upon drying; pores angular, 7–9 per mm. Hyphal system monomitic; generative hyphae thick-walled bearing both clamp connections and simple septa, slightly amyloid, CB+. Basidiospores ellipsoid, hyaline, thin-walled, smooth, IKI–, CB–, 1.9–2.5 × 1.5–2 µm.
Microscopic structures of Elaphroporiaailaoshanensis (drawn from the holotype). A Basidiospores B Basidia and basidioles C Hyphae from trama D Hyphae from subiculum.
https://binary.pensoft.net/fig/182397Holotype
CHINA. Yunnan Province: Jingdong county, Ailaoshan Nature Reserve, 2 October 2016, on the angiosperm trunk, CLZhao 595 (Holotype in SWFC).
Etymology.
Ailaoshanensis (Lat.): referring to the locality (Ailaoshan) of the type specimens.
Basidiocarps.
Annual, resupinate, easy to separate from substrate, soft corky when fresh, without odour or taste when fresh, becoming rigid and light-weight up on drying, up to 5 cm long, 3.5 cm wide, 4 mm thick at centre. Pore surface cream to pale yellow when fresh, turning to yellow upon drying; pores angular, 7–9 per mm; dissepiments thin, entire. Sterile margin narrow, cream, up to 1 mm wide. Subiculum thin, cream, corky, up to 0.2 mm thick. Tubes concolorous with pore surface, hard corky, up to 3.8 mm long.
Hyphal structure.
Hyphal system monomitic; generative hyphae thick-walled, slightly amyloid, CB+; tissues unchanged in KOH.
Subiculum.
Generative hyphae hyaline, thick-walled bearing both clamp connections and simple septa, simple septa more frequent than clamps, occasionally branched, interwoven, 3.5–5.5 µm in diam.
Tubes.
Generative hyphae hyaline, thick-walled bearing simple septa only, occasionally branched, 3–5 µm in diameter. Cystidia and cystidioles absent; basidia clavate, with four sterigmata and a basal clamp connection, 10.5–14.5 × 3.5–4.5 µm; basidioles dominant, in shape similar to basidia, but slightly smaller.
(paratypes). CHINA. Yunnan Province: Jingdong county, Ailaoshan Nature Reserve, 2 October 2016, on the angiosperm trunk, CLZhao 596, CLZhao 597, CLZhao 598 (SWFC).
Discussion
In the present study, a new genus, Elaphroporia, is described based on phylogenetic analyses and morphological characters. The genus has unique morphological characters in Meruliaceae.
Previously, seven clades were found in the Polyporales: antrodia clade, core polyporoid clade, fragiliporia clade, gelatoporia clade, phlebioid clade, residual polyporoid clade and tyromyces clade (Binder et al. 2013, Zhao et al. 2015). According to these results based on the combined ITS+nLSU sequence data (Fig. 1), the new genus is nested into the residual polyporoid clade with strong support (100 % BS, 100 % BP, 1.00 BPP).
Miettinen et al. (2011) analysed a higher-level phylogenetic classification of the residual polyporoid clade morphological plasticity in a group of the polypores, and showed that the natural genera could mostly be characterised morphologically and poroid and hydnoid species belong to separate genera. The current phylogeny shows that the genus Elaphroporia falls into the residual polyporoid clade and belongs to the family Meruliaceae (Figs 1, 2). Furthermore, the new genus is closely related to Junghuhnia and then grouped with Flaviporus and Steccherinum based on ITS+LSU-nrRNA gene regions with a strong support (100 % BS, 100 % BP, 1.00 BPP; Fig. 1). However, morphologically Junghuhnia differs from Elaphroporia by a dimitic hyphal system and presence of cystidia (Núñez and Ryvarden 2001, Ryvarden and Melo 2014). Flaviporus is separated from Elaphroporia by the dark brown to bay pileus, a dimitic hyphal system and presence of the metuloid cystidia (Murrill 1905). Steccherinum differs in its odontioid to hydnoid hymenophore and cyanophilous basidiospores (Bernicchia and Gorjón 2010).
Morphologically, Elaphroporia resembles Ceriporia Donk and Phlebiporia Jia J. Chen, B.K. Cui & Y.C. Dai. Ceriporia is similar to Elaphroporia in an annual growth habit with poroid hymenophore, a monomitic hyphal structure and hyaline, thin-walled and smooth basidiospores. In addition, both genera cause a white rot. However, Ceriporia differs from Elaphroporia by the generative hyphae IKI–, CB– (Jia et al. 2014). Additionally, in molecular studies, Ceriporia fell into the phlebia clade (Miettinen and Larsson 2011, Miettinen and Rajchenberg 2012, Miettinen et al. 2011, Binder et al. 2013) which is also the same as in the authors’ study (Fig. 1). Phlebiporia is similar to Mellipora by having the poroid hymenophore and the generative hyphae bearing both simple septa and clamp connections, but it is separated from Elaphroporia by having dextrinoid generative hyphae, tissues becoming brownish in KOH and presence of thin-walled quasi-binding hyphae in the subiculum (Chen and Cui 2014).
Polypores are an extensively studied group of Basidiomycota (Gilbertson and Ryvarden 1987, Núñez and Ryvarden 2001, Dai 2012, Ryvarden and Melo 2014), but the Chinese polypore diversity is still not well known, especially in subtropics and tropics, from where many recently described taxa of polypores were discovered (Song et al. 2014, 2016, Zhou et al. 2015, 2016, Nie et al. 2017, Yuan et al. 2017). The new genus in the present study, Elaphroporia, is also from the subtropics. It is possible that new polypore taxa will be found after further investigations and molecular analyses.
Acknowledgments
We express our gratitude to Yong-He Li (Yunnan Academy of Biodiversity, Southwest Forestry University, P.R. China) for his support on molecular work. The research is supported by the National Natural Science Foundation of China (Project No. 31700023) and the Science Foundation of Southwest Forestry University (Project No. 111715) and the Science and Technology Talent Support Programme of Three Areas in Yunnan Province (Project No. 21700329).
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