Front. Cell. Infect. Microbiol. Frontiers in Cellular and Infection Microbiology Front. Cell. Infect. Microbiol. 2235-2988 Frontiers Media S.A. 10.3389/fcimb.2023.1205669 Cellular and Infection Microbiology Original Research Taxonomic and phylogenetic contributions to Fuscoporia (Hymenochaetales, Basidiomycota): two new species from Hawaii with a key to North American species Chen Qian 1 2 Liu Lu 1 Si Jing 3 * Vlasák Josef 4 * 1 College of Architecture and Urban Planning, Chongqing Jiaotong University, Chongqing, China 2 College of Architecture and Urban Planning, Tongji University, Shanghai, China 3 Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University, Beijing, China 4 Biology Centre, Institute of Plant Molecular Biology, Czech Academy of Sciences, Branišovská, České Budějovice, Czechia

Edited by: Tingting Dai, Nanjing Forestry University, China

Reviewed by: Sandrine Giraud, Université d’Angers, France; Haonan Wang, Zhejiang Agriculture and Forestry University, China

*Correspondence: Jing Si, jingsi1788@126.com; Josef Vlasák, vlasak@umbr.cas.cz

21 06 2023 2023 13 1205669 14 04 2023 23 05 2023 Copyright © 2023 Chen, Liu, Si and Vlasák 2023 Chen, Liu, Si and Vlasák

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Fuscoporia is a cosmopolitan, poroid, wood-decaying genus, belonging to the Hymenochaetales. During a study of wood-inhabiting fungi in the USA, four unknown specimens were collected from Hawaii. Both morphological criteria and molecular genetic analyses based on the ITS+nLSU+EF1-α datasets and the nLSU dataset confirmed that these four specimens represent two new species of Fuscoporia, and they are described as F. hawaiiana and F. minutissima. Fuscoporia hawaiiana is characterized by pileate basidiocarps, the absence of cystidioles, hooked hymenial setae, broadly ellipsoid to subglobose basidiospores measuring 4−6 × 3.5−4.5 μm. Fuscoporia minutissima is distinguished by small pores (10−13 per mm) and basidiospores (3.4−4 × 2.4−3 μm). The taxonomic status of the two new species is briefly discussed. A key to the North American species of Fuscoporia is provided.

 Hymenochaetaceae phylogeny polypore taxonomy new taxa 32100014, 32070016 National Natural Science Foundation of China10.13039/501100001809 section-in-acceptance Fungal Pathogenesis Introduction

Most wood-rotting fungi, belonging to Basidiomycetes, can use different types of wood as their nutritional source and are indispensable participants and important biological regeneration resources in natural ecosystems. The genus Fuscoporia Murrill (Hymenochaetales, Basidiomycota), erected by Murrill (1907) with F. ferruginosa (Schrad.) as generic type, is a cosmopolitan fungal group that encompasses also important species with medicinal effects, such as F. gilva (Schwein.) T. Wagner & M. Fisch. and F. torulosa (Pers.) T. Wagner & M. Fisch. (Wu et al., 2019). Fuscoporia is widely distributed in Asia, Europe, Oceania, and America. The genus has been considered a synonym of Poria Adans. or Phellinus Quél. for a long time (Overholts, 1953; Lowe, 1966; Ryvarden and Johansen, 1980; Gilbertson and Ryvarden, 1987; Larsen and Cobb-Poulle, 1990). However, Fiasson and Niemela (1984) recognized the genus as monophyletic morphologically, characterized by annual to perennial and resupinate to pileate basidiomata, a dimitic hyphal system with crystal encrustations on generative hyphae, the presence of hymenial setae, and hyaline, thin-walled and smooth basidiospores. Later, Wagner and Fischer, 2001; Wagner and Fischer, 2002) by means of nLSU sequence data and morphological studies of European Hymenochaetales reconfirmed the taxonomic status of the genus of Fuscoporia, with six species of Phellinus being classified into it.

Large number of Fuscoporia new species and new combinations have been discovered in recent years, and at present, more than 90 species are accepted in the genus (Chen et al., 2020; Tchoumi et al., 2020; Vlasák et al, 2020; Yuan et al., 2020; Dai et al., 2021; Chen et al., 2022; Hussain et al., 2022; Wu et al., 2022a, b). Fuscoporia is considered to be a complex genus with still unsettled taxonomy.

Striking diversity in the Hawaiian Islands with multiple co-occurring species leads to many species unreported in the literature. During a study of wood-inhabiting fungi, four unknown specimens were collected from Hawaii, with the features of Fuscoporia. To explore their relationships, phylogenetic analyses based on the ITS+nLSU+EF1-α datasets and the nLSU dataset were carried out. Both morphological criteria and molecular genetic analyses confirmed that these four specimens represent two new species of Fuscoporia. So, we describe them as F. hawaiiana and F. minutissima in the present paper. A key to the North American species of Fuscoporia is also provided.

