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Article

Poriella subacida Gen. & Comb Nov. for Perenniporia subacida (Peck) Donk

1
Key Laboratory for Forest Resources Conservation and Utilization in the Southwest Mountains of China, Ministry of Education, Southwest Forestry University, Kunming 650224, China
2
College of Biodiversity Conservation, Southwest Forestry University, Kunming 650224, China
3
CAS Key Laboratory for Plant Biodiversity and Biogeography of East Asia (KLPB), Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China
*
Author to whom correspondence should be addressed.
Agronomy 2021, 11(7), 1308; https://doi.org/10.3390/agronomy11071308
Received: 11 May 2021 / Revised: 20 June 2021 / Accepted: 22 June 2021 / Published: 27 June 2021
(This article belongs to the Special Issue Research Progress on Added-Value Fungi)

Abstract

:
Poriella subacida gen. & comb. nov., previously known as Perenniporia subacida, which causes white rot, has been documented in temperate and tropical forests. Specimens from Asia, North America, and Europe were examined, including the type specimen of Polylorus subacidus. Sequences of the ITS1-5.8S-ITS2 region, the 28S rDNA, the mitochondrial rDNA small subunit (mtSSU), and the gene encoding the translation elongation factor 1-α (EF1) were generated. In multigene phylogenies (maximum parsimony, maximum likelihood, Bayesian inferences), “Perenniporia subacida” formed a well-supported lineage, distinct from the core “Perenniporia” clade (type species: “P. medulla-panis”), and sister to the “Yuchengia narymica” lineage. We therefore conclude that “P. subacida” should be placed in the new genus “Poriella”gen. nov. Morphologically, “Poriella” is characterized by a di- to trimitic hyphal system, non-truncate basidiospores, and strongly dextrinoid, cyanophilic skeletal hyphae.

1. Introduction

Polyporales is one of the most intensively studied clades of fungi, being of interest to both fungal ecologists and applied scientists [1,2]. The family Polyporaceae accommodates 92 genera [3], and recently many new genera of polypores have been described, such as Amylosporia (B.K. Cui, C.L. Zhao & Y.C. Dai), Crystallicutis (El-Gharabawy and Griffith), Hirticrusta Matozaki (T. Hatt. and Sotome), and Murinicarpus (B.K. Cui and Y.C. Dai) [4,5,6,7]. The family is typified by Polyporus P. Micheli ex Adans. (1763) and many taxa have been separated from Polyporus as new genera over time [1,4,6].
Perenniporia subacida (Peck) Donk (basionym Polyporus subacidus Peck, 1885) is a widespread species of wood-decaying polypores with perennial, resupinate to effused-reflex basidiomes and thick-walled, non-truncate basidiospores. The species has been reported to occur in many forest ecosystems, in boreal, temperate, subtropical, and tropical regions [8,9,10,11,12,13,14,15]. This species was described by Peck (1885) as Polyporus subacidus, but its taxonomic position has long been debated. It has been treated in several genera, but none of these placements have been fully satisfactory [16,17,18,19,20]. Currently, the species is widely accepted in Perenniporia Murrill, which is typed by P. medulla-panis (Jacq.) Donk [8,9,10,11,12,13]. However, Decock and Stalpers argued for a different placement [13].
Molecular studies involving Polyporaceae, mainly based on ITS and/or nLSU sequences, have been carried out [21,22,23,24,25,26,27]. Further studies employing a six-gene dataset have helped to clarify the generic relationships of polyporoid fungi for 373 taxa. The latter study showed that Perenniporia subacida clustered in the core polyporoid clade in which it was related to P. medulla-panis (Jacq.) Donk [23]. Further studies of Perenniporia inferred from nuclear ribosomal 28S and ITS sequence data revealed that P. subacida formed a monophyletic lineage distant from the Perenniporia s.s. [15,16,28].
To resolve the placement of this species, phylogenetic research was carried out employing the ITS, 28S, TEF1 and mtSSU and a comparative morphological study of the type specimen was conducted. We conclude that Perenniporia subacida should be treated as a distinct, new genus as described below.

2. Materials and Methods

2.1. Morphological Studies

Specimens studied are deposited at the Farlow Herbarium of Harvard University (FH), MA, USA, Beijing Forestry University (BJFC), Beijing, China and United States National Fungus Collections (BPI), New York, NY, USA. Macromorphological descriptions were on the basis of field notes and study of specimens. Color terms followed previous studies [29]. Microscopic measurements were made from slide preparations of dried specimens stained with Cotton Blue and Melzer’s reagent by light microscopy [30,31]. Sections were studied using an Olympus BX40 compound microscope (Tokyo, Japan). In presenting spore size variation, 5% of measurements were excluded from each end of the range. The following abbreviations are used: KOH = 5% potassium hydroxide, CB = Cotton Blue, CB+ = cyanophilous; IKI = Melzer’s reagent, IKI– = both non-amyloid and non-dextrinoid, L = mean spore length (arithmetic average of all spores), W = mean spore width (arithmetic average of all spores), Q = mean each spore length/width ratio, n (a/b) = number of spores (a) measured from given number of specimens (b).

