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Article

Phylogenetic and Taxonomic Analyses of Three New Wood-Inhabiting Fungi of Xylodon (Basidiomycota) in a Forest Ecological System

by
Kai-Yue Luo
1,2,3,
Zhuo-Yue Chen
3 and
Chang-Lin Zhao
1,2,3,4,5,*
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
Yunnan Key Laboratory of Plateau Wetland Conservation, Restoration and Ecological Services, Southwest Forestry University, Kunming 650224, China
3
College of Biodiversity Conservation, Southwest Forestry University, Kunming 650224, China
4
Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China
5
School of Life Sciences, Tsinghua University, Beijing 100084, China
*
Author to whom correspondence should be addressed.
J. Fungi 2022, 8(4), 405; https://doi.org/10.3390/jof8040405
Submission received: 19 February 2022 / Revised: 3 April 2022 / Accepted: 13 April 2022 / Published: 15 April 2022
(This article belongs to the Special Issue Polyphasic Identification of Fungi)
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Abstract

:
Wood-inhabiting fungi are a cosmopolitan group and show a rich diversity, growing in the vegetation of boreal, temperate, subtropical, and tropical regions. Xylodon grandineus, X. punctus, and X. wenshanensis spp. nov. were found in the Yunnan–Guizhou Plateau, China, suggested here to be new fungal species in light of their morphology and phylogeny. Xylodon grandineus is characterized by a grandinioid hymenophore and ellipsoid basidiospores; X. punctus has a membranous hymenophore, a smooth hymenial surface with a speckled distribution, and absent cystidia; X. wenshanensis has a grandinioid hymenophore with a cream to slightly buff hymenial surface and cystidia of two types. Sequences of the ITS and nLSU rRNA markers of the studied samples were generated, and phylogenetic analyses were performed using the maximum likelihood, maximum parsimony, and Bayesian inference methods. After a series of phylogenetic studies, the ITS+nLSU analysis of the order Hymenochaetales indicated that, at the generic level, six genera (i.e., Fasciodontia, Hastodontia, Hyphodontia, Lyomyces, Kneiffiella, and Xylodon) should be accepted to accommodate the members of Hyphodontia sensu lato. According to a further analysis of the ITS dataset, X. grandineus was retrieved as a sister to X. nesporii; X. punctus formed a monophyletic lineage and then grouped with X. filicinus, X. hastifer, X. hyphodontinus, and X. tropicus; and X. wenshanensis was a sister to X. xinpingensis.

1. Introduction

In forest ecosystems, fungi play essential ecological roles, driving carbon cycling in forest soils, mediating mineral nutrition of plants, and alleviating carbon limitations [1]. Wood-inhabiting fungi are a cosmopolitan group and have a rich diversity related to the high diversity of plants growing in boreal, temperate, subtropical, and tropical regions [2,3,4,5,6,7,8,9]. The order Hymenochaetales Oberw. comprises many representative wood-inhabiting fungal taxa, including hydnoid, corticioid, and polyporoid fungi possessing basidiomes with diverse hymenophoral and cystidial morphology [10,11,12,13,14]. Members of the family Schizoporaceae Jülich are widely found in different countries and areas. In addition, they cause white rot [15].
To accomplish the genome evolution and reconstruction of the phylogenetic relationships of fungi, an increasing number of taxa have been used for the fungal tree of life by using genome-scale data in the molecular systematics by mycologists [16], and both the species diversity and the classification of fungi are still in great, flux mainly in the more basal branches of the tree topology. The true diversity will come to light from genomic analyses and more region surveys worldwide based on some unique fungal groups.
The wood-inhabiting fungal genus Xylodon (Pers.) Gray (Schizoporaceae, Hymenochaetales) is typified by X. quercinus (Pers.) Gray [4]. This genus is characterized by the resupinate or effuse basidiomata with a smooth, tuberculate, grandinioid, odontioid, coralloid, irpicoid, or poroid hymenophore; a monomitic or dimitic hyphal system with clamped generative hyphae; the presence of different types of cystidia; utriform or suburniform basidia; and cylindrical to ellipsoid to globose basidiospores, in addition to causing white rot [4,17]. Based on the MycoBank database (http://www.mycobank.org, accessed on 20 March 2022) and the Index Fungorum (http://www.indexfungorum.org, accessed on 20 March 2022), the genus Xylodon has registered 218 specific and infraspecific names, but the actual number of the species has reached 92 [4,5,12,14,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43].
These pioneering studies of the genus Xylodon were just the prelude to the molecular systematics period [16]. Hyphodontia s.l. was shown to be a polyphyletic genus, in which Xylodon and Kneiffiella P. Karst are the most species rich [10,12,14]. Due to a lack of rDNA sequences for many taxa, the molecular data were not enough to separate many genera clearly; therefore, a broad concept of Hyphodontia s.l. was employed by mycologists [10,12,14,32,34]. Yurchenko et al. described two clades: the Xylodon-Lyomyces-Rogersella clade and the Xylodon-Schizopora-Palifer clade, and they suggested to mix the species of Xylodon, Schizopora Velen., Palifer Stalpers and P.K. Buchanan, Lyomyces P. Karst., and Rogersella Liberta and A.J. Navas within both clades. The research comprised the representative sequences and taxa of Hyphodontia s.l., such as Xylodon, Schizopora, Palifer, Lyomyces, and Rogersella, in which the result demonstrated that it was hard to distinguish the two genera Xylodon and Schizopora on the basis of the morphological and phylogenetic information; therefore, the authors proposed that Xylodon and Schizopora should be united into the genus Xylodon [12]. For the phylogenetic relationship of the Xylodon species, it was confirmed that the two genera Lagarobasidium Jülich and Xylodon should be synonymous based on molecular data from the ITS and nLSU regions, in which the three species X. pumilius (Gresl. and Rajchenb.) K.H. Larss., X. magnificus (Gresl. and Rajchenb.) K.H. Larss., and X. rickii (Gresl. and Rajchenb.) K.H. Larss. were combined into Xylodon [36]. All of the members of the genera Odontipsis Hjortstam and Ryvarden and Palifer were placed in the genus Xylodon based on the molecular analyses of 28S and ITS data, in which they proposed four new species of Xylodon as X. exilis Yurchenko, Riebesehl and Langer, X. filicinus Yurchenko and Riebesehl, X. follis Riebesehl, Yurchenko and Langer, and X. pseudolanatus Nakasone, Yurchenko and Riebesehl [14]. Based on the multiple loci in Hyphodontia s.l., Fasciodontia Yurchenko and Riebesehl, Hastodontia (Parmasto) Hjortstam and Ryvarden, Hyphodontia J. Erikss., Lyomyces, Kneiffiella, and Xylodon in the order Hymenochaetales, they were divided into four clades [41]. The phylogeny of the Xylodon species based on the ITS and nLSU sequences proposed three new taxa from China, in which X. gossypinus C.L. Zhao and K.Y. Luo and X. brevisetus (P. Karst.) Hjortstam and Ryvarden grouped together [40]. Based on the morphological descriptions and molecular analyses, three new species: Xylodon angustisporus Viner and Ryvarden, X. dissiliens Viner and Ryvarden, and X. laxiusculus Viner and Ryvarden, were described and placed in Xylodon, which were found in Africa [42]. A phylogenetic and taxonomic study on Xylodon (Hymenochaetales) described three new species of this genus from southern China, inferred from 61 fungal specimens representing 55 species, which enriched the fungal diversity of this areas [43].
During investigations on wood-inhabiting fungi in the Yunnan–Guizhou Plateau of China, three additional Xylodon species were collected. To clarify the placement and relationships of the three species, we study carried out a phylogenetic and taxonomic study on Xylodon, based on the ITS and nLSU sequences.