 Materials and methods Morphological studies

Some studied specimen materials are deposited in the fungoria of the Institute of Microbiology, Beijing Forestry University (BJFC). The other ones are deposited in private fungoria of Josef Vlasak (JV) and then will be transferred into Prague Museum Herbarium (PRM). Morphological descriptions are based on field notes and herbarium specimens. The macroscopic color terms cited from Anonymous (1969) and Petersen (1996). Sections were studied at a magnification up to × 1,000 using a Nikon Eclipse 80i microscope with phase contrast illumination (Nikon, Tokyo, Japan). The microscopic analyses were performed accroding to Liu et al. (2022) and Si et al. (2023). Microscopic features, measurements, and drawings were prepared from slides stained with Cotton Blue. The following abbreviations are used: KOH = 5% potassium hydroxide, CB− = acyanophilous in Cotton Blue, IKI− = neither amyloid nor dextrinoid in Melzer’s reagent, L = arithmetic average of all spore length, W = arithmetic average of all spore width, Q = variation in the L/W ratios, and (n = x/y) = number of measured spores (x) measured from a given number of specimens (y).

 DNA extraction, PCR, and sequencing

A CTAB rapid plant genome extraction kit (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to extract total genomic DNA from dried specimens following the manufacturer’s instructions with some modifications (Chen et al., 2020; Zhang et al., 2023; Zhou et al., 2023). For generation of PCR amplicons, the following primer pairs were used: ITS5 (GGA AGT AAA AGT CGT AAC AAG G) and ITS4 (TCC TCC GCT TAT TGATAT GC) for internal transcribed spacer (ITS, White et al., 1990); LR0R (ACC CGC TGA ACT TAA GC) and LR7 (TAC TAC CAC CAA GAT CT) for nuclear large subunit rDNA (nLSU, Vilgalys and Hester, 1990); EF1-983F (GCY CCY GGH CAY CGT GAY TTY AT) and EF1-1567R (ACH GTR CCR ATA CCA CCR ATC TT) for translation elongation factor-1 alpha (EF1-α, Rehner and Buckley, 2005). The PCR procedures for ITS and EF1-α were as follows: initial denaturation at 95°C for 3 min, followed by 35 cycles of denaturation at 94°C for 40 s, annealing at 54°C for 45 s and extension at 72°C for 1 min, and a final extension at 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 of denaturation at 94°C for 1 min, annealing at 50°C for 1 min and extension at 72°C for 1.5 min, and a final extension at 72°C for 10 min. The PCR products were purified and sequenced at the Beijing Genomics Institute with the same primers and the sequences are deposited in GenBank. All newly generated sequences were deposited in GenBank (http://www.ncbi.nlm.nih.gov ) and are listed in Table 1 .

Species, specimens, and GenBank accession numbers of sequences used in ITS+nLSU+EF1-α phylogenetic analyses.

