11 urn:lsid:arphahub.com:pub:C004A564-9D6A-5F9F-B058-6A3815DFE9C3 urn:lsid:zoobank.org:pub:A4FD9303-3F01-45D9-B438-FCCFE465D2F7 MycoKeys MC 1314-4057 1314-4049 Pensoft Publishers 10.3897/mycokeys.125.168173 168173 Research Article Agaricomycetes Biodiversity & Conservation Bioinformatics DNA barcoding Molecular systematics Phylogeny Taxonomy Asia Three new species of Fuscoporia (Hymenochaetales, Basidiomycota) from southern India revealed by morphological and multigene phylogenetic analyses Gunaseelan Sugantha https://orcid.org/0000-0001-7089-2292 1 Kezo Kezhocuyi https://orcid.org/0000-0002-3723-0462 1 Arumugam Elangovan https://orcid.org/0009-0003-9720-2978 1 Karunarathna Samantha C. https://orcid.org/0000-0001-7080-0781 2 Fahad Al-Arjani Al-Bandari 3 Elgorban Abdallah M. https://orcid.org/0000-0003-3664-7853 3 Kumla Jaturong https://orcid.org/0000-0002-3673-6541 4 Suwannarach Nakarin https://orcid.org/0000-0002-2653-1913 4 Chukeatirote Ekachai ekachai@mfu.ac.th https://orcid.org/0000-0002-9968-5841 5 Kaliyaperumal Malarvizhi malar.kaliyaperumal@gmail.com https://orcid.org/0000-0002-1218-3778 1 Centre for Advanced Studies in Botany, University of Madras, Guindy Campus, Chennai 600025, Tamil Nadu, India Center for Yunnan Plateau Biological Resources Protection and Utilization & Yunnan International Joint Laboratory of Fungal Sustainable Utilization in South and Southeast Asia, College of Biology and Food Engineering, Qujing Normal University, Qujing 655099, China Botany and Microbiology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia Center of Excellence in Biotechnology Research (CEBR), DSR, King Saud University, Riyadh, Saudi Arabia Center of Excellence in Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai 50200, Thailand School of Science, Mae Fah Luang University, Chiang Rai, 57100, Thailand

Corresponding authors: Ekachai Chukeatirote (ekachai@mfu.ac.th); Malarvizhi Kaliyaperumal (malar.kaliyaperumal@gmail.com)

Academic editor: R. Henrik Nilsson

 2025 18 11 2025 125 147 166 BE3C9D2E-4CB6-5BCF-9203-5C38C3451454 08 08 2025 20 10 2025 Sugantha Gunaseelan, Kezhocuyi Kezo, Elangovan Arumugam, Samantha C. Karunarathna, Al-Bandari Fahad Al-Arjani, Abdallah M. Elgorban, Jaturong Kumla, Nakarin Suwannarach, Ekachai Chukeatirote, Malarvizhi Kaliyaperumal This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Fuscoporia Murrill, a cosmopolitan genus of the Hymenochaetaceae, consists of parasitic and saprotrophic fungi characterized by resupinate to pileate, strictly dimitic hyphal systems, encrusted generative hyphae, the presence of hymenial setae, and hyaline, smooth, thin-walled basidiospores. Based on morpho-microtaxonomic examinations and phylogenetic analyses using a combined ITS, nrLSU, partial rpb2, and tef1-α dataset, three new species of Fuscoporia—F. indica, F. sirumalaiensis, and F. terminalianae—are described from the Eastern Ghats of Tamil Nadu, India. The newly described species form three distinct lineages within the F. torulosa complex. Fuscoporia indica is characterized by imbricate, convex, dimidiate basidiomes, a glabrous, azonate pilear surface, and ellipsoid basidiospores (3.5–4.5 × 2.5–2.9 μm). Fuscoporia sirumalaiensis has a smooth to glabrous, concentrically zonate pilear surface, a duplex context, and smaller basidiospores (3.1–3.6 × 2.1–2.6 μm). Finally, F. terminalianae is recognized by its effused-reflexed to imbricate basidiome, widely zonate and warted pilear surface, and ellipsoidal basidiospores (3.3–4.3 × 2.8–3 μm). This study provides comprehensive descriptions, morphological illustrations, and insights into the differences among these new species and their respective allied taxa, along with the results of phylogenetic analysis.

