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Research Article
Molecular phylogeny and morphology of the genus Fuscoporia (Hymenochaetales, Basidiomycota) and reveal three new species of the F. ferrea group
expand article infoQian Chen§, Han Chen, Cheng-Hang Luo, Xiao-Hong Lai
‡ Chongqing Jiaotong University, Chongqing, China
§ Tongji University, Shanghai, China
Open Access

Abstract

Fuscoporia is a polypore genus of Hymenochaetaceae that causes wood decay, although some species in the genus have medicinal values. Phylogenetic analyses of concatenated ITS1-5.8S-ITS2-nLSU sequence data and morphological features identified three new species, F. eucalypticola, F. resupinata and F. subtropica from Australia, China and Malaysia, and these new species derived from the Fuscoporia ferrea group. These three species are illustrated and described. A key to resupinate species of Fuscoporia without mycelial setae in the world is provided.

Key words

Hymenochaetaceae, phylogeny, polypore, taxonomy

Introduction

The genus Fuscoporia Murrill (Hymenochaetales, Basidiomycota) with F. ferruginosa (Schrad.) Murrill as the type species was first described in 1907 (Murrill 1907). For a long time, it has been considered as a synonym of Phellinus Quél. sensu lato (Gilbertson 1979; Larsen and Cobb-Poulle 1990; Ryvarden and Gilbertson 1994; Ryvarden 2004). Phylogenetic studies confirmed that the currently recognized genus of Fuscoporia is monophyletic (Wagner and Fischer 2001, 2002; Wu et al. 2022a). Fuscoporia is characterized by an almost light to dark brown, resupinate, effused-reflexed to pileate basidiomata, dimitic hyphal system with generative hyphae bearing crystals, presence of hymenial setae in most species, and hyaline, thin-walled, smooth basidiospores (Fiasson and Niemelä 1984; Chen et al. 2020; Wu et al. 2022a). The species is very rich and 104 Fuscoporia species have been recognized (Wu et al. 2022a, b; Chen et al. 2023a, b; https://www.mycobank.org; accessed on 08-9-2024). Among them, 62 species were described during the last five years (Chen and Dai 2019; Chen et al. 2019, 2020, 2022, 2023a, b; Du et al. 2020; Tchoumi et al. 2020; Vlasák et al. 2020; Yuan et al. 2020; Raymundo 2021; Hussain et al. 2022; Wu et al. 2022a; Cho et al. 2023; Olou et al. 2023; Bittencourt et al. 2024).

Some species of Fuscoporia are difficult to identify because most morphological features of these species overlap. The recent studies revealed some traditional species of Fuscoporia are actually the species complex, such as, Fuscoporia contigua (Pers.) G. Cunn. (Cunningham 1948) was considered as a single species with variable basidiospores (oblong-ellipsoid or cylindric; Niemelä 2005; Dai 2010; Ryvarden and Melo 2017), but two Asian species were derived from F. contigua (Chen et al. 2019). A more comprehensive study (Chen et al. 2020, 2022; Wu et al. 2022a) of the genus revealed Fuscoporia contigua is actually the most complex speciesin that its members do not share a common geographic distribution and host preference. So far, fourteen species have been published in F. contigua group. The taxonomic status of some species in the genus Fuscoporia is in need of re-evaluation.

Fuscoporia ferrea (Pers.) G. Cunn. (1948) was characterized by resupinate, annual to perennial basidiomata, cylindric spores and distribution in the Northern Hemisphere (Cunningham 1948; Ryvarden and Gilbertson 1994; Lowe 1966). Based on time divergence, the early divergence of the Fuscoporia species was inferred to occur in subtropics of southern Asia with a resupinate fruiting body, and Fuscoporia ferrea group is the oldest lineage in the genus with stem age estimated around 49.52 Myr (Hussain et al. 2022). Some Asian specimens previously identified as Fuscoporia ferrea were confirmed as different species based on morphological examinations and phylogenetic analyses, and described as F. ramulicola Y.C. Dai & Q. Chen, F. subferrea Q. Chen & Y. Yuan and F. yunnanensis Y.C. Dai (Dai 2010; Chen and Yuan 2017; Chen and Dai 2019). In addition, F. punctatiformis (Murrill) Zmitr., Malysheva & Spirin was combined (Spirin et al. 2006). So five species comprise the F. ferrea complex and are characterized by resupinate basidiomata, absence of mycelial setae, presence of hymenial setae and cystidioles, and cylindric basidiospores (Chen et al. 2020).