Species Specimen no. GenBank accession no. References
ITS nLSU EF1-α
Fuscoporia acutimarginata Dai 15137 MH050751 MH050765 MN848821 Chen and Dai, 2019
F. acutimarginata Dai 16892 MH050752 MH050766 MN848822 Chen and Dai, 2019
F. ambigua Cui 9244 MN816706 MN809995 MN848804 Du et al., 2020
F. ambigua JV 0509/151 MN816707 MN809996 − Du et al., 2020
F. americana JV 1209/3-J − MG008466 − Chen et al., 2019
F. americana JV 1209/100 KJ940022 MG008467 MH636384 Chen et al., 2019
F. atlantica SP 445618 KP058515 KP058517 − Pires et al., 2015
F. atlantica SP 465829 KP058514 KP058516 − Pires et al., 2015
F. australasica Dai 15625 MN816726 MN810018 MN848829 Chen et al., 2020
F. australasica Dai 15636 MG008397 MG008450 MH636408 Chen et al., 2019
F. australiana Dai 18672 MN816703 MN810014 MN848848 Chen et al., 2020
F. australiana Dai 18879 MN816705 MN810015 MN848850 Chen et al., 2020
F. bambusae Dai 16599 MN816711 MN809999 MN848808 Chen et al., 2020
F. bambusae Dai 16615 MN816715 MN810001 MN848810 Chen et al., 2020
F. callimorpha Dai 17388 MN121765 MN121824 − Chen and Dai, 2019
F. callimorpha Doll 868 MN816701 MN809992 MN848840 Chen et al., 2020
F. caymanensis JV 1908/74 MT676832 MT676833 − Vlasák et al., 2020
F. caymanensis JV 1408/5 MW009110 MW009109 − Vlasák et al., 2020
F. centroamericana JV 1607/93 MG008444 MG008460 MH636389 Chen et al., 2019
F. centroamericana O 908267 MG008443 − − Chen et al., 2019
F. chinensis Dai 15713 MN816721 MN810008 MN848846 Chen et al., 2020
F. chinensis Cui 11209 MN121767 MN121826 − Chen and Dai, 2019
F. chrysea JV 1607/106-J MN816736 MN810027 MN848818 Chen et al., 2020
F. cinchonensis Dai 19815 OP603023 OP600561 − Present study
F. contigua Dai 16025 MG008401 MG008454 MH636386 Chen et al., 2019
F. contigua Dai 13567A MG008402 MG008455 MN848817 Chen and Dai, 2019
F. costaricana JV 1407/92 MG008446 MG008461 MH636400 Chen et al., 2019
F. costaricana JV 1504/85 MG008413 MG478454 MH636401 Chen et al., 2019
F. dhofarensis ATN-007 OP593104 OP593105 OP597768 Hussain et al., 2022
F. eucalypti Dai 18783 MN816730 MN810021 MN848832 Chen et al., 2020
F. eucalypti Dai 18792 MN816731 MN810022 MN848831 Chen et al., 2020
F. ferrea MUCL 45984 KX961112 KY189112 MH636403 Chen and Yuan, 2017
F. ferrea Cui 11801 KX961101 KY189101 MN848823 Chen and Yuan, 2017
F. ferruginosa JV 0408/28 KX961103 KY189103 MH636397 Chen and Yuan, 2017
F. ferruginosa Dai 13200 MN816702 MN809993 MN848802 Chen et al., 2020
F. gilva JV 0709/75 USA MN816720 MN810007 MN848852 Chen et al., 2020
F. gilva JV 1209/65 MN816719 MN810006 MN848851 Chen et al., 2020
‘F. gilva’ URM 83957 MH392545 MH407344 − Yuan et al., 2020
‘F. gilva’ URM 91223 MH392550 MH407349 − Yuan et al., 2020
F. hainanensis Dai 16105 − ON520809 ON616518 Chen et al., 2022
F. hainanensis Dai 16110 − ON520810 ON616519 Chen et al., 2022
F. hawaiiana JV 2208/H22-J OQ817709 OQ817855 OQ849746 Present study
F. hawaiiana JV 2208/H30-J OQ817710 OQ817856 OQ849747 Present study
F. insolita Spirin 5251 KJ677113 − − Spirin et al., 2014
F. insolita Spirin 5208 MN816724 MN810016 MN848800 Chen et al., 2020
F. karsteniana Dai 16552 MN816716 MN810002 MN848806 Chen et al., 2020
F. karsteniana Dai 11403 MN816717 MN810003 MN848807 Chen et al., 2020
F. latispora JV 1109/48 MG008439 MG008468 MH636395 Chen et al., 2019
F. latispora JV 0610/VII-Kout MG008436 MG008469 MH636396 Chen et al., 2019
F. licnoides URM 84107 MH392556 MH407355 − Yuan et al., 2020
F. licnoides URM 83001 MH392561 MH407357 − Yuan et al., 2020
F. marquesiana URM83094 MH392544 MH407343 − Yuan et al., 2020
F. minutissima JV 2208/H12-J OQ817711 OQ817857 OQ849748 Present study
F. minutissima JV 2208/H16-J OQ817712 OQ817858 OQ849749 Present study
F. monticola Dai 10909 MG008410 − − Chen et al., 2019
F. monticola Dai 11860 MG008406 MG008457 MH636390 Chen et al., 2019
F. palomari JV 1004/5-J MN816737 − − Chen et al., 2020
F. palomari JV 1305/3-J MN816738 MN810028 MN848801 Chen et al., 2020
F. plumeriae Dai 17814 MN816714 MN810011 MN848845 Chen et al., 2020
F. plumeriae Dai 18858 MN816712 MN810010 MN848843 Chen et al., 2020
F. pulviniformis CMW 48060 MH599101 MH599125 MT108959 Tchoumi et al., 2020
F. pulviniformis CMW 48600 MH599102 MH599127 MT108960 Tchoumi et al., 2020
F. punctatiformis Dai 17443 MH050755 MH050764 − Chen and Dai, 2019
F. punctatiformis Doll#872a MH050753 − − Chen and Dai, 2019
F. ramulicola Dai 15723 MH050749 MH050762 MN848824 Chen and Dai, 2019
F. ramulicola Dai 16155 MH050750 MH050763 MN848825 Chen and Dai, 2019
F. roseocinerea JV 1407/84 MN816740 MN810030 MN848819 Chen et al., 2020
F. roseocinerea JV 1109/78-J MN816742 MN810032 MN848820 Chen et al., 2020
F. rufitincta JV 1008/25 KJ940029 KX058575 − Chen et al., 2020
F. rufitincta JV 0904/142 KJ940030 KX058574 − Chen et al., 2019
F. sarcites JV 0402/20K MZ264225 MZ264218 − Wu et al., 2022a
F. scruposa CMW 47749 MH599106 MH599129 MT108963 Yuan et al., 2020
F. scruposa CMW 48145 MH599105 MH599130 MT108962 Yuan et al., 2020
F. semiarida URM83800 MH392562 MH407361 − Yuan et al., 2020
F. semiarida URM82510 MH392563 MH407362 − Yuan et al., 2020
F. senex MEL 2382630 KP012992 KP012992 − Chen et al., 2020
F. senex KAUNP MK41 KP794600 − − Chen et al., 2020
F. septiseta Dai 12820 MG008405 MN810033 MH636394 Chen et al., 2019
F. septiseta JV 0509/78 MG008404 − − Chen et al., 2019
F. setifera Dai 15710 MH050758 MH050767 MN848841 Chen and Dai, 2019
F. setifera Dai 15706 MH050759 MH050769 MN848842 Chen and Dai, 2019
F. shoreae Dai 17806 MN816734 MN810025 MN848815 Chen et al., 2020
F. shoreae Dai 17818 MN816735 MN810026 MN848816 Chen et al., 2020
F. sinica Dai 15468 MG008412 MG008459 MH636392 Chen et al., 2019
F. sinica Dai 15489 MG008407 MG008458 MH636393 Chen et al., 2019
F. sinuosa Dai 20498 MZ264226 MZ264219 − Wu et al., 2022a
F. sinuosa Dai 20499 MZ264227 MZ264220 − Wu et al., 2022a
F. subchrysea Dai 16201 MN816708 MN809997 MN848811 Chen et al., 2020
F. subchrysea Dai 17656 MN816709 MN809998 MN848812 Chen et al., 2020
F. subferrea Dai 16326 KX961097 KY053472 MN848826 Chen and Yuan, 2017
F. subferrea Dai 16327 KX961098 KY053473 − Chen and Yuan, 2017
F. submurina Dai 19501 MZ264229 MZ264222 − Wu et al., 2022b
F. submurina Dai 19655 MZ264228 MZ264221 − Wu et al., 2022b
F. torulosa JV 1405/2 KX961106 KY189106 MH636405 Chen and Yuan, 2017
F. torulosa Dai 15518 MN816732 MN810023 MN848827 Chen et al., 2020
F. viticola JV 0911/6 KX961110 − − Chen and Yuan, 2017
F. viticola He 2123 MN816725 MN810017 − Chen et al., 2020
F. wahlbergii JV 1312/20-Kout MN816727 MG008462 − Chen et al., 2020
F. wahlbergii JV 0709/169-J MN816728 − − Chen et al., 2020
F. yunnanensis Cui 8182 MH050756 MN810029 − Chen and Dai, 2019
F. yunnanensis Dai 15637 MH050757 MH050768 − Chen and Dai, 2019
Outgroups
Coniferiporia weirii CFS 504 AY829341 AY829345 − Zhou et al., 2016
Phellinidium fragrans CBS 202.90 AY558619 AY059027 − Zhou et al., 2016