2. Materials and Methods

2.1. Sample Collection and Herbarium Specimen Preparation

The fresh fruiting bodies of the fungi growing on fallen angiosperm branches and fallen Pinus armandii branches were collected from Honghe, Wenshan, and Yuxi of Yunnan Province, China. The samples were photographed in situ, and fresh macroscopic details were recorded. Photographs were recorded by a Jianeng 80D camera. All of the photos were focus stacked and merged using Helicon Focus software. Macroscopic details were recorded and transported to a field station where the fruit body was dried on an electronic food dryer at 45 °C. Once dried, the specimens were sealed in an envelope and zip-lock plastic bags and labeled [43]. The dried specimens were deposited in the herbarium of the Southwest Forestry University (SWFC), Kunming, China.

2.2. Morphology

The macromorphological descriptions were based on field notes and photos captured in the field and lab. The color terminology follows that of Petersen [44]. The micromorphological data were obtained from the dried specimens after observation under a light microscope with a magnification of 10 × 100 oil [27]. The following abbreviations are used: KOH = 5% potassium hydroxide water solution, CB− = acyanophilous, IKI− = both inamyloid and indextrinoid, L = mean spore length (arithmetic average for all spores), W = mean spore width (arithmetic average for all spores), Q = variation in the L/W ratios between the specimens studied, and n = a/b (number of spores (a) measured from given number (b) of specimens).

2.3. Molecular Phylogeny

The CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to obtain genomic DNA from the dried specimens according to the manufacturer’s instructions. The nuclear ribosomal ITS region was amplified with ITS5 and ITS4 primers [45]. The nuclear nLSU region was amplified with the LR0R and LR7 primer pair (http://lutzonilab.org/nuclear-ribosomal-dna/, accessed on 22 January 2022). 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 sequenced at Kunming Tsingke Biological Technology Limited Company (Kunming, China). All of the newly generated sequences were deposited in NCBI GenBank (https://www.ncbi.nlm.nih.gov/genbank/, accessed on 22 January 2022) (Table 1).
The sequences were aligned in MAFFT version 7 [56] using the G-INS-i strategy. The alignment was adjusted manually using AliView version 1.27 [57]. The dataset was aligned first, and then the sequences of ITS and nLSU were combined with Mesquite version 3.51. The alignment datasets were deposited in TreeBASE (submission ID 29411). ITS+nLSU sequences and ITS-only datasets were used to infer the position of the three new species in the genus Xylodon and related species. Sequences of Hymenochaete cinnamomea (Pers.) Bres. and H. rubiginosa (Dicks.) Lév. retrieved from GenBank were used as an outgroup in the ITS+nLSU analysis (Figure 1); sequences of Lyomyces orientalis Riebesehl, Yurch. and Langer, and L. sambuca (Pers.) P. Karst. retrieved from GenBank were used as an outgroup in the ITS analysis (Figure 2) [41].
Maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI) analyses were applied to the combined three datasets following a previous study [58], and the tree construction procedure was performed in PAUP* version 4.0b10 [59]. All of the characters were equally weighted, and gaps were treated as missing data. Using the heuristic search option with TBR branch swapping and 1000 random sequence additions, trees were inferred. Max trees were set to 5000, branches of zero length were collapsed, and all parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1000 replicates [60]. Descriptive tree statistics, tree length (TL), the consistency index (CI), the retention index (RI), the rescaled consistency index (RC), and the homoplasy index (HI) were calculated for each maximum parsimonious tree generated. The multiple sequence alignment was also analyzed using maximum likelihood (ML) in RAxML-HPC2 [61]. Branch support (BS) for ML analysis was determined by 1000 bootstrap replicates.
MrModeltest 2.3 [62] was used to determine the best-fit evolution model for each dataset for Bayesian inference (BI), which was performed using MrBayes 3.2.7a with a GTR+I+G model of DNA substitution and a gamma distribution rate variation across sites [63]. A total of four Markov chains were run for two runs from random starting trees for 1.2 million generations for ITS+nLSU (Figure 1) and 9 million generations for ITS (Figure 2) with trees and parameters sampled every 1000 generations. The first one-fourth of all of the generations were discarded as burn-ins. The majority-rule consensus tree of all of the remaining trees was calculated. Branches were considered significantly supported if they received a maximum likelihood bootstrap value (BS) of >70%, a maximum parsimony bootstrap value (BT) of >70%, or Bayesian posterior probabilities (BPP) of >0.95.