In the process of exploring of wood-decaying fungi, brown and resupinate specimens were collected from southern Asia and Australia, and their morphological characteristics, taxonomic relationships and phylogenetic affinities were analyzed. Three new taxa were confirmed within Fuscoporia ferrea group, and they are described and illustrated. A key to resupinate and mycelial setaeless species of Fuscoporia in the world is provided.

Materials and methods

Morphological studies

The research specimens are conserved in the herbarium of Microbiology, Beijing Forestry University (BJFC). The macroscopic color codes follow Petersen (1996) and the microscopic analyses follow Wang et al. (2023, 2024) and Zhao et al. (2023, 2024) using a Nikon Eclipse 80i microscope with phase contrast illumination. Samples for microscopic examination and drawings were prepared from slides stained with Cotton Blue follow Zhang et al. (2023). The following abbreviations are used: CB– = acyanophilous, IKI– = neither amyloid nor dextrinoid, L = mean length of all spore, W = mean width of all spore, Q = L/W ratios, n (a/b) = number of measured spores(a) form number of specimens (b).

Molecular methods

According to the manufacturer’s instructions, a CTAB rapid plant genome extraction kit (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to obtain PCR products from dried samples. To generate PCR amplicons, the following primer pairs were used: ITS4 and ITS5 for the ITS1-5.8S-ITS2 region (White et al. 1990), LR0R and LR7 for the nLSU region (Vilgalys and Hester 1990). The PCR procedure was as follows: for the ITS1-5.8S-ITS2 region initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 54 °C for 45 s and 72 °C for 1 min, and a final extension of 72 °C for 10 min; for nLSU, initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 1 min, 50 °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 by the Beijing Genomics Institute with the same primers. All newly generated sequences were deposited at GenBank and listed in Table 1 (http://www.ncbi.nlm.nih.gov/genbank). Besides the newly generated sequences for this study, other related sequences downloaded from GenBank based on Chen et al. (2023b) and Wu et al. (2022a, b) to explore the phylogenetic position of the newly sequenced specimens in Fuscoporia.

Table 1.

Species name, specimens, origin and GenBank accession number of sequences used in this study.