2.2. Molecular Techniques and Phylogenetic Analyses

CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to extract DNA from dried specimens from China and other regions, and 2xTaq PCR Mix (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to perform PCR products [30]. The primer pair ITS5 and ITS4 were amplified for the ITS region [32]. The primer pair LR0R and LR7 were for the nuclear r28S region [33]. The primer pair MS1 and MS2 were for the mitochondrial SSU region [32]. The primer pair EF1-983F and EF1-2218R were for Tef1 [34]. The PCR cycling program for ITS, mtSSU, 28S and TEF1 followed previous studies [27]. The Beijing Genomics Institute (Beijing, China) was in charge of purifying and direct sequencing of the PCR products. All sequences used in the phylogeny were downloaded from GenBank (Table 1) with references. The sequence alignment was deposited in TreeBase (submission ID 23826).
A previous study [30] was followed for maximum parsimony analysis. The combined multiple genes dataset was analyzed under heuristic search and 1000 homogeneity replicates, giving a P value of 1.000, which is much greater than the 0.01 used in PAUP* version 4.0b10, which means there is no discrepancy among the four loci in reconstructing phylogenetic trees. Trees were constructed in PAUP* version 4.0b10 [44]. 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 bootstraps. 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 1000 replicates [45]. DNA sequence data was also analyzed using Maximum Likelihood (ML) with RAxML-HPC2 on Abe through the Cipres Science Gateway [46], with default settings except that branch support was obtained with 1000 rapid bootstrap replicates.
Mr Modeltest 2.3 was used to estimate the best-fit evolution model for each data set for Bayesian inference (BI). The best fit models were general time reversible (GTR)+G for ITS, and general time reversible (GTR)+I+G for nr28S, mtSSU, the exons of Tef1, and the combined dataset. The partitioned mixed model, which allows for model parameters to be estimated separately for each genetic marker, was used in the Bayesian analysis. BI was performed using MrBayes 3.1.2 [47,48,49]. Two runs of four Markov chains were run from random starting trees for two datasets: (ITS+28S) dataset for 5 million generations and (ITS+28S+mtSSU+Tef1) dataset for 10 million generations. Trees were sampled every 100 generations. The first quarter of the generations was by default discarded as burn-in. A majority rule consensus tree of all remaining trees was calculated. Branches that received bootstrap support for maximum likelihood (ML-BS), maximum parsimony (MP-BT), and Bayesian posterior probabilities (BPP) greater than or equal to 75% (ML-BS and MP-BS) and 0.95 (BPP) were considered significantly supported.

3. Results

3.1. Molecular Phylogeny

The ITS+28S dataset included sequences from 63 fungal specimens representing 56 taxa. The dataset had an aligned length of 2112 characters, of which 1216 characters are constant, 256 are variable and parsimony-uninformative, and 640 are parsimony-informative. Maximum parsimony analysis yielded 12 equally parsimonious trees (TL = 4377, CI = 0.337, HI = 0.664, RI = 0.525, RC = 0.177). The best model for the ITS+28S dataset estimated and applied in the Bayesian analysis is general time reversible (GTR)+I+G, lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). Bayesian analysis and ML analysis resulted in a similar topology to the MP analysis, with an average standard deviation of split frequencies = 0.005569 (BI).
The phylogeny (Figure 1) inferred from ITS+28S sequences resolves seven major clades for 56 species of the Polyporales. Collections of Perenniporia subacida formed a lineage distinct from the Perenniporia s.s. lineage, within the core polyporoid clade. The P. subacida lineage is closely related to the Yuchengia lineage based on Y. narymica (Pilát) (B.K. Cui, C.L. Zhao, and Steffen) with good support (100% BS, 100% BP, 1.00 BPP).
The four gene (ITS+28S+mtSSU+Tef1) sequence dataset did not show any conflicts in tree topology for the reciprocal bootstrap trees, which allowed us to combine them. The combined dataset included sequences from 61 specimens representing 29 species. The dataset had an aligned length of 3515 characters, of which 2489 characters are constant, 219 are variable and parsimony-uninformative, and 807 are parsimony-informative. Maximum parsimony analysis yielded 10 equally parsimonious trees (TL = 3075, CI = 0.498, HI = 0.502, RI = 0.747, RC = 0.372). Best model for the combined ITS+28S+mtSSU+Tef1 estimated and applied in the Bayesian analysis: general time reversible (GTR)+I+G, lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). Bayesian analysis and ML analyses resulted in a topology similar to the MP analysis, with an average standard deviation of split frequencies = 0.003420.
A further phylogeny (Figure 2) inferred from the combined ITS+28S+mtSSU+Tef1 sequences was obtained for ten genera in Perenniporia s.l. and demonstrated that this taxon formed a clade together with Y. narymica, with strong support (100% BS, 100% BP, 1.00 BPP). The clade is distinct from P. medulla-panis (Jacq.) We, along with Donk, conclude that it belongs to a distinct, new genus, hereafter called Poriella.