3. Results

3.1. Molecular Phylogeny

The ITS+nLSU (Figure 1) included sequences from 50 fungal samples representing 50 species. The dataset had an aligned length of 1919 characters, of which 1140 characters were constant, 206 were variable and parsimony uninformative, and 573 were parsimony informative. The maximum parsimony analysis yielded six equally parsimonious trees (TL = 3564, CI = 0.3533, HI = 0.6467, RI = 0.5235, and RC = 0.1849). The best model for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was GTR+I+G. The Bayesian and ML analyses showed a similar topology to that of the MP analysis with split frequencies = 0.008056 (BI), and the effective sample size (ESS) average ESS (avg ESS) = 418.5. The phylogram based on the ITS+nLSU rDNA gene regions (Figure 1) includes four families within Hymenochaetales, which comprises six genera: Fasciodontia, Hastodontia, Hyphodontia, Kneiffiella, Lyomyces, and Xylodon, which indicated that three genera (Fasciodontia, Lyomyces, and Xylodon) fell into the family Schizoporaceae.
The ITS-only dataset (Figure 2) included sequences from 67 fungal specimens representing 61 taxa. The dataset had an aligned length of 582 characters, of which 233 characters were constant, 59 were variable and parsimony uninformative, and 290 were parsimony informative. Maximum parsimony analysis yielded 5000 equally parsimonious trees (TL = 2191, CI = 0.2743, HI = 0.7257, RI = 0.4560, and RC = 0.1251). The best model for the ITS dataset estimated and applied in the Bayesian analysis was GTR+I+G. The Bayesian and ML analyses resulted in a similar topology to that of the MP analysis with split frequencies = 0.022072 (BI), and the effective sample size (ESS) of the average ESS (avg ESS) = 1670.5. The phylogram inferred from the ITS sequences analysis (Figure 2) indicated that three new species grouped into genus Xylodon: the new species X. grandineus was a sister to X. nesporii (Bres.) Hjortstam and Ryvarden; X. punctus formed a monophyletic lineage and then grouped with X. filicinus Yurchenko and Riebesehl, X. hastifer (Hjortstam and Ryvarden) Hjortstam and Ryvarden, X. hyphodontinus (Hjortstam and Ryvarden) Riebesehl, Yurchenko and G. Gruhn, and X. tropicus, while X. wenshanensis was retrieved as a sister of X. xinpingensis C.L. Zhao and X. Ma.

