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. 2023 May 18;13:1189600. doi: 10.3389/fcimb.2023.1189600

Two new species of Scytinostroma (Russulales, Basidiomycota) in Southwest China

Qiu-Yue Zhang 1, Hong-Gao Liu 2, Lu-Sen Bian 1,3,*, Qian Chen 4,5,*
PMCID: PMC10240063  PMID: 37284500

Abstract

Two new species of Scytinostroma viz. S. acystidiatum and S. macrospermum, are described from southwest China. Phylogeny based on ITS + nLSU dataset demonstrates that samples of the two species form two independent lineages and are different in morphology from the existing species of Scytinostroma. Scytinostroma acystidiatum is characterized by resupinate, coriaceous basidiomata with cream to pale yellow hymenophore, a dimitic hyphal structure with generative hyphae bearing simple septa, the absence of cystidia, and amyloid, broadly ellipsoid basidiospores measuring 4.7–7 × 3.5–4.7 μm. Scytinostroma macrospermum is characterized by resupinate, coriaceous basidiomata with cream to straw yellow hymenophore, a dimitic hyphal structure with generative hyphae bearing simple septa, numerous cystidia embedded or projecting from hymenium, and inamyloid, ellipsoid basidiospores measuring 9–11 × 4.5–5.5 μm. The differences between the new species and morphologically similar and phylogenetically related species are discussed.

Keywords: new taxa, Peniophoraceae, phylogeny, taxonomy, wood-rotting fungi

1. Introduction

The genus Scytinostroma Donk (Russulales, Basidiomycota), typified by S. portentosum (Berk. & M.A. Curtis) Donk, was established by Donk (1956). It is traditionally characterized by resupinate, coriaceous basidiomata, smooth to tuberculate hymenophore and a dimitic hyphal structure with simple septa or clamps on generative hyphae, filiform and dichotomously branched skeletal hyphae which are dextrinoid and cyanophilous, and subglobose to ellipsoid, thin-walled, variably amyloid basidiospores, and a white-rotting ecology (Donk, 1956; Rattan, 1974; Bernicchia and Gorjón, 2010; Wang et al., 2020; Tabish and Daniel, 2021).

The genus accommodated seven species derived from Corticium Fr. (without gloeocystidia) and Gloeocystidium P. Karst. (with gloeocystidia) when it was established. Later, Scytinostroma was gradually recognized by taxonomists, and the number of new species and new combinations has been increasing continuously (Donk, 1956; Gilbertson, 1962; Boidin, 1967; Rattan, 1974; Boidin and Lanquetin, 1977; Lanquetin, 1984; Boidin and Lanquetin, 1987; Boidin and Gilles, 1988; Hjortstam, 1990; Stalpers, 1996). So far, 36 species have been accepted in Scytinostroma worldwide (Nakasone, 2008; Liu, 2019; Wang et al., 2020). Recently, molecular phylogenetic studies demonstrated that Scytinostroma nested in Peniophoraceae within Russulales; furthermore, Scytinostroma was polyphyletic and formed four stable clades, as well as related to Gloiothele Bres., Vararia P. Karst., and Dichostereum Pilát (Nakasone and Micales, 1988; Larsson and Larsson, 2003; Miller et al., 2006; Larsson, 2007). Morphologically, Scytinostroma species are separated from other corticioid fungi of Russulales mainly by their tough and leathery texture of the basidiomata, as well as dextrinoid and dichotomously branched skeletal hyphae (Rattan, 1974; Liu, 2019).

During investigations on the diversity of wood-rotting fungi from China, two unknown corticioid specimens were collected from southwest China, and their morphology corresponded to the concepts of Scytinostroma. To confirm their affinity, phylogenetic analyses based on the ITS+ nLSU rDNA sequences were carried out. The two newly sequenced samples from Guizhou and Chongqing formed two well-supported lineages clustered with two sequences from Korea (KJ668461, Jang et al., 2016) and Japan (LC327052, Ogura-Tsujita et al., 2018), respectively. Based on morphological and phylogenetic evidences, we hereby propose two new species of Scytinostroma.

2. Materials and methods

2.1. Morphological studies

The studied specimens are deposited in the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Macro-morphological descriptions are based on field notes and dried specimens. Color terms followed Petersen (1996). Microscopic structures and abbreviations used in this study followed Wu et al. (2020) and Liu et al. (2022).