3.2. Taxonomy

Poriella C.L. Zhao, gen. nov.
MycoBank: 840061.
It is characterized by producing resupinate to effused-reflex basidiomata with dingy-yellowish to pale tan to ochraceous surface and a di-trimitic hyphal system with unbranched and strongly dextrinoid skeletal hyphae, and thick-walled, non-dextrinoid, cyanophilous basidiospores.
Type species: Poriella subacida (Peck) C.L. Zhao.
Etymology: Poriella (Lat.): referring to its similar appearance to the genus Perenniporia.
Basidiomata perennial, resupinate to effused-reflex. Pore surface is dingy-yellowish to pale tan to ochraceous. Pores round to angular. Subiculum cream to buff, thin. Tubes concolorous with pore surface, corky. Hyphal system di-trimitic, generative hyphae hyaline, thin-walled, with clamp connections; skeletal hyphae predominant, unbranched, strongly dextrinoid, cyanophilous, rarely dissolving in KOH. Basidiospores ellipsoid, non-truncate, hyaline, thick-walled, smooth, non-dextrinoid, CB+.
  • Poriella subacida (Peck) C.L. Zhao, comb. nov. Figure 3 and Figure 4.
  • MycoBank: 840062.
  • Basionym: Polyporus subacidus Peck, Ann. Rep. N.Y. St. Mus. nat. Hist. 38: 92, 1885.
  • =Poria subacida (Peck) Sacc., Syll. fung. (Abellini) 6: 325 (1888).
  • =Chaetoporus subacidus (Peck) Bondartsev & Singer, Annls mycol. 39(1): 51 (1941).
  • =Oxyporus subacidus (Peck) Komarova, Mycoth. Eston. 3: 13 (1961).
  • =Perenniporia subacida (Peck) Donk, Persoonia 5(1): 76 (1967).
  • =Poria colorea Overh. & Englerth, Bull. Yale Univ. School For. 50: 21 (1942).
  • =Poria fuscomarginata Berk. ex Cooke, Grevillea 15(no. 73): 24 (1886).
  • =Poria subaurantia Berk. ex Cooke, Grevillea 15(no. 73): 27 (1886).
Fruiting body: Perennial, resupinate to effused-reflex, becomes corky when dried, about 22 cm or more at the longest dimension, 15 cm or more at the widest dimension, and up to 17 mm thick at the center. Pore surface pale yellowish to dingy-yellowish when fresh, dingy-yellowish to pale tan to ochraceous when dry; pores round to angular, 4–6 per mm; dissepiments thin, entire. Subiculum thin, cream to buff, up to 1 mm thick. Tubes concolorous with pore surface, up to 16 mm thick.
Hyphal structure: Hyphal system di-trimitic; generative hyphae with clamp connections; skeletal hyphae strongly dextrinoid, CB+; dissolving in KOH.
Context: Generative hyphae infrequent, hyaline, thin-walled, 2.5–4 µm in diameter; skeletal hyphae dominant, thick-walled with a wide lumen, unbranched, subparallel, 3–5.5 µm in diameter; skeletal-binding hyphae hyaline, thick-walled, frequently branched, flexuous, interwoven, 1–2 µm in diameter.
Tubes: Generative hyphae infrequent, hyaline, thin-walled, 2.5–3.5 µm in diameter; skeletal hyphae dominant, thick-walled with a wide lumen, unbranched, subparallel, 3–4.5 µm in diameter; skeletal-binding hyphae hyaline, thick-walled, frequently branched, flexuous, interwoven, 0.7–1.7 µm in diameter. Cystidia absent, but fusoid cystidioles present, hyaline, thin-walled, 13–16 × 3–4.5 µm. Basidia barrel-shaped, with four sterigmata and a basal clamp connection, 20–22.5 × 7–8 µm; basidioles similar in shape to basidia, but slightly smaller.
Basidiospores: Ellipsoid, not truncate, hyaline, thick-walled, smooth, non-dextrinoid, CB+, (4.2–)4.5–6.2(–6.4) × (3.2–)3.6–4.6(–4.9) µm, L = 5.25 µm, W = 3.92 µm, Q = 1.13–1.42 (n = 450/15).
Associated wood-rot: White.
Substrates and distribution: Mainly on conifers, but also on hardwood, causes white rot in Abies Mill and Tsuga Carr. and common on dead fallen trees in many areas. A boreal eastern species in Europe, widely distributed in forest regions of Asia and North America [8,10,11].