3.2. Taxonomy

Xylodon grandineus K.Y. Luo and C.L. Zhao, sp. nov. Figure 3 and Figure 4.
MycoBank no.: 843054.
Holotype—China, Yunnan Province, Yuxi, Xinping County, Mopanshan National Forestry Park. GPS coordinates: 24°53′ N, 101°57′ E; altitude: 2000 m asl. On fallen Pinus armandii branches, leg. C.L. Zhao, 19 January 2018, CLZhao 6425 (SWFC).
Etymology—grandineus (Lat.): Referring to the hymenial surface grandinioid of the specimens.
Basidiomata—Annual, resupinate, adnate, soft coriaceous when fresh, coriaceous upon drying, up to 10 cm long, 1.5 cm wide, 100–200 µm thick. Hymenial surface grandinioid, without odor or taste when fresh, pale buff when fresh, pale buff to buff when dry. Sterile margin indistinct, cream to buff, 0.5–1 mm wide.
Hyphal system—Monomitic, generative hyphae with clamp connections, colorless, thick-walled, frequently branched, interwoven, 2–4 µm in diameter, IKI−, CB−; tissues unchanged in KOH; subhymenial hyphae densely covered by the crystals.
Hymenium—Cystidia subulate, colorless, thin-walled, smooth, 11–19 × 3–5 µm; basidia barreled, constricted, with four sterigmata and a basal clamp connection, 11–19 × 2.5–4 µm.
Spores—Basidiospores ellipsoid, colorless, thin-walled, smooth, with one oil drop inside, IKI−, CB−, 3–4.5(–5) × 2–3 µm, L = 3.69 µm, W = 2.46 µm, Q = 1.46–1.54 (n = 60/2).
Additional specimenexamined (paratype)—China, Yunnan Province, Wenshan, Pingba Town, Wenshan National Nature Reserve. GPS coordinates: 23°15′ N, 104°06′ E; altitude: 1600 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 25 July 2019, CLZhao 16075 (SWFC).
Xylodon punctus K.Y. Luo and C.L. Zhao, sp. nov. Figure 5 and Figure 6.
MycoBank no.: 843055.
Holotype—China, Yunnan Province, Honghe, Pingbian County, Daweishan National Nature Reserve. GPS coordinates: 23°42′ N, 103°30′ E; altitude: 1500 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 1 August 2019, CLZhao 17691 (SWFC).
Etymology—punctus (Lat.): Referring to the spotted hymenial surface of the specimens.
Basidiomata—Annual, resupinate, adnate, thin, membranous, very hard to separate from substrate, up to 12 cm long, 1.5 cm wide, 20–80 µm thick. Hymenial surface smooth, speckled distribution, white when fresh, white to slightly grey upon drying. Sterile margin indistinct, white, up to 1 mm wide.
Hyphal system—Monomitic, generative hyphae with clamp connections, colorless, thin- to thick-walled, occasionally branched, interwoven, 1–3 µm in diameter, IKI−, CB−; tissues unchanged in KOH.
Hymenium—Cystidia absent; basidia clavate, short and obtused, with four sterigmata and a basal clamp connection, 10–16 × 3.5–5.5 µm.
Spores—Basidiospores ellipsoid to broad ellipsoid, colorless, thin-walled, smooth, IKI−, CB−, (1.5–)2–4(–4.5) × 1.5–2.5(–3) µm, L = 2.71 µm, W = 1.98 µm, Q = 1.32–1.43 (n = 31/3).
Additional specimens examined (paratypes)—China, Yunnan Province, Honghe, Pingbian County, Daweishan National Nature Reserve. GPS coordinates: 23°42′ N, 103°30′ E; altitude: 1500 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 1 August 2019, CLZhao 17908; CLZhao 17916 (SWFC).
Xylodon wenshanensis K.Y. Luo and C.L. Zhao, sp. nov. Figure 7 and Figure 8.
MycoBank no.: 843056.
Holotype—China, Yunnan Province, Wenshan, Xichou County, Jiguanshan Forestry Park. GPS coordinates: 23°15′ N, 104°40′ E; altitude: 1500 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 22 July 2019, CLZhao 15729 (SWFC).
Etymology—wenshanensis (Lat.): Referring to the specimens’ provenance from the Wenshan locality.
Basidiomata—Annual, resupinate, thin, without odor and taste when fresh, coriaceous, up to 9 cm long, 2 cm wide, 50–100 µm thick. Hymenial surface grandinioid, cream when fresh, cream to slightly buff upon drying. Sterile margin indistinct, cream, about 1 mm wide.
Hyphal system—Monomitic, generative hyphae with clamp connections, colorless, thin- to thick-walled, frequently branched, interwoven, 2–3.5 µm in diameter; IKI−, CB−; tissues unchanged in KOH.
Hymenium—Cystidia of two types: (1) capitate cystidia in hymenium and subiculum, colorless, thin-walled, smooth, slightly constricted at the neck, with a globose head, 6–11 × 3–6.5 µm; (2) clavate cystidia, slightly sinuous, 10.5–20 × 2.5–5.5 µm; basidia clavate to subcylindrical, slightly sinuous, with four sterigmata and a basal clamped connection, 8–15.5 × 3–5 µm.
Spores—Basidiospores ellipsoid, colorless, thin-walled, smooth, IKI−, CB−, 3–5 × 2–3.5(–4) µm, L = 3.96 µm, W = 2.82 µm, Q = 1.35–1.47 (n = 120/4).
Additionalspecimens examined (paratypes)—China, Yunnan Province, Wenshan, Xichou County, Xiaoqiaogou, Wenshan National Nature Reserve. GPS coordinates: 23°22′ N, 104°43′ E; altitude: 1500 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 14 January 2019, CLZhao 10790 (SWFC); Jiguanshan Forestry Park. GPS coordinates: 23°15′ N, 104°40′ E; altitude: 1500 m asl. On fallen angiosperm branches, leg. C.L. Zhao, 22 July 2019, CLZhao 15718, CLZhao 15782 (SWFC).