Newly generated sequences for this study and new species are in bold.

 Phylogenetic analysis

The following softwares were used for data processing and phylogenetic analyses: Mesquite, MAFFT 7.110, BioEdit 7.0.1 (Hall, 1999), ClustalX 2.0P10 (Thompson et al., 1997), MrModeltest 2.3 (Posada and Crandall, 1998; Nylander, 2004), PAUP* 4.0b10 (MP, Swofford, 2002), raxmlGUI 1.2 (ML, Stamatakis, 2006; Silvestro and Michalak, 2012), TreeView 1.5.0, and PowerPoint. Maximum parsimony (MP) and maximum likelihood (ML) methods were adopted to perform phylogenetic analyses of the two aligned datasets. The two phylogenetic methods resulted in similar topology for each data set. Thus, only the topology of the MP analysis appears along and branches that received bootstrap supports greater than or equal to 75% (MP and ML) were considered as significantly supported at the nodes.

To explore the phylogenetic position of Fuscoporia in Hymenochaetaceae, representatives of 28 genera of Hymenochaetaceae were included in nLSU dataset (not shown in Table 1 except for Fuscoporia; shown in Figure 1 ). Oxyporus populinus (Schumach.) Donk and Hyphodontia pallidula (Bres.) J. Erikss. were used as outgroups (Larsson et al., 2006; Zmitrovich and Malysheva, 2014; Zhou et al., 2016; Chen et al., 2019).