Additional specimen examined: CANADA, Ontario, Gull lake, on Thuja occidentalis L., July 25 1919, J.H. Faull (5088) (FH 00605379); Lake Timagami, Timagami Is., on fallen trunk of Abies balsamea (L.) Mill., September 10 1918, J.H. Faull (3365) (FH 00605372); Lake Rosseau, on Tsuga Carr., September 1902, Harper (587) (FH 00605380); Lake Rosseau, on Pinus Linn log, October 1903, S.A. Haper (832) (FH); Lake Rosseau, on fallen log of Abies balsamea (L.) Mill., 18 August 1921, J.H. Faull (6013) (FH 00605381); Humber Valley, on roots of Thuja occidentalis L., September 1914, J.H. Faull (169) (FH 00605373); Humber Valley, on dead hardwood, September 1914, J.H. Faull (156) (FH 00605375); Sudbury District, Cleland Tp., on dead coniferous wood, 15 September 1918, J.H. Faull (3502) (FH 00605377). CHINA, Fujian Province, Wuyishan Nature Reserve, on fallen angiosperm trunk, 21 October 2005, Dai 7316 (IFP); Guizhou Province, Jiangkou County, Fanjingshan Nature Reserve, on fallen angiosperm trunk, 21 August 2010, Yuan 5511 (IFP); Heilongjiang Province, Yichun, Fenglin Nature Reserve, on fallen trunk of Picea Dietr., 2 August 2011, Cui 9849, 9853 (BJFC); Yunnan Province, Baoshan, Gaoligong Mountain, on angiosperm trunk, 23 September 2007, Yuan 3850, 3854 (IFP); Zhejiang Province, Linan, Tianmushan Nature Reserve, on fallen trunk of Picea Dietr., 11 October 2005, Cui 2705 2712 (BJFC). FINLAND, Pera-Pohjanma, Pisavaara National Park, on fallen trunk of Picea Dietr., 14 September 1997, Dai 2648 (BJFC); Pisavaara National Park, on Picea Dietr., 4 November 2011, Dai 12619 (BJFC). USA, Arizona, Coronado National Forest, on conifer, 28 August 1958, Lowe (9407) (FH 00605393); Connecticut, New Haven, Sleeping Giant State Park, on fallen trunk of Betula L., 22 July 2012, Dai 12773 (BJFC); as above, on fallen trunk of Tsuga Carr., 22 July 2012, Dai 12785 (BJFC); as above, 24 July 2012, Dai 10287 (BJFC); as above, on fallen trunk of Pinus Linn, 24 July 2012, Dai 10285 (BJFC); Idaho, Bovill County, on dead fallen trunk of Pinus Linn, 2 October 1920, A.S. Rhoads (15891)(FH 00605391); Priest River, on Picea engelmanni, September 1915, J.R. Weir (9551) (FH 00605392); Priest River, on Abies grandis (Dougl ex D Don) Lindl, September 1915, J.R. Weir (8152) (FH 00605394); Kansas, Rooks County, Rockport, on underside of old log, 9 December 1893, (1315) (FH 00605389); Maine, Norcross, on fallen Populus L. log, 20 August 1940, D.H. Linder s.n. (FH 00605344), Kittery Point, on log of Quercus L., 28 July 1922, J.R. Weir (897)(FH 00605347); Linekin, on Picea Dietr., September 1899, Burt (2125B) (FH); as above, 16 August 1899, Burt 2127 (FH); Massachusetts, Norfolk County, Sharon, 28 June 1946, A.P.D.Piguet (54) (FH 00605353); Michigan, Neebish, September 1911, E.T.Harper & S.A.Harper s.n. (FH 00605383); Isle Royal, Rock Harbor, August 1904, E.T.Harper & S.A.Harper s.n. (FH 00605384); Pleasant Ridge, on fallen Tsuga Carr., 25 September 1915, P.Spaulding & J.F.Collins s.n. (FH 00605345); New York, Osceola, on Tsuga Carr., August 1885, C.H. Peck, BPI 844697 (Holotype, BPI); as above, on Picea Dietr. log, FH 0053717 (FH); Dryden, Ringwood Preserve, on old wood, 9 June 1952, W.B.Cooke & V.G.Cooke (29105) (FH 00605356); Ithaca, on Betula lutea F.Michx., nom. illeg. 21 March 1935, W.L. White (1568) (FH 00605360); Floodwood, on conifer, August 1900, Burt 2122B (FH); Floodwood, on Picea Dietr. log, 20 August 1900, 2123B (FH); Pennsylvania: Bedford County, New Paris, on fallen Quercus L., 1 September 1916, J.H. Faull (1490) (FH 00605366); Lackawanna County, Carbondale, on Tsuga Carr. log, 13 December 1898, Burt 2124C (FH); Tennessee, Knox County, Ball Camp Pike, on Pinus Linn log, 18 December 1938, L.R. Hesler (11907) (FH 00605387); Vermont: Addison County, Ripton, on hardwood log, 31 October 1896, (FH 00605346); Addison County, Ripton, on Picea Dietr., 4 November 1896, Burt 2126A (FH); Addison County, Ripton, on Picea Dietr., 31 October 1896, Burt 2126B (FH); Abby, on Tsuga Carr. log, 31 October 1896, Burt 2123A (FH); Underhill, on hardwood, 18 May 1880, Pringle (1025) (FH 00605350); as above, on dead conifer, August 1897, Burt 2121A, B, C (FH); as above, on fallen trunk of Tsuga Carr., September 1902, Harper 625 (FH); Dunamase, on Tsuga Carr. log, 14 September 1896, Burt 2121E (FH); Dunamase, on Tsuga Carr., October 1896, Burt 2122A (FH); Snake Mountain, January 1896, 2124A (FH); as above, September 1902, Harper 746 (FH); Washington: Marymere Falls, Olympic National Park, on Pseudotsuga taxifolia (Lamb.) Britton, WB (27590) (FH 00605395); Clallam County, Sol Duc Hot Springs, on fallen Tsuga heterophylla (Raf.) Sarg., 4 July 1920, J.R. Weir (650) (FH 00605396); as above, on fallen trunk of Picea sitchensis (Bong.) Carr., 14 July 1920, J.R. Weir (609) (FH 00605397); Wisconsin, June 21 1946, Neuman (162) (FH 00605386).