4. Discussion

Many recently described wood-inhabiting fungi taxa have been reported in the subtropics and tropics, including those of the genus Xylodon [64,65,66,67,68,69,70], which were collected on rotten trunks and stumps of conifers and angiosperms, bamboo, and ferns [2,3,6,15,23,25,40,41,43,52,71,72,73,74,75,76,77,78,79,80,81,82,83]. The present study reports three new taxa of Xylodon: X. grandineus, X. punctus, and X. wenshanensis, based on a combination of morphological features and molecular evidence.
Phylogenetically, the molecular relationships of Xylodon and related genera located in Hyphodontia s.l. (Hymenochaetales), on the basis of the combined datasets of ITS, nLSU, and mt-SSU regions, indicated that seven families—Chaetoporellaceae Jülich, Coltriciaceae Jülich, Hymenochaetaceae Donk, Neoantrodiellaceae Y.C. Dai, B.K. Cui, Jia J. Chen and H.S. Yuan, Nigrofomitaceae Jülich, Oxyporaceae Zmitr. and Malysheva, and Schizoporaceae—were monophyletic lineages, which nested in the order Hymenochaetales, in which some genera grouped into Hyphodontia s.l. as independent genera, including Xylodon [41]. In the present study (Figure 1), four families in the order Hymenochaetales were analyzed by the ITS+nLSU data, which showed that the genus Xylodon nested into the family Schizoporaceae.
The ITS-based evolution phylogram for Xylodon and related species revealed four species—X. cystidiatus (A. David and Rajchenb.) Riebesehl and Langer; X. hyphodontinus; X. serpentiformis (Langer) Hjortstam and Ryvarden; and X. subclavatus (Yurchenko, H.X. Xiong and Sheng H. Wu) Riebesehl, Yurch. and Langer—were in the genus Xylodon [14]. In the current study (Figure 2), the three new species also nested into the genus Xylodon, in which X. grandineus was a sister to X. nesporii; X. punctus formed a monophyletic lineage and then grouped with X. filicinus, X. hastifer, X. hyphodontinus, and X. tropicus, while X. wenshanensis was retrieved as a sister to X. xinpingensis. However, morphologically, Xylodon nesporii can be delimited from X. grandineus has an odontioid hymenial surface and narrowly ellipsoid to cylindrical basidiospores (4.5–6 × 2–2.5 µm) [70]. Xylodon filicinus differs from X. punctus by its odontioid hymenial surface and larger, globose to subglobose basidiospores (4–5 × 4–4.5 µm) [14]. Xylodon hastifer could be delimited from X. punctus by its odontioid hymenial surface and larger, subglobose basidiospores (4.5–5 × 4–4.5 µm) [15]. Xylodon hyphodontinus is differs from X. punctus in its odontioid hymenial surface and globose to subglobose basidiospores (4.5–5 μm in diameter) [64]. Xylodon tropicus differs from X. punctus in its coriaceous basidiomata with a grandinioid hymenial surface and subglobose basidiospores [43]. Xylodon xinpingensis can be delimited from X. wenshanensis by its soft-membranaceous basidiomata, a reticulate hymenial surface, fusiform cystidia (19.5–31 × 2–6 µm), and larger subglobose basidiospores (5–6.4 × 3.5–5 µm) [52].
Morphologically, Xylodon grandineus is similar to X. follis Riebesehl, Yurchenko and Langer; X. laceratus C.L. Zhao; X. macrosporus C.L. Zhao and K.Y. Luo; X. tropicus; and X. sinensis C.L. Zhao and K.Y. Luo due to its the grandinioid hymenial surface. However, Xylodon follis differs from X. grandineus in its effused basidiomata with a cream-colored hymenial surface, capitate cystidia (17–30 × 4.5–9 µm), and larger globose to subglobose basidiospores (8–9.5 × 7–8.5 µm) [14]. Xylodon laceratus differs from X. grandineus in its capitate cystidia (15.4–24.7 × 3.8–4.7 µm) and fusiform cystidia (20.3–26.8 × 5.3–6.4 µm) [43]. Xylodon macrosporus differs from X. grandineus by having cystidia of three types: capitate cystidia (8–25.5 × 3–10 µm), cylindrical cystidia (44–79.5 × 3–6 µm), and cystidia (11–21 × 6–11 µm), as well as larger thick-walled basidiospores (8–10.5 × 7.5–9 µm) [40]. Xylodon tropicus can be delimited from X. grandineus by its buff to pale brown hymenial surface; absent cystidia; and subglobose, slightly thick-walled basidiospores [43]. Xylodon sinensis is distinguishable from X. grandineus by its buff to brown hymenial surface, fusiform cystidia, and subglobose basidiospores [40].
Xylodon grandineus resembles X. attenuatus Spirin and Viner; X. borealis (Kotir. and Saaren.) Hjortstam and Ryvarden; X. bresinskyi (Langer) Hjortstam and Ryvarden; X. dimiticus (Jia J. Chen and L.W. Zhou) Riebesehl and E. Langer; and X. vesiculosus Yurchenko, Nakasone and Riebesehl with its ellipsoid basidiospores. However, Xylodon attenuatus differs from X. grandineus in its cream-colored, grandinioid to odontoid hymenial surface with rather regularly arranged projections; cystidia of two types: subcapitate or capitate cystidia (13.5–25.1 × 3.5–5 µm) and hyphoid cystidia (16–38.3 × 2.8–4.5 µm); and wider basidiospores (4.1–5.5 × 3.4–4.5 µm) [36]. Xylodon borealis differs from X. grandineus by having effused basidiomata; cystidia of two types: capitate cystidia (20–50 × 4–6 µm) and slender hypha-like cystidia (40–70 × 3–5 µm); and larger basidiospores (4.5–5.5 × 3.5–4 µm) [4]. Xylodon bresinskyi differs from X. grandineus in its poroid hymenial surface with rudimentary console shaping and larger basidiospores (4.5–5.5 × 3–3.5 µm) [84]. Xylodon dimiticus is distinguishable from X. grandineus by its poroid hymenial surface with angular pores (2–4 per mm) and absent cystidia [28]. Xylodon vesiculosus can be delimited from X. grandineus by its membranaceous basidiomata with an odontioid hymenial surface and larger basidiospores (5.3–6.3 × 3–4 µm) [14].