Phylogeny of Fuscoporia and related species generated inferred from nLSU sequences. Statistical values (MP/ML) are indicated for each node. Names of new species are in bold.

In addition to the newly generated sequences, additional ITS, nLSU, and EF1-α sequences of Fuscoporia based on formerly studies (Chen et al., 2019; Chen et al., 2020; Du et al., 2020; Wu et al., 2022a) were obtained from GenBank ( Table 1 ) to explore the phylogenetic position of our specimens in Fuscoporia. A total of 251 sequences, 100 samples of Fuscoporia were analyzed in the ITS+nLSU+EF1-α dataset ( Figure 2 ). Coniferiporia weirii (Murrill) L.W. Zhou et al (2016) and Phellinidium fragrans (M.J. Larsen & Lombard) Nuss (1986) were selected as outgroups (Zhou et al., 2016; Chen and Yuan, 2017).

Phylogeny of Fuscoporia and related species generated inferred from ITS+nLSU+EF1-α dataset. Statistical values (MP/ML) are indicated for each node. Names of new species are in bold.

 Results Phylogeny

To explore the phylogenetic position of Fuscoporia in Hymenochaetaceae, 81 taxa of the family Hymenochaetaceae were included in nLSU-alone dataset ( Figure 1 ). The dataset had an aligned length of 1436 characters, of which 1019 were constant, 98 variable but parsimony-uninformative, and 291 parsimony-informative. Maximum parsimony analysis yielded 140 equally topologies (TL = 1723, CI = 0.327, RI = 0.672, RC = 0.220, HI = 0.673). Fuscoporia is a powerfully supported lineage (100/100) within the Hymenochaetaceae family based on phylogenetic tree inferred from the nLSU dataset ( Figure 1 ).

The ITS+nLSU+EF1-α dataset ( Figure 2 ) included 98 ITS, 92 nLSU, and 62 EF1-α sequences from 100 fungal specimens representing 52 pecies of Fuscoporia. The dataset had an aligned length of 2705 characters, of which 1721 were constant, 99 variable but parsimony-uninformative, and 885 parsimony-informative. Maximum parsimony analysis yielded six equally topologies (TL = 4489, CI = 0.389, RI = 0.807, RC = 0.314, HI = 0.611). Two well-supported lineages (100/100) clustered within Fuscoporia and distincted from other species in phylogeny ( Figure 2 ). Taking morphological characters into consideration, these two new lineages represent two new species, F. hawaiiana and F. minutissima, which are described below.

 Taxonomy

Fuscoporia hawaiiana Q. Chen, Jing Si & Vlasák, sp. nov., Figures 3 , 4

Basidiocarps of Fuscoporia hawaiiana (holotype, JV 2208/H22-J). Bars: (A) = 1 cm. (B) = 1 mm.

Microscopic structures of Fuscoporia hawaiiana (holotype, JV 2208/H22-J). (A) Basidiospores. (B) Basidia and basidioles. (C) Hymenial setae. (D) Generative hyphae at dissepiment edge. (E) Hyphae from tube trama. (F) Hyphae from subiculum.

MycoBank no. — MB 847960

Holotype. — USA. Hawaii, Big Island, Kalopa State Park, on living tree of Eucalyptus, August 2022, JV 2208/H22-J (PRM, isotype JV, BJFC 039915).

Etymology. — Hawaiiana (Lat.): refers to the place (Hawaii) where the species was collected.

Fruiting body. — Basidiocarps perennial, pileate, laterally fused to imbricate, without odor or taste when fresh, hard corky when dry. Pilei mostly semicircular, projecting up to 11 cm long, 5 cm wide, and 5 cm thick at the base, more or less convex towards margin. Pileal surface reddish brown, concentrically sulcate with zones, glabrous, sometimes covered with mosses; margin obtuse, yellowish brown, up to 5 mm wide. Pore surface honey-yellow to deep olive, slight glancing; margin narrow, olivaceous buff, up to 1 mm wide; pores circular, 5−7 per mm; dissepiments thin to fairly thick, entire, abundant hymenial setae in tube cavities (under anatomical lens). Context clay-buff, hard corky, about 8 mm thick. Tubes olivaceous buff, hard corky, up to 2 cm long.

Hyphal structure. — Hyphal system dimitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH.

Context. — Generative hyphae rare, hyaline, thin- to slightly thick-walled, branched, frequently simple septate, 2−2.5 μm in diam; skeletal hyphae dominant, rust-brown, thick-walled with a medium to wide lumen, unbranched, occasionally septate, straight, more or less straight and regularly arranged, 2.5−3.5 μm in diam.