2.2. DNA extraction and sequencing

A CTAB rapid plant genome extraction kit (Aidlab Biotechnologies, Co., Ltd., Beijing, China) was used to obtain DNA products from voucher specimens, according to the manufacturer’s instructions with some modifications (Yuan et al., 2021; Yuan et al., 2022). The following primer pairs were used to amplify the DNA: ITS5 (5′-GGA AGT AAA AGT CGT AAC AAG G-3′) and ITS4 (5′-TCC TCC GCT TAT TGATAT GC-3′) for the internal transcribed spacer (ITS) regions (White et al., 1990); LR0R (5′-ACC CGC TGA ACT 6 TAA GC-3′) and LR7 (5′-TAC TAC CAC CAA GAT CT-3′) for nuclear large subunit (nLSU) rDNA (Vilgalys and Hester, 1990).

The procedures for DNA extraction and polymerase chain reaction (PCR) used in this study were the same as described by Wu et al. (2022b). The PCR products were purified and sequenced by Beijing Genomics Institute (BGI), China. All newly generated sequences in this study were deposited in GenBank (http://www.ncbi.nlm.nih.gov/genbank/) and listed in Table 1 .

Table 1.

Taxa information and GenBank accession numbers of sequences used in this study.

Species Specimen no. Locality ITS nLSU Literature
Confertobasidium olivaceoalbum FP 90196 USA AF511648 AF511648 Larsson and Larsson, 2003
Metulodontia nivea NH 13108 Russia AF506423 AF506423 Larsson and Larsson, 2003
Scytinostroma acystidiatum Dai 24608 China OQ689127 OQ629351 Present study
S. acystidiatum KUC20121019-32 Korea KJ668461 − Jang et al., 2016
S. aluta CBS 762.81 France MH861482 MH873221 Vu et al., 2019
S. alutum CBS 763.81 France MH861483 MH873222 Vu et al., 2019
S. alutum CBS 764.81 France MH861484 MH873223 Vu et al., 2019
S. alutum CBS 765.81 France MH861485 MH873224 Vu et al., 2019
S. alutum CBS 766.81 France MH861486 MH873225 Vu et al., 2019
S. caudisporum CBS 746.86 Gabon MH862030 NG073580 Vu et al., 2019
S. crispulum CBS 716.86 Reunion MH862013 MH873703 Vu et al., 2019
S. crispulum CBS 717.86 France MH862014 MH873704 Vu et al., 2019
S. crispulum CBS 718.86 France MH862015 MH873705 Vu et al., 2019
S. crispulum CBS 776.86 France MH862053 MH873741 Vu et al., 2019
S. decidens CBS 714.86 France MH862011 MH873701 Vu et al., 2019
S. decidens CBS 715.86 France MH862012 MH873702 Vu et al., 2019
S. duriusculum CBS 757.81 France MH861477 MH873216 Vu et al., 2019
S. duriusculum CBS 758.81 France MH861478 MH873217 Vu et al., 2019
S. hemidichophyticum CBS 702.84 Belgium MH861818 MH873509 Vu et al., 2019
S. hemidichophyticum CBS 759.81 France MH861479 MH873218 Vu et al., 2019
S. hemidichophyticum CBS 760.81 France MH861480 MH873219 Vu et al., 2019
S. jacksonii NH 6626 Canada AF506467 AF506467 Larsson and Larsson, 2003
S. jacksonii CBS 239.87 Canada MH862071 MH873759 Vu et al., 2019
S. macrospermum Dai 24606 China OQ689126 OQ629350 Present study
S. macrospermum M2138 Japan LC327052 − Ogura‐Tsujita et al., 2018
S. mediterraneense CBS 764.86 France MH862045 MH873732 Vu et al., 2019
S. mediterraneense CBS 765.86 France MH862046 MH873733 Vu et al., 2019
S. mediterraneense CBS 766.86 France MH862047 MH873734 Vu et al., 2019
S. microspermum CBS 238.87 Guadeloupe MH862070 − Vu et al., 2019
S. ochroleucum CBS 767.86 France MH862048 − Vu et al., 2019
S. ochroleucum CBS 768.86 France MH862049 MH873735 Vu et al., 2019
S. ochroleucum CBS 126049 USA MH864062 MH875517 Vu et al., 2019
S. odoratum KHL 8546 Sweden AF506469 AF506469 Larsson and Larsson, 2003
S. phaeosarcum CBS 728.81 Cote d’Ivoire MH861481 MH873205 Vu et al., 2019
S. portentosum CBS 503.48 Canada MH856447 MH873220 Vu et al., 2019
S. pseudopraestans CBS 737.91 − MH862322 MH873994 Vu et al., 2019
S. pseudopraestans CBS 738.91 − MH862323 MH873995 Vu et al., 2019
S. pseudopraestans CBS 739.91 − MH862324 MH873996 Vu et al., 2019
S. pseudopraestans CBS 740.91 − MH862325 MH873997 Vu et al., 2019
S. pseudopraestans CBS 741.91 − MH862326 MH873998 Vu et al., 2019
S. pseudopraestans CBS 742.91 − MH862327 − Vu et al., 2019
S. quintasianum CBS 749.86 Cote d’Ivoire MH862031 MH873719 Vu et al., 2019
S. quintasianum CBS 750.86 − MH862032 MH873720 Vu et al., 2019
S. quintasianum CBS 751.86 − MH862033 − Vu et al., 2019
S. renisporum CBS 771.86 Indonesia MH862051 MH873738 Vu et al., 2019
S. renisporum CBS 772.86 Indonesia MH862052 MH873739 Vu et al., 2019
S. yunnanense CLZhao 10758 China MT611445 − Wang et al., 2020
S. yunnanense CLZhao 10802 China MT611446 − Wang et al., 2020
S. yunnanense CLZhao 11010 China MT611447 − Wang et al., 2020
S. sp1 KUC20130725-13 Korea KJ668460 − Jang et al., 2016
S. sp2 MEL:2382679 Australia KP013042 − Rosenthal et al., 2017
S. sp3 UC2022985 USA KP814265 − Rosenthal et al., 2017
S. sp3 UC2022946 USA KP814564 − Rosenthal et al., 2017
S. sp4 MEL:2382745 Australia KP012928 − Rosenthal et al., 2017
S. sp5 LR-40 Chile MT366713 − Direct Submission
S. sp6 Het 803-1 USA OL989828 − Otto et al., 2021
S. sp6 NO 6-1-B USA OK173822 − Otto et al., 2021
S. sp6 iNAT:30809947 USA MZ267776 − Direct Submission
S. sp7 UoA SVB-F86 − MT975590 − Direct Submission
S. sp7 UC2023098 Canada KP814402 − Rosenthal et al., 2017
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The new species are in bold.