Xylodon punctus is similar to X. acystidiatus Xue W. Wang and L.W. Zhou, X. gossypinus C.L. Zhao and K.Y. Luo, X. montanus C.L. Zhao, and X. nudisetus (Warcup and P.H.B. Talbot) Hjortstam and Ryvarden in having a smooth hymenial surface. However, Xylodon acystidiatus differs from X. punctus by having brittle basidiomata with a cracked hymenial surface and larger basidiospores (4.7–5.3 × 2.7–3.7 µm) [41]. Xylodon gossypinus differs from X. punctus in its cotton hymenial surface and wider basidiospores (3–5.5 × 2.5–4 μm) [40]. Xylodon montanus can be delimited from X. punctus by its absence of a speckled distribution on the hymenial surface and wider basidiospores (3.9–5.3 × 3.2–4.3 μm) [43]. Xylodon nudisetus differs from X. punctus in having larger basidiospores (4.5–6 × 3–4.5 µm) [4].
Xylodon punctus resembles X. bambusinus C.L. Zhao and X. Ma; X. mussooriensis Samita, Sanyal and Dhingra ex L.W. Zhou and T.W. May; X. rhododendricola Xue W. Wang and L.W. Zhou; X. pruinosus (Bres.) Spirin and Viner; and X. ussuriensis Viner in having ellipsoid to broad ellipsoid basidiospores. However, Xylodon bambusinus is distinguished from X. punctus by its ceraceous basidiomata with a grandinoid hymenial surface and larger basidiospores (4–5 × 2.6–3.7 µm) [52]. Xylodon mussooriensis differs from X. punctus by the presence of an odontioid hymenial surface and larger basidiospores (5.2–5.8 × 3.1–3.5 µm) [41]. Xylodon rhododendricola differs from X. punctus in having an odontioid hymenial surface and larger basidiospores (4.8–6.5 × 3.8–5.1 µm) [41]. Xylodon pruinosus differs from X. punctus in having a grandinioid to odontoid hymenial surface with greyish-white or pale cream-colored and larger, clearly thick-walled basidiospores (4.5–5.9 × 3.7–4.8 µm) [36]. Xylodon ussuriensis is distinguished from X. punctus by its grandinioid to odontoid hymenial surface with larger, pale ochraceous, and clearly thick-walled basidiospores (5.1–6 × 3.8–4.6 µm) [36].
Xylodon wenshanensis is similar to X. laceratus, X. macrosporus, X. sinensis, X. victoriensis Xue W. Wang and L.W. Zhou, and X. yarraensis Xue W. Wang and L.W. Zhou in having a grandinioid hymenial surface. However, Xylodon laceratus is distinguished from X. wenshanensis by its capitate cystidia (15.4–24.7 × 3.8–4.7 µm) and fusiform cystidia (20.3–26.8 × 5.3–6.4 µm) [43]. Xylodon macrosporus is differentiated from X. wenshanensis in having three types cystidia and larger, thick-walled basidiospores (8–10.5 × 7.5–9 µm) [40]. Xylodon sinensis differs from X. wenshanensis in its fusiform cystidia (10–21 × 3–6 µm) and subglobose basidiospores (3–5 × 2.5–4 µm) [40]. Xylodon victoriensis can be delimited from X. wenshanensis by its brittle basidiomata with a cracked hymenophore, leptocystidia (30–40 × 4.5–5 μm), and globose to subglobose basidiospores (3.8–4.6 × 3.2–3.7 μm) [41]. Xylodon yarraensis is different from X. wenshanensis in its cracked and brittle basidiomata and capitate cystidia (25–30 × 2.5–3.5 µm) [41].
Xylodon wenshanensis resembles X. asper (Fr.) Hjortstam and Ryvarden; X. flaviporus (Berk. and M.A. Curtis ex Cooke) Riebesehl and Langer; X. ovisporus (Corner) Riebesehl and Langer; X. pseudolanatus Nakasone, Yurchenko and Riebesehl; and X. rimosissimus (Peck) Hjortstam and Ryvarden in having capitate cystidia. However, Xylodon asper is different from X. wenshanensis in having an odontioid hymenial surface with scattered aculei and larger basidiospores (5–6 × 3.5–4 μm) [4]. Xylodon flaviporus is distinguished from X. wenshanensis by its poroid hymenial surface with deep pores (up to 2 mm) and a pseudodimitic hyphal system [15]. Xylodon ovisporus is differentiated from X. wenshanensis by having a poroid hymenophore with pinkish-cream or buff hymenial surface [19]. Xylodon pseudolanatus differs from X. wenshanensis by its emebranaceous basidiomata, odontioid hymenial surface, and longer basidiospores (5–6 × 3–3.5 µm) [14]. Xylodon rimosissimus can be delimited from X. wenshanensis by its subceraceous basidiomata with a dense odontioid hymenial surface and larger basidiospores (5–6 × 3.5–4 μm) [4].
The macromorphology of the basidiomata and hymenophore construction do not reflect monophyletic groups based on a higher-level phylogenetic classification of polypores [82]. The current phylogeny (Figure 2) shows that the morphological characteristics do not follow the phylogenetic grouping of different taxa in Xylodon based on the ITS datasets.
Wood-inhabiting fungi are a characteristic group of Basidiomycota, which has a number of corticioid, poroid, and hydnoid genera based on the results of morphological, phylogenetic, and cytological studies in China [8,9]. To date, thirty-six species of Xylodon have been recorded in China [12,14,33,36,40,41,43,46,52,83], but the species diversity of Xylodon is still not well known in China, especially in the country’s subtropical and tropical areas. This paper enriches our knowledge of fungal diversity in this area, and it is likely that more new taxa will be found with further fieldwork and molecular analyses.