4. Discussion

The taxonomic position of this taxon has been long debated [16,17,18,19,20]. It was transferred to Perenniporia by Donk [20], and this placement has been generally accepted [8,9,10,11,12]. Due to a combination of characteristics (completely unbranched skeletal hyphae and ellipsoid and non-truncate basidiospores), Decock and Staplers [13] mentioned that the species did not belong to Perenniporia. Based on phylogenetic and morphological grounds, Robledo et al. confirmed that this taxon should be separated from Perenniporia, and could be recognized as a distinct genus morphologically as well [28]. Previous studies also confirmed that this species formed a clade distinct from the P. medulla-panis clade [5,6,28]. The present study confirms these results from previous studies and formally describes the new genus, Poriella with P. subacida as type species.
Other species of Perenniporia and Poriella subacida share the morphological features of unbranched skeletal hyphae and non-truncate basidiospores, such as Perenniporia africana (Ipulet and Ryvarden) in Uganda [50,51], P. contraria (Berk. and M.A. Curtis) Ryvarden in Cuba, and P. ellipsospora (Ryvarden and Gilb.) in North America [10]. A comparison of Poriella and related genera is presented in Table 2.
Three species have been mentioned as possible synonyms of this highly variable species. Poria colorea Overh. & Englerth, described from Western Tsuga Carr. and has generally been considered to be conspecific with Polyporus subacidus. Poria fuscomarginata Berk. ex Cooke was commented on by Murrill [52] who found the type material to be badly preserved and scanty. He concluded that “it was suggested little.” Poria subaurantia Berk. ex Cooke was considered a synonym of Polyporus subacidus by Murrill [53]. Judging by their descriptions, these fungi show a range of variation that is acceptable for P. subacidus.
Our molecular phylogenetic analyses revealed that the genus Poriella is closely related to Yuchengia narymica and then grouped with Vanderbylia based on the ITS+28S gene regions and the combined ITS+28S+mtSSU+Tef1 sequences (Figure 1 and Figure 2). Morphologically, two species share the same hyphal system and basidiospores. However, Y. narymica differs markedly from the acyanophilous, amyloid skeletal hyphae [38,53,54,55]. Vanderbylia mainly differs in its pileate basidiomata, a dimitic hyphal system with distinctly arboriform vegetative hyphae in the hymenophoral trama and obovoid basidiospores [10,56].
Poriella subacida is primarily a boreal taxon and is widely distributed in forest regions of northern Asia, North America and Europe [8,10,11]. It mainly grows on conifers, especially Picea Dietr., but also on Larix and Pinus Linn. In Europe, it has also been found occasionally on hardwoods like Populus L. and Prunus L. [8], and other hardwoods in North America and Asia [10,11]. The species is also present in tropical areas (e.g., in Africa) [57], but these records should be treated with caution. Ipulet and Ryvarden [50] recently described Perenniporia africana Ipulet and Ryvarden, from Uganda, with seemingly the same combination of characteristicss, i.e., unbranched skeletal hyphae and non-truncate basidiospores [57,58]. A list of characteristics of Poriella subacida comb. nov. from different regions is presented in Table 3.
Poriella subacida causes white rot of conifers and hardwoods and also butt and root rots of living conifers. Due to the cream to golden yellow mycelia felts that develop in the decayed wood, this rot is commonly called “feather rot” [10].

Author Contributions

Conceptualization, C.-L.Z. and R.C.; methodology, C.-L.Z. and R.C.; software, R.C.; validation, R.C., C.-L.Z. and S.C.K.; formal analysis, R.C.; resources, C.-L.Z.; data curation, R.C.; writing—original draft preparation, R.C.; writing—review and editing, C.-L.Z. and S.C.K.; visualization, R.C.; supervision, C.-L.Z. and S.C.K.; project administration, C.-L.Z.; funding acquisition, C.-L.Z. and S.C.K. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the Yunnan Fundamental Research Project (Grant No. 202001AS070043) and the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111).

Data Availability Statement

Publicly available datasets were analyzed in this study. This data can be found here: https://www.ncbi.nlm.nih.gov/; https://www.mycobank.org/page/Simple%20names%20search; http://purl.org/phylo/treebase, submission ID 23826; accessed on 25 June 2021.

Acknowledgments

Special thanks are conveyed to Yu-Cheng Dai (BJFU, China), Bao-Kai Cui (BJFU, China), and Donald H. Pfister (Harvard University) for allowing me to study their specimens.

Conflicts of Interest

The authors declare no conflict of interest. All the experiments undertaken in this study comply with the current laws of the People’s Republic of China.