Species name Specimens Origin GenBank accession no.
ITS nLSU
Fuscoporia acutimarginata Dai 15137 China MH050751 MH050765
F. acutimarginata Dai 16892 China MH050752 MH050766
F. ambigua Cui 9244 China MN816706 MN809995
F. ambigua JV 0509/151 USA MN816707 MN809996
F. americana JV 1209/3-J USA MG008466
F. americana JV 1209/100 USA KJ940022 MG008467
F. atlantica SP 445618 Brazil KP058515 KP058517
F. atlantica SP 465829 Brazil KP058514 KP058516
F. australasica Dai 15625 China MN816726 MN810018
F. australasica Dai 15636 China MG008397 MG008450
F. australiana Dai 18672 Australia MN816703 MN810014
F. australiana Dai 18879 Australia MN816705 MN810015
F. bambusae Dai 16599 Thailand MN816711 MN809999
F. bambusae Dai 16615 Thailand MN816715 MN810001
F. caymanensis JV 1908/74 French Guiana MT676832 MT676833
F. caymanensis JV 1408/5 Costa Rica MW009110 MW009109
F. callimorpha Dai 17388 Brazil MN121765 MN121824
F. callimorpha Doll 868 Unknown MN816701 MN809992
F. chinensis Dai 15713 China MN816721 MN810008
F. chinensis Cui 11209 China MN121767 MN121826
F. chrysea JV 1607/106 Costa Rica MN816736 MN810027
F. centroamericana JV 1607/93 Costa Rica MG008444 MG008460
F. centroamericana O 908267 Costa Rica MG008443
F. contigua Dai 16025 USA MG008401 MG008454
F. contigua Dai 13567A Romania MG008402 MG008455
F. costaricana JV 1407/92 Costa Rica MG008446 MG008461
F. costaricana JV 1504/85 Costa Rica MG008413 MG478454
F. dhofarensis ATN-007 Oman OP593104 OP593105
F. dolichoseta SFC20191015-23 Korea ON427765 ON427795
F. dolichoseta SFC20161006-16 Korea ON427789 ON427817
F. eucalypti Dai 18783 Australia MN816730 MN810021
F. eucalypti Dai 18792 Australia MN816731 MN810022
F. eucalypticola Dai 18592A Australia PP732562 PP732631
F. eucalypticola Dai 18683 Australia PP732563 PP732632
F. ferrea MUCL 45984 France KX961112 KY189112
F. ferrea Cui 11801 China KX961101 KY189101
F. ferruginosa JV 0408/28 Czech Republic KX961103 KY189103
F. ferruginosa Dai 13200 France MN816702 MN809993
F. gilva JV 0709/75 USA MN816720 MN810007
F. gilva JV 1209/65 USA MN816719 MN810006
gilva URM 83957 Brazil MH392545 MH407344
gilvoides SFC2018042‐12 Korea ON427763 ON427793
gilvoides MUGBt Pakistan ON427781 ON427810
F. hainanensis Dai 16105 China ON520809
F. hainanensis Dai 16110 China ON520810
F. hawaiana JV 2208/H-22-J USA OQ817709 OQ817855
F. hawaiana JV 2208/H-30-J USA OQ817710 OQ817856
F. insolita Spirin 5251 Russia KJ677113
F. insolita Spirin 5208 Russia MN816724 MN810016
F. karsteniana Dai 16552 China MN816716 MN810002
F. karsteniana Dai 11403 China MN816717 MN810003
F. kenyana Dai 19205 Kenya OP580527 OP580521
F. kenyana Dai 19202 Kenya OP580526 OP580520
F. koreana SFC20150625-05 Korea ON427776 ON427805
F. koreana SFC20160726-93 Korea ON427762 ON427792
F. latispora JV 1109/48 USA MG008439 MG008468
F. latispora JV 0610/VII-Kout Mexico MG008436 MG008469
F. licnoides URM 84107 Brazil MH392556 MH407355
F. licnoides URM 83001 Brazil MH392561 MH407357
F. marquesiana URM 83094 Brazil MH392544 MH407343
F. minutissima JV 2208/H12-J USA OQ817711 OQ817857
F. minutissima JV 2208/H16-J USA OQ817712 OQ817858
F. monticola Dai 10909 China MG008410
F. monticola Dai 11860 China MG008406 MG008457
F. palomari JV 1004/5-J USA MN816737
F. palomari JV 1305/3-J USA MN816738 MN810028
F. plumeriae Dai 17814 Singapore MN816714 MN810011
F. plumeriae Dai 18858 Australia MN816712 MN810010
F. punctatiformis Dai 17443 Brazil MH050755 MH050764
F. punctatiformis Doll#872a Brazil MH050753
F. pulviniformis CMW 48060 South Africa MH599101 MH599125
F. pulviniformis CMW 48600 South Africa MH599102 MH599127
F. ramulicola Dai 15723 China MH050749 MH050762
F. ramulicola Dai 16155 China MH050750 MH050763
F. reticulata SFC20121010-19 Korea ON427766
F. reticulata SFC20160115-16 Korea ON427761 ON427791
F. rhabarbarina Dai 16550 China MN816744 MN810036
F. rhabarbarina Dai 16226 China MN816743 MN810035
F. resupinata Dai 20455 China PP732567 PP732636
F. resupinata Dai 20422 China PP732568 PP732637
F. resupinata Dai 21201 Malaysia PP732569 PP732638
F. roseocinerea JV 1407/84 Costa Rica MN816740 MN810030
F. roseocinerea JV 1109/78-J USA MN816742 MN810032
F. rufitincta JV 1008/25 USA KJ940029 KX058575
F. rufitincta JV 0904/142 USA KJ940030 KX058574
F. sarcites JV 0402/20K Venezuela MZ264225 MZ264218
F. scruposa CMW 48145 South Africa MH599105 MH599130
F. scruposa CMW 47749 South Africa MH599106 MH599129
F. semiarida URM 83800 Brazil MH392562 MH407361
F. semiarida URM 82510 Brazil MH392563 MH407362
F. semicephala SFC20170524-08 Korea ON427764 ON427794
F. semicephala SFC20170712-20 Korea ON427787 ON427815
F. senex MEL 2382630 Australia KP012992
F. senex KAUNP MK41 Sri Lanka KP794600
F. septoseta Dai 12820 USA MG008405 MN810033
F. septoseta JV 0509/78 USA MG008404
F. setifera Dai 15710 China MH050758 MH050767
F. setifera Dai 15706 China MH050759 MH050769
F. shoreae Dai 17806 Singapore MN816734 MN810025
F. shoreae Dai 17818 Singapore MN816735 MN810026
F. sinica Dai 15468 China MG008412 MG008459
F. sinica Dai 15489 China MG008407 MG008458
F. sinuosa Dai 20498 China MZ264226 MZ264219
F. sinuosa Dai 20499 China MZ264227 MZ264220
F. subchrysea Dai 16201 China MN816708 MN809997
F. subchrysea Dai 17656 China MN816709 MN809998
F. subferrea Dai 16326 China KX961097 KY053472
F. subferrea Dai 16327 China KX961098 KY053473
F. submurina Dai 19501 Sri Lanka MZ264229 MZ264222
F. submurina Dai 19655 Sri Lanka MZ264228 MZ264221
F. subtropica Dai 20476 China PP732564 PP732633
F. subtropica Dai 19957 China PP732565 PP732634
F. subtropica Dai 22604 China PP732566 PP732635
F. torulosa JV 1405/2 Czech Republic KX961106 KY189106
F. torulosa Dai 15518 China MN816732 MN810023
F. viticola JV 0911/6 Czech Republic KX961110
F. viticola He 2123 USA MN816725 MN810017
F. wahlbergii JV 1312/20-Kout Spain MN816727 MG008462
F. wahlbergii JV 0709/169-J USA MN816728
F. yunnanensis Cui 8182 China MH050756 MN810029
F. yunnanensis Dai 15637 China MH050757 MH050768
Outgroups
Coniferiporia weirii CFS 504 Canada AY829341 AY829345
Phellinidium fragrans CBS 202.90 USA AY558619 AY05027