 Hymenochaetaceae multigene phylogeny Phellinus sensu lato taxonomy wood-rotting fungi EMR-SERB, DST (EMR/2016/003078), Government of India Tamil Nadu State Council for Higher Education (RGP/2019-20/MU/HECP-0040) The National Natural Science Foundation of China (Number 32260004) The High-Level Talent Recruitment Plan of Yunnan Province (High-End Foreign Experts program) The Ongoing Research Funding Program (ORF-2025-558), King Saud University, Riyadh, Saudi Arabia Citation

Gunaseelan S, Kezo K, Arumugam E, Karunarathna SC, Fahad Al-Arjani A-B, Elgorban AM, Kumla J, Suwannarach N, Chukeatirote E, Kaliyaperumal M (2025) Three new species of Fuscoporia (Hymenochaetales, Basidiomycota) from southern India revealed by morphological and multigene phylogenetic analyses. MycoKeys 125: 147–166. https://doi.org/10.3897/mycokeys.125.168173

 Introduction

The genus Fuscoporia Murrill was erected with F. ferruginosa (Schrad.) Murrill as the type species (Murrill 1907). It is widely recognized as a forest pathogen that causes white rot in both coniferous and deciduous trees (Panconesi et al. 1994; Spirin et al. 2014; Luana et al. 2015). Fuscoporia is characterized by its annual to perennial, resupinate to pileate basidiomes, a dimitic hyphal system with encrusted generative hyphae, and the presence of hymenial setae in most species except for F. shoreae, F. longisetulosa, and F. discipes. Marginal setae may be present or absent, and the basidiospores are hyaline, thin-walled, smooth, and range in shape from subglobose to cylindrical (Dai 2010; Wu et al. 2022). Fuscoporia is distributed worldwide; more than 1,561 nucleotide sequences belonging to 63 known and 55 unspecified species are publicly available in GenBank (Sayers et al. 2025), and 130 taxa are registered in MycoBank (Robert et al. 2013) as of October 2025.

Phylogenetic studies of North American and European members of the Hymenochaetaceae based on nrLSU redefined Phellinus sensu lato into several monophyletic genera. These include Fomitiporia Murrill, Fomitiporella Murrill, Fulvifomes Murrill, Fuscoporia Murrill, Phellinus s. str., Porodaedalea Murrill, Phellinidium (Kotl.) Fiasson and Niemelä, Phellopilus Niemelä, T. Wagner and M. Fisch., Aurificaria D.A. Reid, Onnia P. Karst., and Phylloporia Murrill (Fiasson and Niemelä 1984; Fischer 1996; Niemelä et al. 2001; Wagner and Fischer 2001, 2002), all of which were later accepted as distinct taxa (Niemelä et al. 2001; Spirin et al. 2006; Groposo et al. 2007; Baltazar et al. 2009; Baltazar and Gibertoni 2010; Dai 2010; Raymundo et al. 2013; Chen and Yuan 2017; Chen et al. 2019; Chen and Dai 2019). Recent molecular analyses of Fuscoporia have revealed that within this taxon, there are several major groupings supported by substantial morphological and phylogenetic evidence. These are the F. contigua complex, the F. ferra complex, the F. ferruginosa complex, the F. gilva complex, the F. torulosa complex, and the F. viticola complex (Chen et al. 2022, 2024; Wu et al. 2022; Cho et al. 2023).

Chen et al. (2019, 2020, 2023a, b) proposed 18 new species—F. acutimarginata, F. australasica, F. australiana, F. americana, F. bambusae, F. centroamericana, F. chinensis, F. costaricana, F. eucalypti, F. karsteniana, F. latispora, F. monticola, F. plumeriae, F. ramulicola, F. septiseta, F. sinica, F. shoreae, and F. subchrysea—from Australia, China, Costa Rica, Mexico, Singapore, Thailand, and the United States. From 2022 to 2024, numerous new species—F. dolichoseta, F. dollingeri, F. eucalypticola, F. gilvoides, F. hawaiiana, F. koreana, F. minutissima, F. naditirana, F. resupinate, F. reticulata, F. semicephala, F. sinuosa, F. submurina, and F. subtropica—have been reported worldwide (Wu et al. 2022; Chen et al. 2023a, b; Cho et al. 2023; Chen et al. 2024; Crous et al. 2024). Recently, Chen et al. (2025) reported Fuscoporia reflexoides, a new species belonging to the F. gilva complex from China.