Tubes. — Generative hyphae rare, mostly present at dissepiment edges and subhymenium, hyaline, thin-walled, frequently branched and simple septate, 1.5−2.5 μm in diam, some of them encrusted at dissepiment edges and in the hymenium; skeletal hyphae dominant, yellowish brown, thick-walled with a medium to wide lumen, frequently septate, more or less straight, subparallel along the tubes, 2.5−3.5 μm in diam. Hymenial setae subulate, occasionally hooked, mostly originating from tramal hyphae, dark brown, thick-walled, 30−45 × 4−7 μm; basidia broadly clavate, with four sterigmata and a simple septum at the base, 12−18 × 5−7 μm; basidioles dominating the hymenium, in shape similar to basidia, but slightly smaller.

Spores. — Basidiospores broadly ellipsoid, hyaline, thin-walled, smooth, IKI−, CB−, some of them bearing a guttule, 4−6 × (3.4−)3.5−4.5(−4.7) μm, L = 4.77 μm, W = 3.90 μm, Q = 1.14−1.31 (n = 60/2).

Other material examined (paratype). — USA. Hawaii, Big Island, Volcano, on dead tree of Metrosideros polymorpha, August 2022, JV 2208/H30-J (JV, BJFC 039918).

Fuscoporia minutissima Q. Chen, Jing Si & Vlasák, sp. nov., Figures 5 , 6 MycoBank no. — MB 847964

Basidiocarps of Fuscoporia minutissima. (A–C)JV 2208/H16-J (holotype). (D) JV 2208/H12-J. Bars: (A, B, D) = 1 cm. (C) = 1 mm.

Microscopic structures of Fuscoporia minutissima (holotype, JV 2208/H16-J). (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.

Holotype. — USA. Hawaii, Big Island, Makuala O’oma Trail, August 2022, JV 2208/H16-J (PRM, isotype JV, BJFC 039911).

Etymology. — Minutissima (Lat.): refers to the very small size of the pores exhibiting by these species.

Fruiting body. — Basidiocarps perennial, pileate, imbricate, without odor or taste when fresh, hard corky when dry. Pilei mostly imbricate, projecting up to 5 cm long, 3 cm wide, and 1 cm thick at the base. Pileal surface reddish brown, concentrically sulcate with zones, nodulose; margin obtuse to slightly acute, honey-yellow, up to 1 mm wide. Pore surface greyish brown to deep olive, glancing; margin narrow, honey-yellow, up to 1 mm wide; pores circular, 10−13 per mm; dissepiments fairly thick, entire and matted, abundant hymenial setae in tube cavities (under anatomical lens). Context honey-yellow, hard corky, up to 5 mm thick. Tubes olivaceous buff, paler than pores, hard corky, up to 4 mm long.

Hyphal structure. — Hyphal system dimitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH.

Context. — Generative hyphae rare, hyaline, thin- to slightly thick-walled, unbranched, frequently simple septate, 2−3 μm in diam; skeletal hyphae dominant, rust-brown, thick-walled with a medium to narrow lumen, unbranched, occasionally septate, straight, regularly arranged, 3−4 μm in diam.

Tubes. — Generative hyphae rare, mostly present at dissepiment edges and subhymenium, hyaline, thin-walled, frequently branched and simple septate, 2−3 μm in diam, some of them encrusted at dissepiment edges and in the hymenium; skeletal hyphae dominant, yellowish brown, thick-walled with a medium to narrow lumen, frequently septate, more or less straight, subparallel along the tubes, 3−4 μm in diam. Hymenial setae subulate, mostly originating from tramal hyphae, dark brown, thick-walled, 18−40 × 6−9 μm; fusoid cystidioles hyaline and thin-walled, 9.5−12 × 4−5.5 μm; basidia short clavate to barrel-shaped, with four sterigmata and a simple septum at the base, 10−15 × 4−6.5 μm; basidioles dominating the hymenium, in shape similar to basidia, but slightly smaller.

Spores. — Basidiospores broadly ellipsoid to subglobose, hyaline, thin-walled, smooth, IKI−, CB−, bearing a guttule, (3−)3.4−4 × (2.2−)2.4−3(−3.8) μm, L = 3.60 μm, W = 2.79 μm, Q = 1.24−1.30 (n = 52/2).

Other material examined (paratype). — USA. Hawaii, Kauai Island, Koke’e State Park, on dead tree of Acacia koa, August 2022, JV 2208/H12-J (JV, BJFC 039910).