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Figure 1. Maximum parsimony strict consensus tree illustrating the phylogeny of Poriella subacida and related species in Polyporales based on ITS+28S sequences. Branches are labeled with maximum likelihood bootstrap higher than 70%, parsimony bootstrap proportions higher than 50% and Bayesian posterior probabilities more than 0.95, respectively. Clade names [23].
Figure 1. Maximum parsimony strict consensus tree illustrating the phylogeny of Poriella subacida and related species in Polyporales based on ITS+28S sequences. Branches are labeled with maximum likelihood bootstrap higher than 70%, parsimony bootstrap proportions higher than 50% and Bayesian posterior probabilities more than 0.95, respectively. Clade names [23].
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Figure 2. Maximum Parsimony strict consensus tree illustrating the phylogeny of Poriella subacida, and related species in Perenniporia s.l. based on the combined ITS+28S+mtSSU+TEF1 sequence datasets. Branches are labeled with maximum likelihood bootstrap higher than 70%, parsimony bootstrap proportions higher than 50% and Bayesian posterior probabilities more than 0.95, respectively. Clade names follow [15].
Figure 2. Maximum Parsimony strict consensus tree illustrating the phylogeny of Poriella subacida, and related species in Perenniporia s.l. based on the combined ITS+28S+mtSSU+TEF1 sequence datasets. Branches are labeled with maximum likelihood bootstrap higher than 70%, parsimony bootstrap proportions higher than 50% and Bayesian posterior probabilities more than 0.95, respectively. Clade names follow [15].
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Figure 3. Basidiomata of Poriella subacida (Holotype, BPI 844697). Scale bar = 1 cm.
Figure 3. Basidiomata of Poriella subacida (Holotype, BPI 844697). Scale bar = 1 cm.
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Figure 4. Microscopic structures of Poriella subacida (Holotype). (a) Basidiospores. (b) Basidia and basidioles. (c) Cystidioles. (d) Hyphae from trama. (e) Hyphae from context. Bars: A = 5 µm; B–E = 10 µm.
Figure 4. Microscopic structures of Poriella subacida (Holotype). (a) Basidiospores. (b) Basidia and basidioles. (c) Cystidioles. (d) Hyphae from trama. (e) Hyphae from context. Bars: A = 5 µm; B–E = 10 µm.
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Table 1. Information of the sequences used in this study.
Table 1. Information of the sequences used in this study.
Species NameSample No.GenBank AccessionsReferences
ITSLSUmtSSUTEF1
Abortiporus biennisTFRI 274EU232187EU232235 [21]
A. biennisEL65-03JN649325JN649325 [21]
Abundisporus roseoalbusDai 12269KC415908KC415910KF051037KF181131[5]
A. pubertatisCui 5776KC787565KC787572KF051029KF181129[5]
A. sclerosetosusMUCL 41438FJ411101FJ393868 [28]
A. violaceusRyvarden 10775KF018126KF018134KF051058KF181152[5]
Amylocystis lapponicaKHL 11755EU118603EU118603 [21]
Antrodia albidaCBS 308.82DQ491414 [35]
A. albidaFP 105979EU232272EU232272 [36]
A. heteromorphaCBS 200.91DQ491415AY515350 [35]
A. macraMUAF 887EU340898 [23]
Bjerkandera adustaNBRC 4983AB733156AB733333 [23]
Cinereomyces lindbladiiFBCC 177HQ659223HQ659223 [23]
Climacocystis borealisKH 13318JQ031126JQ031126 [23]
Coriolopsis caperataLE(BIN)-0677AB158316AB158316 [37]
Donkioporia expansaMUCL 35116FJ411104FJ393872 [28]
Earliella scabrosaPR1209JN165009JN164793 [38]
Fragiliporia fragilisDai 13080KJ734260KJ734264KJ734268KJ790245[5]
F. fragilisDai 13559KJ734261KJ734265KJ734269KJ790246[5]
F. fragilisDai 13561KJ734262KJ734266KJ734270KJ790247[5]
Ganoderma sichuanenseWu 1006-38JQ781858 JX029989JX029976[5]
G. sichuanenseDai 12479JQ781864 JX029988JX029975[5]
G. australeCui 9511JN048773JN048792 [39]