"-" represents the absence of a certainsequence in the species.

2.3. Phylogenetic analyses

Phylogenetic analyses were performed with the Maximum Parsimony (MP), Maximum Likelihood (ML), and Bayesian Inference (BI) methods. New sequences generated in this study, along with reference sequences retrieved from GenBank ( Table 1 ), were aligned by MAFFT 7 (Katoh et al., 2019; http://mafft.cbrc.jp/alignment/server/) using the “G-INS-i” strategy and manually adjusted in BioEdit (Hall, 1999). Unreliably aligned sections were removed before the analyses, and efforts were made to manually inspect and improve the alignment. The data matrix was edited in Mesquite v3.70. Confertobasidium olivaceoalbum (Bourdot & Galzin) Jülich and Metulodontia nivea (P. Karst.) Parmasto were selected as outgroups (Larsson and Larsson, 2003).

MP topology and bootstrap (BT) values obtained from 1,000 replicates were computed in PAUP* version 4.0b10 (Swofford, 2002). All characters were equally weighted, and the gaps were treated as missing data. Trees were inferred using the heuristic search option with tree-bisection reconnection (TBR) branch swapping and 1,000 random sequence additions. Max-trees were set to 5,000, branches of zero length were collapsed, and all parsimonious trees were saved. Clade robustness was assessed by a BT analysis with 1,000 replicates (Felsenstein, 1985). Descriptive tree statistics, such as 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.

RAxML 7.2.8 was used to construct ML trees for the combined dataset with the GTR+I+G model of site substitution, including estimation of Gamma-distributed rate heterogeneity and a proportion of invariant sites (Stamatakis, 2006). The branch support was evaluated with a bootstrapping method of 1000 replicates (Hillis and Bull, 1993).