Phylogenetic analysis

The following software was used for data processing and phylogenetic analysis: BioEdit (Hall 1999), ClustalX (Thompson et al. 1997) and MAFFT (http://mafft.cbrc.jp/alignment/server/, Katoh et al. 2017) for sequences and manually adjusted, PhyloSuite v.1.2.2 (Zhang et al. 2020) for concatenated the separate alignments, PAUP* 4.0b10 (Swofford 2002) for maximum parsimony (MP) analysis, raxmlGUI 1.2 (Silvestro and Michalak 2012) for maximum likelihood (ML) analysis and MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003) for Besian Inference (BI), TreeView 1.5.0 and FigTree version 1.4.4 (Rambaut 2018) to show the phylogenetic tree. The best topologies from ML analyses are shown in this study and the final alignments and the retrieved topologies has been deposited at TreeBASE (http://treebase.org/treebase-web/home.html), study ID: 31700.

In Maximum likelihood (ML) methods, statistical support values were obtained by using nonparametric bootstrapping with 1000 replicates, with default settings for all parameters. For BI analysis, the best-fit partitioning scheme and substitution model were determined by using ModelFinder (Kalyaanamoorthy et al. 2017). Tree was sampled every 1000 generations, starting from random trees with four chains for 2.5 million generations. In maximum parsimony (MP) analysis, tree was inferred using the heuristic search option with tree bisection reconnection (TBR) branch swapping and 1000 random sequence additions. The maxtrees parameter was set to 5000, branches of zero length were collapsed, and all parsimonious trees were saved. Clade robustness was assessed by a bootstrap analysis with 1000 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. The three phylogenetic methods produced a similar topology for each dataset, so, only the topology of the ML tree is presented along. Branches that received bootstrap support for ML and MP not less than 75% and BPP not less than 0.95 were considered as significantly supported.