 Discussion

The islands far from the mainland have attracted scientific researchers for a long time, due to the close combination of ecological environment and evolution process here that deepen our understanding of the formation process of biological diversity (Cotoras et al., 2018). Hawaii is a biological hotspot with a variety of climates and habitats. 160 genera and 400 species of wood-rotting basidiomycetes are reported from 110 native and exotic substrate species on the Hawaiian Islands (Gilbertson et al., 2002; Ashiglar et al., 2015). There are few natural forests on the Hawaii islands (Chambers et al., 2007). Most forests are filled with introduced trees, and our new species may be introduced as well. The two new wood-rotting fungal species Fuscoporia hawaiiana and F. minutissima were collected from the northernmost and oldest Kauai Island, and the largest island Big Island in the Hawaiian Islands, both with a tropical sea climate.

Fuscoporia hawaiiana is characterized by perennial and pileate basidiocarps, circular and medium pores (5−7 per mm), the absence of cystidioles, hooked hymenial setae, broadly ellipsoid to subglobose basidiospores measuring 4−6 × 3.5−4.5 μm. Fuscoporia hawaiiana may also be distributed in Yunnan Province, China, considered to be F. torulosa derived from CLZhao 10146 (OM959398) on GenBank. The ITS of CLZhao 10146 shows that only 2 base differences from our new samples, but unfortunately, it lacks nLSU or EF1-α sequence. Furthermore, F. torulosa can be easily distinguished from F. hawaiiana by its subungulate basidiocarps and straight hymenial setae. Morphologically, seven species, F. semihispida (Ryvarden) Y.C. Dai & F. Wu, F. australasica Q. Chen, F. Wu & Y.C. Dai, F. marquesiana Gibertoni & C.R.S. de Lira, F. atlantica Motato-Vásq., R.M. Pires & Gugliotta, F. wahlbergii (Fr.) T. Wagner & M. Fisch., F. eucalypti Q. Chen, F. Wu & Y.C. Dai, and F. rufa (Bres.) Y.C. Dai & F. Wu., are similar to F. hawaiiana by sharing similar pileate basidiocarps and hooked hymenial setae. However, F. hawaiiana is distant from F. atlantica, F. marquesiana, F. wahlbergii, F. eucalypti, and F. australasica both in the phylogenetic analyses ( Figures 1 , 2 ) and morphology. F. atlantica differs from F. hawaiiana by its annual basidiocarps (Pires et al., 2015); F. australasica is described from Southern China and Viet Nam and differs from F. hawaiiana by its glabrous basidiocarps and the presence of cystidioles (Chen et al., 2020); F. wahlbergii is described from Europe, East-African, Australia, and USA, and distinguishes from F. hawaiiana by its smaller pores (7−9 per mm) and the presence of cystidioles (Chen et al., 2020); F. eucalypti is distributed Australia, and differs from F. hawaiiana by its spores effused-reflexed to pileate and wider (4.3−5.5 × 4−4.5 μm) (Chen et al., 2020); F. marquesiana is a South American species, and can be easily distinguished from F. hawaiiana by its smaller pores (8−9 per mm) (Wu et al., 2022b). Even without molecular data, F. semihispida and F. rufa can be easily distinguished from F. hawaiiana by narrower or wider spores (2−2.5 μm in F. semihispida; 4−4.5 μm in F. rufa) (Wu et al., 2022a).

Fuscoporia minutissima stands out in the genus by its perennial and pileate basidiocarps, and very small pores (10−13 per mm) and spores (3.4−4 × 2.4−3 μm). Fuscoporia minutissima is usually sterile after collecting, and such specimens are very similar to F. gilva by pileate basidiocarps, glabrous to rugose sometimes with nodulose pileal surface and broadly ellipsoid to subglobose spores, but the later differs by having bigger pores (6−8 per mm). Fuscoporia plumeriae Q. Chen, F. Wu & Y.C. Dai and F. chinensis Q. Chen, F. Wu & Y.C. Dai clustered together with F. minutissima into a group with a strong support (100/100, Figures 1 , 2 ). Morphologically, these species share the effused-reflexed to pileate basidiocarps with lacerate dissepiments and ellipsoid basidiospores; but F. plumeriae and F. chinensis have annual basidiocarps and bigger pores (10−13 per mm in F. minutissima; 8−10 per mm in F. plumeriae; 7−8 per mm in F. chinensis; Chen et al., 2020). Fuscoporia rhabarbarina (Berk.) Groposo, Log.-Leite & Góes-Neto is a common fungus on hardwoods along streams in South America and subtropical and tropical Asia. But the type locality of this species remains unknown, probably in Brazil (Wu et al., 2022b). Fuscoporia minutissima may be confused with F. rhabarbarina in field, but the pileal surface of F. rhabarbarina is glabrous and dark brown crust at the base. Moreover, the two species are not closely related in the phylogenetic tree ( Figure 1 ).