The BI was conducted with MrBayes 3.2.6 in two independent runs, each of which had four chains for 5 million generations and started from random trees (Ronquist and Huelsenbeck, 2003). Trees were sampled every 1,000 generations. The first 25% of the sampled trees were discarded as burn-in, and the remaining ones were used to reconstruct a majority rule consensus and calculate Bayesian Posterior Probabilities (BPP) of the clades.

Branches that received BT supports for Maximum Parsimony (BP) and Maximum Likelihood (BS) greater than or equal to 75%, and BPP greater than or equal to 0.95 were considered as significantly supported. FigTree v1.4.4 and Treeview (Page, 1996) were used to visualize the resulting tree.

3. Results

3.1. Phylogenetic results

Two ITS and two nLSU sequences were generated in this study and were deposited in GenBank. Their accession numbers are specified in the phylogenetic tree ( Figure 1 ). The final ITS + nLSU dataset included 60 sequences representing 28 species and resulted in an alignment of 1,826 characters. Maximum parsimony analysis yielded one equally parsimonious tree (TL = 2833, CI = 0.502, HI = 0.846, RI = 0.424, and RC = 0.498). BI analysis and ML analysis resulted in a similar topology to the MP analysis, with an average standard deviation of split frequencies of 0.002601 (BI).

Figure 1.

Figure 1

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Phylogeny of Scytinostroma by Maximum Parsimony (MP) analysis based on combined ITS + nLSU dataset. Branches are labeled with bootstrap supports for Maximum Parsimony (BP) > 70%, Maximum Likelihood bootstrap (BS) > 70%, and Bayesian Posterior Probabilities (BPP) > 0.95, respectively. The new species are in bold.

The phylogeny ( Figure 1 ) inferred from the ITS + nLSU dataset demonstrated that two new species, Scytinostroma acystidiatum and S. macrospermum, clustered in the Scytinostroma clade. Moreover, Scytinostroma acystidiatum clustered with one sample from Korea (KUC20121019-32) formed an independent lineage with a robust support (BP = 100%, BS = 100%, and BPP = 1.00) and then closely related to S. renisporum Boidin, Lanq. & Gilles. S. macrospermum clustered with one sample from Japan (M2138), forming an independent lineage with a strong support (BP = 100%, BS = 100%, and BPP = 1.00).

3.2. Taxonomy

Scytinostroma acystidiatum Q.Y. Zhang, L.S. Bian & Q. Chen, sp. nov., Figures 2 , 3

Figure 2.

Figure 2

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Basidiomata of Scytinostroma acystidiatum (Holotype, Dai 24608). Scale bar = 1.0 cm. Photo by: Qiu-Yue Zhang.

Figure 3.

Figure 3

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Microscopic structures of Scytinostroma acystidiatum (Holotype, Dai 24608). (A) Basidiospores. (B) Cystidioles. (C) Basidia and basidioles. (D) A section of basidiomata. Drawings by: Qiu-Yue Zhang.

MycoBank no.: 848524

Type — China, Guizhou Province, Tongren, Fanjingshan, on fallen angiosperm branch, 13 July 2022, Dai 24608 (BJFC039842).

Etymology — Acystidiatum (Lat.): refers to the species lacking cystidia.

Basidiomata —Annual, resupinate, coriaceous, not separable from substrate, up to 7 cm long, 2 cm wide, and less than 0.1 mm thick at center. Hymenial surface smooth to locally tuberculate, cream to pale yellow; margin concolorous with hymenial surface, thinning out, and adnate.

Hyphal structure —Hyphal system dimitic; generative hyphae infrequent, simple septate, hyaline, thin-walled, rarely branched, 2–3 μm in diameter, IKI–, CB–; skeletal hyphae dominant, frequently dichotomously branched, tortuous, interwoven, thick-walled, dextrinoid, cyanophilous, 1–2.5 μm in diameter; tissues unchanged in KOH.

Hymenium —Cystidia absent; cystidioles present, clavate, some gradually tapering to the apex, thin-walled, hyaline, smooth, 12–18 × 2–4 μm; basidia clavate, with a basal simple septum and four sterigmata, thin-walled, smooth, 13–21 × 3.5–5 μm; basidioles similar to basidia in shape, but slightly smaller.