Results

Molecular phylogeny

In this study, the data set of ITS and nLSU region included 118 ITS and 110 nLSU sequences from 121 samples, representing 61 species of Fuscoporia and Coniferiporia weirii (Murrill) L.W. Zhou & Y.C. Dai and Phellinidium fragrans (M.J. Larsen & Lombard) Nuss as the outgroups (Table 1, Fig. 1) based on previous studies (Chen and Yuan 2017). The dataset had an aligned length of 2224 characters, of which 1392 were constant, 120 variable but parsimony-uninformative, and 712 parsimony-informative. MP analysis yielded four similar topologies (TL = 3361, CI = 0.406, RI = 0.840, RC = 0.341, HI = 0.594). The BI analysis resulted in a concordant topology with an average standard deviation of split frequencies of 0.002648. The best model suggested by MrModeltest and applied in Bayesian analysis was GTR+F+I+G4 for ITS1+ITS2, K2P for 5.8s and K2P+I+G4 for nLSU. MP and BI analysis also resulted in a topology similar to that of the ML analysis. The seven specimens formed three lineages, named as Fuscoporia eucalypticola, F. resupinata and F. subtropica, with high support (100 in ML/1.00 in BI/100 in MP, respectively), which clustered together with F. ferrea, F. punctatiformis, F. ramulicola, F. subferrea and F. yunnanensis, in the F. ferrea clade with strongly support (100 in ML/1.00 in BI/100 in MP).

Figure 1. 

Maximum Likelihood (ML) tree illustrating the phylogeny of Fuscoporia and related species generated inferred from a combined ITS1-5.8S-ITS2-nLSU dataset. Statistical values (ML//BI/MP) are indicated for each node that received bootstrap support from ML and MP ≥ 75% and BPP ≥ 0.90. Names of new species are in bolds.

Taxonomy

Fuscoporia eucalypticola Q. Chen, sp. nov.

MycoBank No: 853957
Figs 2, 3

Holotype

Australia • Victoria, Yarra Ranges National Park, on fallen branch of Eucalyptus, 9 May 2018, Dai 18592A (BJFC 027061).

Etymology

Eucalypticola (Lat.): refere to the species growing on Eucalyptus.

Description

Basidiomata. Annual, resupinate, inseparable from the substrate, without odor or taste and corky when fresh, rigid when dry, up to 20 cm long, 3 cm wide and 1.5 mm thick at center. Pore olivaceous buff to greyish brown; sterile margin narrow or almost lacking, buff, up to 1 mm wide; pores irregular or sinuous, 3–5 per mm; dissepiments thin, entire, abundant setae seen in tube cavities (under lens). Subiculum clay-buff, corky, about 0.1 mm thick. Tubes olivaceous buff, up to 1 mm long.

Figure 2. 

Microscopic structures of Fuscoporia eucalypticola (Dai 18592A, holotype) A basidiospores B basidia and basidioles C cystidioles D hymenial setae E generative hyphae at dissepiment edge F hyphae from tube trama G hyphae from subiculum.

Hyphal structure. Hyphal system dimitic; generative hyphae simple septate; tissue becoming black in KOH.

Subiculum. Generative hyphae infrequently, thin-walled, frequently branched, simple septate, 1.5–2.5 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a medium lumen, unbranched, aseptate, flexuous, strongly interwoven, 2–3 μm in diam.

Figure 3. 

Basidiomata of Fuscoporia eucalypticola A Dai 18592A (holotype) B Dai 18683.

Tubes. Generative hyphae infrequent, mostly present at subhymenium, hyaline, thin-walled, frequently branched, simple septate, 1.5–2.5 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a wide lumen, unbranched, aseptate, loosely interwoven to subparallel along the tubes, 2–3 μm in diam. Setae frequent, mostly originating from hymenium, subulate, dark brown, thick-walled, 30–60 × 4–6 μm; fusoid cystidioles frequent, hyaline and thin-walled, 25–32 × 2–4 μm; basidia barrel-shaped, with four sterigmata and a simple septum at the base, 20–25 × 4–7 μm; basidioles in shape similar to basidia, but slightly smaller.

Basidiospores. Basidiospores cylindric, hyaline, thin-walled, smooth, IKI–, CB–, sometimes with a small guttule, 6.2–8 × (2–)2.1–3 μm, L = 7.03 μm, W = 2.37 μm, Q = 2.86–3.05 (n = 60/2).

Additional specimen examined

Australia • Melbourne, Dandenong Ranges Botanical Garden, on fallen branch of Eucalyptus, 12 May 2018, Dai 18683 (BJFC 027152).

Fuscoporia resupinata Q. Chen, sp. nov.