Fuscoporia hawaiiana and F. minutissima are described in this study. Traditionally, they were most probably treated as F. wahlbergii and F. gilva, respectively. The phylogenetic analyses provide molecular evidence to support these two new species derive from the F. wahlbergii complex and the F. gilva complex ( Figures 1 , 2 ). The similar results are demonstrated in many polypores complex, for instances, Megasporoporia setulosa (Henn.) Rajchenb. (Wang et al., 2021; Wang et al., 2022), Heterobasidion annosum (Fr.) Bref. (Yuan et al., 2021), Porogramme epimiltina (Berk. & Broome) Y.C. Dai (Mao et al., 2023), Sidera vulgaris (Fr.) Miettinen (Liu et al., 2021), and Phaeolus schweinitzii (Fr.) Pat. (Yuan et al., 2022). It seems that the diversity of wood-inhabiting fungi is extremely rich in tropics, some traditional definition on tropical species should be re-evaluated, and the concepts of taxa in tropics should be modified after molecular phylogeny.

A key to North American species of Fuscoporia.

1. Basidiocarps completely resupinate...........................................1

- Basidiocarps pileate, effused-reflexed or substipitate................7

2. Mycelial setae absent .........................F. ferrea (Pers.) G. Cunn.

- Mycelial setae present.....................................................................3

3. Pores 5−9 per mm..........................................................................4

- Pores 2−4 per mm...........................................................................5

4. Pores 7−9 per mm, skeletal hyphae septate ................ ............................................................................................................ ......F. rufitincta (Berk. & M.A. Curtis ex A.L. Sm.) Murrill

- Pores 5−6 per mm, skeletal hyphae aseptate F. ambigua P. Du, Q. Chen & J. Vlasák

5. Basidiocarps perennial ..........................F. contigua (Pers.) G. Cunn.

- Basidiocarps annual.................................................................................6

6. Spores broadly ellipsoid, 4.8−6.0 × 3.2−4.2 μm ....................................F. americana Y.C. Dai, Q. Chen & J. Vlasák

- Spores cylindric, 6.0−7.0 × 2.0−3.0 μm ............................F. septiseta Y.C. Dai, Q. Chen & J. Vlasák

7. Hymenial setae hooked ........................................................................8

- Hymenial setae straight...........................................................................9

8. Pores 7−9 per mm, cystidioles present..........................F. wahlbergii

- Pores 5−7 per mm, cystidioles absent.............................F. hawaiiana

9. Basidiocarps annual.........................F. palomari Vlasák & Ryvarden

- Basidiocarps perennial..........................................................................10

10. Spores cylindric, 7−9 × 1.5−2 μm, Q > 2...........................F. viticola (Schwein.) Murrill

- Spores broadly ellipsoid to ellipsoid, Q < 2.......................................11

11. Pileal surface velutinate to glabrous.......................F. rhabarbarina

- Pileal surface tomentose, hispid, rugose............................................12

12. Pores 10−13 per mm...................................................F. minutissima

- Pores 6−9 per mm..................................................................................13

13. Pileal surface not sulcate...........................................................F. gilva

- Pileal surface concentrically sulcate....................................................14

14. Basidiocarps subungulate, pileal surface grayish brown.....................................................................................F. torulosa

- Basidiocarps usually applanate, pileal surface fuscous to black........................................F. senex (Nees & Mont.) Ghob.-Nejh

 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: F. hawaiiana JV 2208/H22-J: OQ817709; OQ817855; OQ849746. F. hawaiiana JV 2208/H30-J: OQ817710; OQ817856; OQ849747. F. minutissima JV 2208/H12-J: OQ817711; OQ817857; OQ849748. F. minutissima JV 2208/H16-J: OQ817712; OQ817858; OQ849749. F. cinchonensis OP603023; OP600561.

 Author contributions

QC, LL, JS, and JV designed the research and contributed to data analysis and interpretation. JV prepared the samples. QC and LL conducted the molecular experiments and analyzed the data. QC prepared the samples and drafted the manuscript. JV and JS discussed the results and edited the manuscript. All authors contributed to the article and approved the submitted version.

 Funding

This study was financed by the National Natural Science Foundation of China (Nos. 32100014 and 32070016), the Science and Technology Research Program of Chongqing Municipal Education Commission (No. KJQN202100737), and the Academy of Sciences of the Czech Republic RVO (No. 60077344).

 Acknowledgments

The authors would like to express their deep appreciations to Prof. Yu-Cheng Dai (Beijing Forestry University, China) for forwarding specimens on loan and Dr. Josef Vlasák Jr. for collecting fresh specimens on Hawaii islands.

 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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