Spores —Basidiospores broadly ellipsoid with an apiculus, hyaline, thin-walled, smooth, occasionally with one or two guttules, amyloid, acyanophilous, (4.5–)4.7–7 × (3–)3.5–4.7(–5) μm, L = 5.68 μm, W = 4.02 μm, Q = 1.41 (n = 30/1).

Scytinostroma macrospermum Q.Y. Zhang, L.S. Bian & Q. Chen, sp. nov., Figures 4 , 5

Figure 4.

Figure 4

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Basidiomata of Scytinostroma macrospermum (Holotype, Dai 24606). Scale bar = 1.0 cm. Photo by: Qiu-Yue Zhang.

Figure 5.

Figure 5

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Microscopic structures of Scytinostroma macrospermum (Holotype, Dai 24606). (A) Basidiospores. (B) Basidia and basidioles. (C) Cystidia. (D) A section of basidiomata. Drawings by: Qiu-Yue Zhang.

MycoBank no.: 848525

Type —China, Chongqing, Jiangjin District, Simianshan National Scenic Spot, on rotten angiosperm wood, 10 July 2022, Dai 24606 (BJFC039838).

Etymology — Macrospermum (Lat.): refers to the species having large basidiospores.

Basidiomata —Annual, resupinate, coriaceous, not separable from substrate, up to 13.5 cm long, 3 cm wide, and less than 0.2 mm thick at center. Hymenial surface smooth to locally tuberculate, cream to straw yellow; margin concolorous with hymenial surface, slightly fimbricate.

Hyphal structure —Hyphal system dimitic; generative hyphae infrequent, simple septate, thin-walled, hyaline, rarely branched, 1.5–3 μm in diameter, IKI–, CB–; skeletal hyphae dominant, frequently dichotomously branched, interwoven, thick-walled, dextrinoid, cyanophilous, 1–3 μm in diameter; tissues unchanged in KOH.

Hymenium —Cystidia numerous, narrowly fusoid to cylindrical, thin-walled, smooth, 25–107 × 2.5–10 μm, embedded or projecting from hymenium up to 25 µm; basidia clavate, with a basal simple septum and four sterigmata, thin-walled, smooth, with some guttules, 30–45 × 6–8 μm; basidioles dominant, similar to basidia in shape, but slightly smaller.

Spores —Basidiospores ellipsoid with an apiculus, hyaline, thin-walled, smooth, occasionally with one or two guttules, inamyloid, acyanophilous, 9–11(–12) × (4–)4.5–5.5(–6) μm, L = 9.89 μm, W = 4.94 μm, Q = 2.00 (n = 30/1).

4. Discussion

Two new species, Scytinostroma acystidiatum and S. macrospermum, are described in this study based on morphological characteristics and phylogenetic analyses. The ITS + nLSU-based phylogeny ( Figure 1 ) shows the phylogenetic positions of the two new species in the genus Scytinostroma. In detail, the sequence of KUC20121019-32 from Korea, clustered together with Scytinostroma acystidiatum, and shares less than 1.5% sequence (ITS) dissimilarity (Jang et al., 2016). The sample KUC20121019-32 was collected in Odaesan National Park, South Korea, which has geographical proximity (eastern Asia) and a similar climate (subtropical climate) to Guizhou, China. So, we treat KUC20121019-32 as Scytinostroma acystidiatum. In addition, Scytinostroma acystidiatum grouped with S. renisporum with strong support (100% BP, 100% BS, 1.00 BPP, Figure 1 ). Scytinostroma renisporum is morphologically distinguished from S. acystidiatum by its membranaceous to paper-like basidiomata and larger gloeocystidia measuring 20–35 × 6–10 µm (Boidin and Lanquetin, 1987).