MycoBank No: 853956
Figs 4, 5

Holotype

China • Yunnan Province, Pu’er, Taiyanghe National Forest Park, on dead angiosperm tree, 17 August 2019, Dai 20455 (BJFC032123).

Etymology

Resupinata (Lat.): refers to the species having resupinate basidiomata.

Description

Basidiomata. Annual, resupinate, inseparable from the substrate, without odor or taste and corky when fresh, rigid when dry, up to 10.6 cm long, 4 cm wide and 1.2 mm thick at center. Pore surface fawn when fresh, snuff brown when dry; sterile margin indistinct, honey-yellow when dry, up to 1 mm wide, paler than color than the pore surface; pores circular to angular, 5–7 per mm; dissepiments thin, entire, abundant setae seen in tube cavities (under lens). Subiculum honey yellow, corky, about 0.2 mm thick. Tubes grayish brown, paler contrasting with pores, rigid, up to 1 mm long.

Figure 4. 

Microscopic structures of Fuscoporia resupinata (holotype, Dai 20455) A basidiospores B basidia and basidioles C cystidioles D hymenial setae E generative hyphae at dissepiment edge F hyphae from tube trama G hyphae from subiculum.

Hyphal structure. Hyphal system dimitic; generative hyphae simple septate; tissue becoming black in KOH.

Subiculum. Generative hyphae infrequently, thin-walled, frequently branched, simple septate, 1–1.5 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a narrow to medium lumen, unbranched, aseptate, flexuous, strongly interwoven, 3–4 μm in diam.

Figure 5. 

Basidiomata of Fuscoporia resupinata A Dai 20455 (holotype) B Dai 20422.

Tubes. Generative hyphae infrequent, mostly present at subhymenium, hyaline, thin-walled, frequently branched, simple septate, 1–2 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a narrow to medium lumen, unbranched, aseptate, loosely interwoven, 2–5 μm in diam. Setae frequent, mostly originating from hymenium, subulate, dark brown, thick-walled, 20–30 × 5–7 μm; fusoid cystidioles frequent, hyaline and thin-walled, sometimes covered with crystals, 8–12 × 3.5–5 μm; basidia barrel-shaped, with four sterigmata and a simple septum at the base, 16–20 × 6–8 μm; basidioles in shape similar to basidia, but slightly smaller.

Basidiospores. Basidiospores cylindric, hyaline, thin-walled, smooth, usually glued in tetrads, IKI–, CB–, sometimes with guttules, (5.4–)5.5–7(–7.2) × (2.4–)2.5–3 μm, L = 6.38 μm, W = 2.71 μm, Q = 2.29–2.44 (n = 60/2).

Additional specimens examined

China • Yunnan Province, Xinping County, Longquan Park, on fallen angiosperm branch, 16 August 2019, Dai 20422 (BJFC 032090). Malaysia • Selangor, Kota Damansara, Community Forest Reserve, on dead angiosperm tree, 7 December 2019, Dai 21201 (BJFC 032855).

Fuscoporia subtropica Q. Chen, sp. nov.

MycoBank No: 853958
Figs 6, 7

Holotype

China • Yunnan Province, Wenshan Zhuang and Miao Autonomous Region, Xichou County, Xiaoqiaogou Forest Farm, on fallen angiosperm trunk, 29 June 2019, Dai 19957 (BJFC 031631).

Etymology

Subtropica (Lat.): refers to the species being found in subtropical area.

Description

Basidiomata. Annual, resupinate, inseparable from the substrate, without odor or taste and corky when fresh, rigid when dry, up to 15 cm long, 8 cm wide and 2.5 mm thick at center. Pore surface grayish brown to honey-yellow; sterile margin indistinct, curry-yellow, up to 1 mm wide; pores irregular to angular, sometimes sinuous, 3–5 per mm; dissepiments thin, entire, abundant setae seen in tube cavities (under lens). Subiculum clay-buff, corky, about 0.5 mm thick. Tubes olivaceous buff, up to 2 mm long.

Figure 6. 

Microscopic structures of Fuscoporia subtropica (holotype, Dai 19957) A basidiospores B basidia and basidioles C cystidioles D hymenial setae E generative hyphae at dissepiment edge F hyphae from tube trama G hyphae from subiculum.