Author Contributions

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

Funding

The research was supported by the National Natural Science Foundation of China (Project No. 32170004, U2102220) to Chang-Lin Zhao, Yunnan Fundamental Research Project (Grant No. 202001AS070043) to Chang-Lin Zhao, the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111) to Chang-Lin Zhao, and Yunnan Key Laboratory of Plateau Wetland Conservation, Restoration and Ecological Services (202105AG070002) to Kai-Yue Luo.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

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 29411; accessed on 17 February 2022.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Maximum parsimony strict consensus tree illustrating the phylogeny of Xylodon and related genera in the order Hymenochaetales based on ITS+nLSU sequences. The families and genera represented by each color are indicated in the upper left of the phylogenetic tree.
Figure 1. Maximum parsimony strict consensus tree illustrating the phylogeny of Xylodon and related genera in the order Hymenochaetales based on ITS+nLSU sequences. The families and genera represented by each color are indicated in the upper left of the phylogenetic tree.
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Figure 2. Maximum parsimony strict consensus tree illustrating the phylogeny of three new species in Xylodon based on ITS sequences. Branches are labeled with a maximum likelihood bootstrap value > 70%, a parsimony bootstrap value > 50%, and Bayesian posterior probabilities > 0.95, respectively. The new species are in bold. The red stars representative holotypes.
Figure 2. Maximum parsimony strict consensus tree illustrating the phylogeny of three new species in Xylodon based on ITS sequences. Branches are labeled with a maximum likelihood bootstrap value > 70%, a parsimony bootstrap value > 50%, and Bayesian posterior probabilities > 0.95, respectively. The new species are in bold. The red stars representative holotypes.
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Figure 3. Basidiomata of Xylodon grandineus (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 3. Basidiomata of Xylodon grandineus (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
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Figure 4. Microscopic structures of Xylodon grandineus (holotype): basidiospores (A), basidia and basidioles (B), subulate cystidia (C), a section of the hymenium (D). Bars: (A) = 5 μm, (B–D) = 10 µm.
Figure 4. Microscopic structures of Xylodon grandineus (holotype): basidiospores (A), basidia and basidioles (B), subulate cystidia (C), a section of the hymenium (D). Bars: (A) = 5 μm, (B–D) = 10 µm.
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Figure 5. Basidiomata of Xylodon punctus (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 5. Basidiomata of Xylodon punctus (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
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Figure 6. Microscopic structures of Xylodon punctus (holotype): basidiospores (A), basidia and basidioles (B), a section of the hymenium (C). Bars: (A) = 5 μm, (B,C) = 10 µm.
Figure 6. Microscopic structures of Xylodon punctus (holotype): basidiospores (A), basidia and basidioles (B), a section of the hymenium (C). Bars: (A) = 5 μm, (B,C) = 10 µm.
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Figure 7. Basidiomata of Xylodon wenshanensis (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 7. Basidiomata of Xylodon wenshanensis (holotype): the front of the basidiomata (A), characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
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Figure 8. Microscopic structures of Xylodon wenshanensis (holotype): basidiospores (A), basidia and basidioles (B), capitate cystidia in subiculum and hymenium (C), clavate cystidia (D), a section of the hymenium (E). Bars: (A) = 5 μm, (B–E) = 10 µm.
Figure 8. Microscopic structures of Xylodon wenshanensis (holotype): basidiospores (A), basidia and basidioles (B), capitate cystidia in subiculum and hymenium (C), clavate cystidia (D), a section of the hymenium (E). Bars: (A) = 5 μm, (B–E) = 10 µm.
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Table
Table 1. List of species, specimens, and GenBank accession numbers of sequences used in this study.
Table 1. List of species, specimens, and GenBank accession numbers of sequences used in this study.
Species NameSpecimen No.GenBank Accession No.ReferencesCountry
ITSnLSU
Fasciodontia brasiliensisMSKF 7245aMK575201MK598734[46]Brazil
F. bugellensisKASFD 10705aMK575203MK598735[46]France
F. yunnanensisCLZhao 6280MK811275MZ146327[47]China
Hastodontia halonataHHB 17058MK575207MK598738[46]Mexico
Hymenochaete cinnamomeaHe 2074KU975460KU975500UnpublishedChina
Hym. rubiginosaHe 1049JQ716407JQ279667[48]China
Hyphodontia alutariaGEL 3183DQ340318DQ340373UnpublishedGermany
Hyp. argutaKHL 11938EU118632EU118633[49]Sweden
Hyp. densisporaLWZ 20170908-5MT319426MT319160[41]China