Morphologically, Scytinostroma alutum Lanq., S. arachnoideum (Peck) Gilb., S. cystidiatum Boidin, S. hemidichophyticum Pouzar, S. portentosum (Berk. & M.A. Curtis) Donk, and S. yunnanense C.L. Zhao are similar to S. acystidiatum by sharing amyloid basidiospores. However, S. alutum differs from S. acystidiatum by its resupinate to effuse-reflexed basidiomata with cracked hymenophore, larger basidia (40–65 × 5–6 µm vs. 13–21 × 3.5–5 μm), and bigger basidiospores (5.3–7.2 × 5.7–7.3 μm vs. 4.7–7 × 3.5–4.7 μm; Lanquetin, 1984). Scytinostroma arachnoideum is separated from S. acystidiatum by its cottony basidiomata with white rhizomorphs and smaller basidiospores (3.5–4.5 × 3–3.5 μm vs. 4.7–7 × 3.5–4.7 μm; Gilbertson, 1962). Scytinostroma cystidiatum, S. hemidichophyticum, and S. portentosum are separated from S. acystidiatum by the presence of cystidia (Donk, 1956; Boidin, 1960; Pouzar, 1966). S. yunnanense differs from S. acystidiatum by its white to cream basidiomata and shorter basidiospores (4.5–5.5 μm vs. 4.7–7 μm in length; Wang et al., 2020).

Phylogenetically, the sequence of M2138 from Japan, clustered together with Scytinostroma macrospermum and formed an independent lineage with less than 1.5% sequence (ITS) dissimilarity (Ogura-Tsujita et al., 2018). The sample M2138 was collected in Kagoshima, Japan, which has geographical proximity (eastern Asia) and a similar climate (subtropical climate) to Chongqing, China. So, we treat M2138 as Scytinostroma macrospermum ( Figure 1 ). Morphologically, Scytinostroma ochroleucum (Bres. & Torrend) Donk resembles S. macrospermum by resupinate, cream-colored to pale ochraceous basidiomata, but the former is different from the latter by its larger basidia (35–85 × 6.5–9 µm vs. 30–45 × 6–8 μm), and larger basidiospores (9–14 × 5–7 µm vs. 9–11 × 4.5–5.5 μm; Donk, 1956). Scytinostroma phaeosarcum Boidin & Lanq. resembles S. macrospermum by the approximately same size of basidiospores (8–10 × 4.5–5.5 μm), while S. phaeosarcum differs from S. macrospermum by its basidiomata becoming brown when bruised and thinner basidia (3–5 μm vs. 6–8 μm in width; Boidin and Lanquetin, 1977). In addition, Scytinostroma macrospermum is similar to S. decidens Boidin, Gilles & Lanq., S. jacksonii Boidin and S. mediterraneense Boidin & Lanq. by sharing large cystidia (> 100 μm in length) and inamyloid basidiospores. However, the latter three species distinctly differ from S. macrospermum by their obviously narrower basidiospores (2.5–3.5 μm in width vs. 4.5–5.5 μm in width, Boidin, 1981; Boidin and Lanquetin, 1987; Nakasone and Micales, 1988).

Wood-rotting fungi as an important group within the Basidiomycota are known for their ecological role in the forest ecosystem in terms of decaying living and dead trees and recycling nutrients in forest ecosystems (Dai et al., 2007; Yuan et al., 2021; Yuan et al., 2022). However, the diversity and taxonomy of these fungi remain not well known, and many new species have been described recently because of the application of molecular phylogeny (Dai et al., 2021; Mao et al., 2023; Wang et al., 2021; Wang et al., 2022; Wu et al., 2022a; Wu et al., 2022b; Zhou et al., 2021). Similarly, despite numerous species of Scytinostroma have been described, many unknown species or unnamed sequences still exist (Scytinostroma sp., Figure 1 ). Consequently, with the application of molecular phylogeny, the diversity and systematics will be outlined by further studies based on more samples worldwide.

Data availability statement

The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found below: https://www.ncbi.nlm.nih.gov/genbank/, OQ629350, OQ629351, OQ689126, OQ689127.

Author contributions

Q-YZ, H-GL, L-SB, and QC designed the research and contributed to data analysis and interpretation. Q-YZ prepared the samples and drafted the manuscript. H-GL, L-SB and QC discussed the results and edited the manuscript. All authors contributed to the article and approved the submitted version.

Acknowledgments

The authors would like to express their deep appreciations to Prof. Yu-Cheng Dai (Beijing Forestry University, China) for allowing us to study his specimens.

Funding Statement

The research was financed by the National Natural Science Foundation of China (Project nos. 32100014, 31800018) and Fundamental Research Funds for the Central Non-profit Research Institution of the Chinese Academy of Forestry (Project No. CAFYBB2021MA007).

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Publisher’s note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found below: https://www.ncbi.nlm.nih.gov/genbank/, OQ629350, OQ629351, OQ689126, OQ689127.

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