Hyphal structure. Hyphal system dimitic; generative hyphae simple septate; tissue becoming black in KOH.

Subiculum. Generative hyphae infrequently, thin-walled, frequently branched, simple septate, 2–3 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a medium lumen, unbranched, aseptate, flexuous, strongly interwoven, 3–4 μm in diam.

Figure 7. 

Basidiomata of Fuscoporia subtropica A Dai 19957 (holotype) B Dai 22604.

Tubes. Generative hyphae infrequent, mostly present at subhymenium, hyaline, thin-walled, frequently branched, simple septate, 2–3 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a narrow to medium lumen, unbranched, aseptate, loosely interwoven, 2–4 μm in diam. Setae frequent, mostly originating from hymenium, subulate, dark brown, thick-walled, 35–55 × 4–7 μm; fusoid cystidioles frequent, hyaline and thin-walled, 18–26 × 4–6 μm; basidia barrel-shaped, with four sterigmata and a simple septum at the base, 14–18 × 4–6 μm; basidioles in shape similar to basidia, but slightly smaller.

Basidiospores. Basidiospores cylindric, hyaline, thin-walled, smooth, IKI–, CB–, (5.5–)6–7.5(–8) × 2–3(–3.2) μm, L = 6.91 μm, W = 2.66 μm, Q = 2.32–2.71 (n = 50/2).

Additional specimens examined

China • Yunnan Province, Pu’er, Pu’er Forest Park, Xiniuping Scenic Area, on fallen angiosperm branch, 17 August 2019, Dai 20476 (BJFC 032144); • Taiyanghe National Forest Park, on fallen angiosperm branch, 8 July 2021, Dai 22604 (BJFC 037178).

Discussion

Fuscoporia is a polypore genus causing wood decay, associated with angiosperms and gymnosperms (Dai et al. 2007; Wu et al. 2022a, b; Yuan et al. 2023; Zhao et al. 2024). The medicinal potential of Fuscoporia, such as F. gilva and F. torulosa, was confirmed by modern studies (Deveci et al. 2019; Wu et al. 2019; Duong and Dang 2022). Fuscoporia is widely distributed in Asia (Bakshi et al. 1970; Dai 2010; Chen and Dai 2019; Chen et al. 2019, 2020; Du et al. 2020), Africa (Reid 1975; Ryvarden and Johansen 1980; Chen et al. 2023b), Australia (Chen et al. 2020), Europe (Donk 1960; Ryvarden and Gilbertson 1994, Ryvarden and Melo 2017), South America and North America (Larsen and Cobb-Poulle 1990; Chen et al. 2019; Wu et al. 2022a, b; Chen et al. 2023a). In this study, three new species of Fuscoporia are described based on molecular analyses and morphological features in Australia and southern Asia.

The recent studies (Chen and Dai 2019; Chen et al. 2020) demonstrated that the species of Fuscoporia ferrea was a complex species. We recognized eight species in the group: Fuscoporia ferrea sensu stricto (Ryvarden and Gilbertson 1994; Lowe 1966) in the Northern Hemisphere, such as Northern China, Europe and North America; three new species reported in this study, F. resupinata and F. subtropica from southern Asia, F. eucalypticola from Australia; F. ramulicola (Chen and Dai 2019), F. subferrea (Chen and Yuan 2017) and F. yunnanensis (Dai 2010) also distribution in south China; F. punctatiformis in Neotropics (Spirin et al. 2006), such as Brazil and USA. Eight species clustered into a clade with high statistical support (100/1.00/100) in phylogenetic analysis published in this study. The members of the Fuscoporia ferrea group differ from other species in the genus by its resupinate basidiomata, presence of hymenial setae and cystidioles, absence of mycelial setae, and cylindric basidiospores (Dai 2010; Chen and Yuan 2017; Chen and Dai 2019).

The species in the Fuscoporia ferrea group have similar morphological characteristics, which sometimes may be confused. However, F. ferrea and F. punctatiformis can be segregated from the three new species by their perennial basidiomata (Lowe 1966; Spirin et al. 2006). The remaining species of the F. ferrea group have annual basidiomata and are distributed in southern Asia, except for F. eucalypticola, which is from Australia and grows on Eucalyptus. Furthermore, two samples of F. eucalypticola formed a well-supported lineage (100/1.00/100), indicating that they are phylogenetically distinct from other species in Fig. 1. Fuscoporia eucalypticola is closely related to F. subtropica in the phylogenetic tree and also has similar macromorphology in sharing annual basidiomata, irregular to angular, sometimes sinuous and bigger porse (3–5 per mm), but the latter differs in being without guttule in basidiospores and its distribution in Yunnan provinces, China.