Despite significant advancements, several regions of the world remain under-sampled, including Oceania, parts of South America, tropical Africa, and Southeast Asian islands. For a more comprehensive understanding of Fuscoporia, widespread sampling from Neotropical, Paleotropical, tropical, and temperate countries of Asia is essential. Earlier, knowledge of the species diversity of hymenochaetoid fungi in India was confined to the northern regions, primarily based on morpho-taxonomic characteristics. Kumari et al. (2021) documented 14 species of Phellinus (P. callimorphus, P. chryseus, P. contiguus, P. discipes, P. ferreus, P. ferruginosus, P. gilvus, P. orientalis, P. punctatiformis, P. rhabarbarinus, P. rufitinctus, P. senex, P. torulosus, and P. wahlbergii), which have since been validated and transferred to Fuscoporia through phylogenetic analysis. From the southern region of India, Bakshi (1971) reported P. discipes (syn. F. discipes). Our previous study represented the first multigene approach in revealing F. naditirana from the Eastern Ghats of Tamil Nadu (Crous et al. 2024). In the present study, we report three novel species of Fuscoporia from the Eastern Ghats of Tamil Nadu and provide detailed descriptions, illustrations, and phylogenetic placement based on ITS+nrLSU+rpb2+tef1-α sequence data.

 Materials and methods Morphological analyses

The studied specimens were collected from various locations in the Eastern Ghats of Tamil Nadu, India. Morphological and microscopic characteristics were analyzed as described earlier (Gunaseelan et al. 2024), and the phenetic color codes follow the Methuen Handbook (Kornerup and Wanscher 1978). Freehand sections of dried basidiomes were mounted in water, 5% potassium hydroxide (KOH), cotton blue (CB), and Melzer’s reagent (IKI) to analyze micro-morphological features. Microscopic observations, measurements, and line drawings were performed in 5% KOH using a LABOMED CxL2 compound microscope. Photomicrographs were taken with a LABOMED OPTIC-CX BINO LED microscope at magnifications up to 1000×. Basidiospore measurements (minimum–mean–maximum) and Q values (length/width ratios) were recorded. Abbreviations: CB− = acyanophilous, CB+ = cyanophilous, IKI− = inamyloid, IKI+ = amyloid, Q = L/W ratio (basidium length excluding sterigmata), L = mean spore length (arithmetic mean of all spores), W = mean spore width (arithmetic mean of all spores), n = number of spores measured. Measurements were based on 50 basidiospores, 30 cystidioles, hymenial setae, basidioles, and basidia per specimen. The identified specimens were deposited at the Madras University Botany Laboratory (MUBL), Centre for Advanced Studies in Botany, University of Madras, Chennai 600025, Tamil Nadu, India.

 Genomic DNA extraction, PCR amplification, and sequencing

The nuclear ribosomal internal transcribed spacer (ITS) region was amplified using primers ITS1 and ITS4 (White et al. 1990). PCR amplification was carried out under the following conditions: initial denaturation at 95 °C for 3 minutes, followed by 32 cycles of 95 °C for 30 seconds, 52 °C for 30 seconds, and 68 °C for 1 minute, with a final extension at 68 °C for 3 minutes. The nuclear ribosomal large subunit (nrLSU) region was amplified using primers LR0R and LR7 (Vilgalys and Hester 1990). PCR amplification conditions for nrLSU were as follows: initial denaturation at 94 °C for 1 minute, followed by 34 cycles at 94 °C for 30 seconds, 45 °C for 30 seconds, and 72 °C for 1.5 minutes, and a final extension at 72 °C for 10 minutes. The RNA polymerase II gene (rpb2) was amplified with primers bRPB2-6F and bRPB2-7.1R (Matheny 2005), and PCR conditions were as follows: initial denaturation at 94 °C for 2 minutes; 35 cycles of 94 °C for 45 seconds, 53 °C for 90 seconds, and 72 °C for 90 seconds; and a final extension at 72 °C for 10 minutes. The translation elongation factor 1-alpha gene (tef1-α) was amplified with primer pair EF1-983F/EF1-1567R (Rehner and Buckley 2005). PCR amplification conditions for tef1-α were as follows: initial denaturation at 94 °C for 3 minutes, followed by 34 cycles of denaturation at 94 °C for 40 seconds, annealing at 54 °C for 45 seconds, extension at 72 °C for 1 minute, and a final extension at 72 °C for 10 minutes. Sequencing was outsourced to Eurofins Genomics India Pvt. Ltd., Karnataka, India.