G. sinenseWei 5327KF494998KF495008 KF494976[5]
G. applanatumDai 12483KF494999KF495009 KF494977[5]
Gelatoporia subvermisporaBRNU 592909FJ496694FJ496706 [37]
Grammetheliopsis subtropicaCui 9041JQ845096JQ845099KF051039KF181133[39]
Heterobasidion annosumPFC 5252KC492906KC492906 [23]
Hornodermoporus latissimaCui 6625HQ876604JF706340KF051040KF181134[5]
H. martiusCui 7992HQ876603HQ654114KF051041KF181135[5]
H. martiusMUCL 41677FJ411092FJ393859 [28]
H. martiusMUCL 41678FJ411093FJ393860 [28]
Hydnopolyporus fimbriatusLR 40855JN649347JN649347 [23]
Hypochnicium lyndoniaeNL 041031JX124704JX124704 [23]
Lentinus tigrinusDSH93-181AY218419AF518627U27050 [38]
Microporellus violaceocinerascensMUCL 45229FJ411106FJ393874 [28]
M. violaceocinerascensCui 8459HQ876606HQ654113KF051042KF181136[5]
Obba rivulosaKCTC 6892FJ496693FJ496710 [4]
Perenniporia hainanianaCui 6364JQ861743JQ861759KF051044KF181138[5]
P. hainanianaCui 6365JQ861744JQ861760KF051045KF181139[5]
P. hainanianaCui 6366JQ861745JQ861761KF494996KF494981[5]
P. medulla-panisMUCL 49581FJ411088FJ393876 [28]
P. medulla-panisMUCL 43250FJ411087FJ393875 [28]
P. medulla-panisCui 3274JN112792JN112793KF051043KF181137[5]
P. substramineaCui 10177JQ001852JQ001844KF051046KF181140[5]
P. substramineaCui 10191JQ001853JQ001845KF051047KF181141[5]
P. substramineaDai 10781KF495007KF495018KF494995KF494983[5]
Perenniporiella chaqueniaMUCL 47647FJ411083FJ393855 HM467609[28]
P. chaqueniaMUCL 47648FJ411084FJ393856 HM467610[28]
P. microporaMUCL43581FJ411086FJ393858 HM467608[28]
P. neofulvaMUCL 45091FJ411080FJ393852 HM467599[28]
P. pendulaMUCL 46034FJ411082FJ393853 HM467601[28]
Phanerochaete chrysosporiumBKM-F-1767HQ188436GQ470643 [37]
Phlebia unicaKHL 11786EU118657EU118657 [37]
Physisporinus sanguinolentusBRNM 699576FJ496671FJ496725 [37]
Piloporia sajanensisMannine 2733aHQ659239HQ659239 [37]
Podoscypha venustulaCBS 65684JN649367JN649367 [23]
Polyporus tuberasterCulTENN 8976AF516598AJ488116 [40]
Poriella subacidaDai 8224HQ876605JF713024KF218322KF286328[15]
P. subacidaCui 3643FJ613655AY336753KF218320KF286326[15]
P. subacidaCui 10053KF495006KF495017KF218321KF286327[15]
P. subacidaMUCL 31402FJ411103FJ393880 [28]
P. subacidaCBS 463.50FJ805245 Direct submission
P. subacidaDLL 2009-125JQ673136 [41]
P. subacidaDLL 2009-150JQ673014 [41]
P. subacidaDLL 2009-154JQ673015 [41]
P. subacidaDai 8859FJ613656 [15]
P. subacidaHHb-14877-TAY089739AY089739 Direct submission
P. subacidaB 37AF218403 [42]
Postia alniX 1400KC595932KC595932 [23]
P. caesiaCIEFAP 174JX090109JX090129 [23]
P. guttulataKHL 11739EU11865EU11865 [43]
P. venataCIEFAP 346JX090113JX090133 [43]
P. lacteaX 1391KC595939KC595939 [23]
Pyrofomes demidoffiiMUCL 41034FJ411105FJ393873 [28]
Sebipora aquosaMiettinen 8680HQ659240HQ659240 [22]
Skeletocutis amorphaMiettinen 11038FN907913FN907913 [37]
Stereum hirsutumNBRC 6520AB733150AB733325 [23]
Trametes elegansFP105679JN048766JN048785 [15]
T. hirsutaCui 7784JN048768JN048787 [15]
T. hirsutaRLG5133TJN164854JN164801AF042154JN164891[38]
T. pubescensPRM 900586AY684173AY855906 [37]
Truncospora ochroleucaDai 11486HQ654105JF706349KF051048KF181142[5]
T. ochroleucaMUCL 39726FJ411098FJ393865 [28]
T. ochroleucaCui 5671JX941584JX941602KF218309KF286315[5]
T. ochroleucaCui 5673JX941585JX941603KF218308KF286314[5]
T. ornataCui 5714HQ654103HQ654116KF051056KF181150[5]
T. ohiensisMUCL 41036FJ411096FJ393863 [28]
Tyromyces chioneusCui 10225KF698745KF698756 [5]
T. kmetiiPenttila 13474KF705040KF705041 [5]
Vanderbylia delavayiDai 6891JQ861738KF495019KF218287KF286293[5]
V. fraxineaDP 83AM269789AM269853 [28]
V. fraxineaCui 7154HQ654095HQ654110KF218288KF286294[5]
V. fraxineaCui 8885HQ876611JF706344KF218289KF286295[5]
V. fraxineaCui 8871JF706329JF706345KF051050KF181144[5]