Southern Asia is among the regions with the highest fungal biodiversity, especially in southern China (Dai et al. 2021; Zhou et al. 2023). Fuscoporia subtropica, F. yunnanensis, F. ramulicola and F. resupinataare distributed in Yunnan provinces, China, F. resupinataare also distributed in Malaysia, F. subferrea is distributed in Hainan provinces, China, which is an island. Macromorphologically the two new species, Fuscoporia resupinata and F. subtropica, are also similar to F. subferrea, F. ramulicola and F. yunnanensis, but F. resupinata differs from F. yunnanensis and F. subferrea by its medium-sized pores (5–7 per mm in F. resupinata vs. 3–4 per mm in F. yunnanensis, 7–10 per mm in F. subferrea; Chen and Yuan 2017); differs from F. ramulicola by its wider spores (2.5–3 μm, Q = 2.29–2.44 in F. resupinata vs. 2–2.5 μm, Q = 2.57–2.88 in F. ramulicola; Chen and Dai 2019). Fuscoporia subtropica differs from F. ramulicola and F. subferrea by its larger pores (3–5 per mm in F. subtropica vs. 6–7 per mm in F. ramulicola, 7–10 per mm in F. subferrea), differs from F. yunnanensis by its irregular pores (Dai 2010). Fuscoporia resupinata resembles F. subtropica by having annual and resupinate basidiomata, cylindric spores, but the former has smaller pores (5–7 per mm vs. 3–5 per mm), shorter fusoid cystidioles (8–12 μm vs. 18–26 μm), and its basidiospores sometimes with guttules.

A key to resupinate and mycelial setaeless species of Fuscoporia in the world

1 Basidiomata perennial 2
Basidiomata annual to biennial 4
2 Basidiospores narrowly ovoid to narrow ellipsoid F. montana Y.C. Dai & Niemela
Basidiospores cylindric to subcylindrical 3
3 Basidiospores 4–6 × 1.5–2 μm F. punctatiformis
Basidiospores 6–7.8 × 2–2.5 μm F. ferrea
4 Pores 3–5 per mm 5
Pores 5–10 per mm 7
5 Pores circular, dissepiments entire and matted F. yunnanensis
Pores sinuous or irregular or daedaleoid, dissepiments entire and slightly lacerate with age 6
6 Basidiospores without guttule, Q = 2.32–2.71, distribution in China F. subtropica
Basidiospores occasionally with a small guttule, Q = 2.86–3.05, distribution in Australia F. eucalypticola
7 Pores 7–10 per mm; basidiospores 4.2–6.2 μm long, Q = 2.15–2.27 F. subferrea
Pores 5–7 per mm, basidiospores 5.5–7 μm long, Q > 2.27 8
8 Basidiospores 2.5–3 μm wide, Q = 2.29–2.44 F. resupinata
Basidiospores 2–2.5 μm wide, Q = 2.57–2.88 F. ramulicola

Acknowledgements

We are grateful to Prof. Yu-Cheng Dai (Beijing Forestry University) who allowed us to study his specimens. We would like to extend our thanks to the executive editor and anonymous reviewers for their suggestions and corrections to improve our work.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was financed by the National Natural Science Foundation of China (project no. 32100014), the Science and Technology Research Program of Chongqing Municipal Education Commission (grant no. KJQN202100737) and the Natural Science Foundation of Chongqing Science and Technology Bureau (grant no. CSTB2022NSCQ-MSX1345).

Author contributions

Data curation: HC. Investigation: CHL. Writing - original draft: QC. Writing - review and editing: XHL.

Author ORCIDs

Qian Chen https://orcid.org/0000-0001-7355-714X

Han Chen https://orcid.org/0009-0001-8424-9916

Cheng-Hang Luo https://orcid.org/0009-0005-6059-6373

Xiao-Hong Lai https://orcid.org/0000-0001-5393-4307

Data availability

All of the data that support the findings of this study are available in the main text.

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