 Phylogenetic analyses

For the phylogenetic analyses, concatenated sequences (ITS+nrLSU+rpb2+tef1-α) generated from this study and allied taxa retrieved from GenBank (Table 1), along with the outgroups Coniferiporia weirii (CFS 504) and Phellinidium fragrans (CBS 202.90), were aligned in MEGA X v10.0.2, followed by manual adjustments to enhance alignment accuracy (Kumar et al. 2018). To facilitate dataset partitioning, the most appropriate partitioning scheme and substitution models were determined using PartitionFinder v1.10 (Lanfear et al. 2012) under the AIC criterion with the “greedy” search option. The dataset was subdivided into 10 partitions: ITS1, 5.8S, ITS2, nrLSU, rpb2 introns, rpb2 exon, tef1-α introns, tef1-α 1st codon, tef1-α 2nd codon, and tef1-α 3rd codon. The best-fit evolutionary models selected by MrModeltest v2.3 (Nylander 2004) were implemented for each partition: GTR+G (ITS1), GTR+I+G (5.8S), JC69 (ITS2), HKY85 (nrLSU), GTR+G (rpb2 introns), GTR+G (rpb2 exon), HKY85+I+G (tef1-α introns), GTR+G (tef1-α 1st codon), HKY85+G (tef1-α 2nd codon), and K80+G (tef1-α 3rd codon). The combined dataset with partition-specific models was used for Bayesian analysis using MrBayes version 3.2.7a (Ronquist et al. 2012) with two independent runs comprising four Metropolis-coupled Markov chain Monte Carlo chains, each running for 4,000,000 generations with sampling every 1,000 generations. The first 25% of sampled trees were discarded as burn-in, and the remaining trees were used to infer a majority-rule consensus and calculate Bayesian posterior probabilities (BPP) for the clades. Maximum likelihood (ML) analyses were performed using raxmlGUI 2.0 (Edler et al. 2020), involving 1,000 replicates under the GTR+GAMMAI model. Additionally, 1,000 rapid bootstrap replicates were performed using the GTRCAT model to evaluate the ML bootstrap values of the nodes. A clade was considered strongly supported if bootstrap (BS) ≥ 60% and posterior probability (PP) ≥ 0.80. Newly generated sequences were deposited in GenBank (http://www.ncbi.nlm.nih.gov/) (Table 1). The alignment was submitted to Figshare (DOI: 10.6084/m9.figshare.29844794).

Names, strain numbers, countries of collection, and corresponding GenBank accession numbers of the sequences used in this study. Newly generated sequences are in bold, ‘T’ indicates type specimens, and em dashes (—) indicate data unavailability.