V. robiniophilaCui 5644HQ876609JF706342KF051051KF181145[5]
V. vicinaMUCL 44779FJ411095AF518666 [28]
Yuchengia narymicaDai 7050JN048776JN048795KF051053KF181147[39]
Y. narymicaDai 10510HQ654101JF706346KF051054KF181148[39]
Y. narymicaDai 6998JN048775JN048794KF051055KF181149[5]
Y. narymica0709/42JN641258JN641265 [39]
Y. narymica0709/157JN641259JN641266 [39]
Y. narymica0809/3JN641261JN641268KF051049KF181143[39]
Table 2. A comparison of Poriella subacida and related genera and species.
Table 2. A comparison of Poriella subacida and related genera and species.
GeneraHyphal SystemBasidiospore MorphologyChemical ReactionsReference
Skeletal HyphaeBasidiospores
Perenniporia s.s.dimiticellipsoid, truncate or notdextrinoid or not, CB+variable dextrinoid, CB+[13]
Perenniporiopsistrimiticoblong-ellipsoid, truncatedextrinoid, CB– in the context, CB+ in the tramadextrinoid, CB+[6]
Perenniporielladimiticglobose to subglobose, non-truncatenon-dextrinoid to strongly dextrinoid, slightly to distinctly cyanophilousslightly dextrinoid, CB+[51]
Poriella subacidadi-trimiticellipsoid, non-truncatestrongly dextrinoid, CB+non-dextrinoid, CB+This study
Yuchengia narymicadimiticellipsoid, non-truncateamyloid, CB–IKI–, CB+[38]
CB+ = cyanophilous, CB– = acyanophilous.
Table 3. A list of characteristics of Poriella subacida comb. nov. from different regions.
Table 3. A list of characteristics of Poriella subacida comb. nov. from different regions.
SpecimensLocalityBasidiospores (µm)AverageQPores/mmSubstrate
BPI 844697 (Type)USA, NY(4.2–)4.4–5.7(–6.2) × (3.1–)3.5–4.3(–4.5)5.2 × 3.91.354–5on Tsuga Carr.
BPI 844698USA, NY(4.7–)5–6(–6.4) × (3.6–)3.9–4.6(–4.8)5.4 × 4.11.364–5on Tsuga Carr.
BPI 844699USA, NY(4.3–)4.5–5.7(–6.2) × (3.2–)3.6–4.4(–4.6)5.2 × 3.91.354–5on Tsuga Carr.
BPI 885858USA, NY(4.9–)5.1–6.2(–6.5) × (3.4–)3.7–4.6(–4.8)5.6 × 4.21.34–5on Tsuga Carr.
FH0053717USA, NY(4.8–)5–5.6(–5.8) × (3.7–)3.9–4.5(–4.7)5.3 × 4.21.334–5on the log of Picea asperata Mast.
Hesler 11907USA, TN(4.4–)4.6–5.3(–5.5) × (3.5–)3.7–4.3(–4.6)4.9 × 41.24–5on the log of Pinus Linn
00605389USA, KS(4.1–)4.5–5.1(–5.3) × (3–)3.4–4.5(–4.8)4.8 × 3.81.34–6on underside of old log
Lowe 9407USA, AZ(4.6–)4.9–5.8(–6) × (3.9–)4.3–4.9(–5.1)5.4 × 4.51.24–6on the log of Picea asperata Mast.
00605344USA, ME(4.3–)4.6–5.2(–5.4) × (3.4–)3.6–4.2(–4.6)4.9 × 3.91.24–5on fallen Populus L. log
00605372Canada, Ontario(4.4–)4.6–5.8(–6) × (3–)3.3–4.2(–4.5)5.3 × 3.81.24–6on fallen trunk of Abies Mill
Cui 9849China, Heilongjiang(4.9–)5.2–5.7(–5.9) × (3.9–)4.1–4.4(–4.6)5.4 × 4.21.34–5on fallen trunk of Picea Dietr.
Yuan 3854China, Yunnan(5–)5.2–5.8(–6) × (3.9–)4.1–4.4(–4.5)5.6 × 4.21.34–6on angiosperm trunk
Cui 2712China, Zhejiang(4.9–)5.1–5.7(–6) × (3.9–)4–4.4(–4.6)5.4 × 4.21.34–5on fallen trunk of Picea Dietr.
Dai 2648Finland, Pohjanmaa(4.3–)4.7–5.7(–6.1) × (3–)3.3–3.9(–4.1)5.3 × 3.61.34–6on fallen trunk of Picea Dietr.
Dai 12619Finland, Pohjanmaa(4.5–)4.6–5.8(–6.4) × (3.1–)3.5–4.1(–4.6)5.2 × 3.81.34–5on Picea Dietr.
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Chen, R.; Karunarathna, S.C.; Zhao, C.-L. Poriella subacida Gen. & Comb Nov. for Perenniporia subacida (Peck) Donk. Agronomy 2021, 11, 1308. https://doi.org/10.3390/agronomy11071308

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Chen R, Karunarathna SC, Zhao C-L. Poriella subacida Gen. & Comb Nov. for Perenniporia subacida (Peck) Donk. Agronomy. 2021; 11(7):1308. https://doi.org/10.3390/agronomy11071308

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Chen, Rui, Samantha C. Karunarathna, and Chang-Lin Zhao. 2021. "Poriella subacida Gen. & Comb Nov. for Perenniporia subacida (Peck) Donk" Agronomy 11, no. 7: 1308. https://doi.org/10.3390/agronomy11071308

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