Hyp. pallidulaKASGEL 2097DQ340317DQ340372UnpublishedGermany
Hyp. zhixiangiiLWZ 20180903-5MT319423MT319158[41]China
Kneiffiella barba-jovisKHL 11730DQ873609DQ873610[14]Sweden
K. eucalypticolaLWZ 20180515-9MT319411MT319143[41]Australia
K. palmaeKASGEL 3456DQ340333DQ340369[46]China
K. subalutaceaGEL 2196DQ340341DQ340362[46]Norway
Lyomyces allantosporusFR 0249548KY800397KY795963[13]Réunion
L. bambusinusCLZhao 4831MN945968MW264919[50]China
L. fimbriatusWu 911204-4MK575210MK598740[46]China
L. mascarensisKASGEL 4833KY800399KY795964[46]Réunion
L. orientalisLWZ 20170909-7MT319436MT319170[41]China
L. sambuciKASJR 7KY800402KY795966[13]Germany
Xylodon acystidiatusLWZ 20180514-9MT319474MT319211[41]Australia
X. apacheriensisWu 0910-58KX857797 [11]China
X. asperKHL 8530AY463427AY586675[51]Sweden
X. astrocystidiatusWu 9211-71JN129972 [26]China
X. attenuatusSpirin 8775MH324476 [36]America
X. australisLWZ 20180509-8MT319503MT319248[41]China
X. bambusinusCLZhao 9174MW394657 [52]China
X. borealisJS 26064AY463429 [51]Norway
X. brevisetusJS 17863AY463428AY586676[51]Norway
X. crystalligerLWZ 20170816-33MT319521 [41]China
X. cystidiatusFR 0249200MH880195MH884896[14]Réunion
X. damansaraensisLWZ 20180417-23MT319499MT319244[41]Malaysia
X. detriticusZíbarová 30.10.17MH320793MH651372[36]Czech Republic
X. filicinusMSKF 12869MH880199NG067836[14]China
X. flaviporusFR 0249797MH880201 [14]Réunion
X. follisFR 0249814MH880204MH884902[14]Réunion
X. grandineusCLZhao 6425 *OM338090OM338099Present studyChina
X. grandineusCLZhao 16075OM338091OM338100Present studyChina
X. gossypinusCLZhao 8375MZ663804MZ663813[40]China
X. hastiferK(M) 172400NR166558 [12]USA
X. heterocystidiatusWei 17-314MT731753MT731754UnpublishedChina
X. hyphodontinusKASGEL 9222MH880205MH884903[14]Kenya
X. kunmingensisTUBFO 42565MH880198 [14]China
X.laceratusCLZhao 9892OL619258 [43]China
X. lagenicystidiatusLWZ 20180513-16MT319634 [41]Australia
X. lenisWu 890714-3KY081802 [12]China
X. macrosporusCLZhao 10226MZ663809MZ663817[40]China
X. mollissimusLWZ 20160318-3KY007517MT319347[41]China
X.montanusCLZhao 8179OL619260 [43]China
X. nesporiiLWZ 20180921-35MT319655MT319238[41]China
X. niemelaeiLWZ 20150707-13MT319630 [41]China
X. nongravisGC 1412-22KX857801KX857818[11]China
X. nothofagiICMP 13842AF145583 [53]China
X. ovisporusLWZ 20170815-31MT319666 [41]China
X. papillosusCBS 114.71MH860026 [54]The Netherlands
X. paradoxusDai 14983MT319519 [41]China
X. pruinosusSpirin 2877MH332700 [36]Estonia
X. pseudolanatusFP 150922MH880220 [14]Belize
X. pseudotropicusDai 16167MT319509MT319255[41]China
X. punctusCLZhao 17691 *OM338092OM338101Present studyChina
X. punctusCLZhao 17908OM338093 Present studyChina
X. punctusCLZhao 17916OM338094OM338102Present studyChina
X. quercinusKHL 11076KT361633 [51]Sweden
X. ramicidaSpirin 7664NR138013 UnpublishedUSA
X. rhododendricolaLWZ 20180513-9MT319621 [41]Australia
X. rimosissimusRyberg 021031DQ873627 [55]Sweden
X. serpentiformisLWZ 20170816-15MT319673 [41]China
X. sinensisCLZhao 11120MZ663811 [40]China
X. spathulatusLWZ 20180804-10MT319646 [41]China
X. subclavatusTUBFO 42167MH880232 [14]China
X. subflaviporusWu 0809-76KX857803 [11]China
X. subserpentiformisLWZ 20180512-16MT319486 [41]Australia
X. subtropicusLWZ 20180510-24MT319541 [41]China
X. taiwanianusCBS 125875MH864080 [54]The Netherlands
X. tropicusCLZhao 3351OL619261OL619269[43]China
X. ussuriensisKUN 1989NR166241 UnpublishedUSA
X. verecundusKHL 12261DQ873642 [55]Sweden
X. victoriensisLWZ 20180510-29MT319487 [41]Australia
X. wenshanensisCLZhao 10790OM338095OM338103Present studyChina
X. wenshanensisCLZhao 15718OM338096 Present studyChina
X. wenshanensisCLZhao 15729 *OM338097OM338104Present studyChina
X. wenshanensisCLZhao 15782OM338098OM338105Present studyChina
X. xinpingensisCLZhao 11224MW394662MW394654[52]China
X. yarraensisLWZ 20180510-5MT319639 [41]Australia
X. yunnanensisLWZ 20180922-47MT319660MT319253[41]China
* is shown holotype.
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Luo, K.-Y.; Chen, Z.-Y.; Zhao, C.-L. Phylogenetic and Taxonomic Analyses of Three New Wood-Inhabiting Fungi of Xylodon (Basidiomycota) in a Forest Ecological System. J. Fungi 2022, 8, 405. https://doi.org/10.3390/jof8040405

AMA Style

Luo K-Y, Chen Z-Y, Zhao C-L. Phylogenetic and Taxonomic Analyses of Three New Wood-Inhabiting Fungi of Xylodon (Basidiomycota) in a Forest Ecological System. Journal of Fungi. 2022; 8(4):405. https://doi.org/10.3390/jof8040405

Chicago/Turabian Style

Luo, Kai-Yue, Zhuo-Yue Chen, and Chang-Lin Zhao. 2022. "Phylogenetic and Taxonomic Analyses of Three New Wood-Inhabiting Fungi of Xylodon (Basidiomycota) in a Forest Ecological System" Journal of Fungi 8, no. 4: 405. https://doi.org/10.3390/jof8040405

APA Style

Luo, K. -Y., Chen, Z. -Y., & Zhao, C. -L. (2022). Phylogenetic and Taxonomic Analyses of Three New Wood-Inhabiting Fungi of Xylodon (Basidiomycota) in a Forest Ecological System. Journal of Fungi, 8(4), 405. https://doi.org/10.3390/jof8040405

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