Species Strain Country ITS LSU tef1-α rpb2
Fuscoporia acutimarginata T Dai 16892 China MH050752 MH050766 MN848822 MH079393
F. ambigua T JV 0509/151 United States MN816707 MN809996 — MN848792
F. americana T BJFC 020644 United States NR_173723 — — —
F. atlantica VRTO24 Brazil ON808610 ON795835 — —
F. australasica T Dai 15636 China MG008397 MG008450 MH636408 MH079402
F. australiana T Dai 18879 Australia MN816705 MN810015 MN848850 MN848767
F. bambusae T Dai 16599 Thailand MN816711 NG_075315 MN848808 —
F. bambusicola T Cui 8692 China MN816739 MT032486 MN848813 —
F. callimorpha Doll 868 China MN816701 MN809992 MN848840 —
F. caymanensis T JV 1908/74 French Guiana MT676832 MT676833 — —
F. centroamericana JV 1606/93 Costa Rica MG008444 MG008460 MH636389 —
F. chinensis T Dai 15713 China MN816721 MN810008 MN848846 MN848771
F. chrysea JV 1607/106-J Costa Rica MN816736 MN810027 MN848818 MN848773
F. cinchonensis CBS 447.76 South Korea AY558613 — — —
F. contigua T Dai 16025 United States MG008401 MG008454 MH636386 MH079406
F. costaricana JV 1407/92 Costa Rica MG008446 MG008461 — —
F. dhofarensis ATN-007 Oman OP593104 OP593105 — —
F. dolichoseta T SFC20191015-23 Republic of Korea ON427765 ON427795 — ON464731
F. dollingeri T Doll623 United States MW908540 MW898444 — —
F. eucalypti T Dai 18792 Australia MN816731 MN810022 MN848831 MN848778
F. eucalypticola T Dai 18593A Australia PP732562 PP732631 — —
F. ferrea JV 1606/2.2-J United States KX961100 KY189100 MH636402 MH079394
F. ferruginosa Cui 9244 China MN816706 MN809995 MN848804 MN848791
F. formasona T VRTOBFO3 Brazil ON808603 ON795827 — —
F. formasona VRTO83 Brazil — ON795830 — —
F. gilva JV 1209/65 United States MN816719 MN810006 MN848851 MN848768
F. gilvoides T SFC20180426-12 Republic of Korea ON427763 ON427793 ON479802 ON464729
F. hawaiiana JV 2208/H22-J United States OQ817709 OQ817855 OQ849746 —
F. indica T MUBL1104 India — PP390498 PV638743 PV638735
F. indica HRS-15B India — PQ113747 PV638744 PV638736
F. insolita T JV1208/5208 Russia MN816724 MN810016 MN848800 —
F. karsteniana T Dai 11403 China MN816717 MN810003 MN848795 MN848807
F. koreana T SFC20160726-93 Republic of Korea ON427762 ON427792 ON479801 ON464728
F. latispora JV 1109/482 United States MG008439 MG008468 MH636395 MN848799
F. licnoides URM 83001 Brazil MH392561 MH407357 — —
F. minutissima T JV 2208/H16-J United States OQ817711 OQ817857 OQ849748 —
F. montana 175856 Taiwan JX484015 JX484007 — —
F. monticola T Dai 11860 China MG008406 MG008457 MH636390 —
F. marquesiana URM83094 Brazil MH392544 MH407343 — —
F. nadiatariana MUBL1105 India PQ098039 PP390499 PQ346367 PQ346369
F. nadiatariana SP2F2A India PQ098040 PQ113748 PQ346368 PQ346370
F. palomari JV 1305/3-J United States MN816738 MN810028 — —
F. plumeriae T Dai 18858 Australia MN816712 MN810010 MN848843 MN848769
F. pulviniformis T CMW48600 South Africa MH599102 MH599127 MT108960 —
F. punctatiformis Dai 17443 China MH050755 MH050764 — —
F. ramulicola T Dai 15723 China MH050749 MH050762 MN848824 MH079398
F. resupinata Dai 20455 China PP732567 PP732636 — —
F. reticulata T SFC20160115-16 Republic of Korea ON427761 ON427791 ON479800 ON464727
F. rhabarbarina Dai 16226 China MN816743 MN810035 MN848838 MN848784
F. roseocinerea JV 1407/84 China MN816740 MN810030 MN848819 —
F. rufitincta JV 1008/25 United States KJ940029 KX058575 — —
F. sarcites JV 0402/20K Venezuela MZ264225 MZ264218 — —
F. semiarida URM 82510 Brazil MH392563 MH407362 — —
F. scruposa VRTOV473 Brazil ON795836 — —
F. senex Dai 15775 China MN816746 MN810038 — —
F. septiseta T Dai 12820 United States MG008405 MN810033 MH636394 —
F. setifera Dai 15706 China MH050759 MH050769 MN848842 MN159391
F. shoreae T Dai 17818 Singapore MN816735 MN810026 MN848816 —
F. sinica T Dai 15468 China MG008412 MG008459 MH636392 —
F. sinuosa Dai 20499 China MZ264227 MZ264220 — —
F. sirumalaiensis T MUBL1106 India PQ098037 PP390500 PV638741 PV638737
F. sirumalaiensis SRM21 India PQ098038 PQ113746 PV638742 PV638738
F. subchrysea T Dai 16201 China MN816708 MN809997 MN848811 MN848796
F. subferrea T Dai 16327 China KX961098 KY053473 — MH079401
F. submurina T Dai 19655 China MZ264228 MZ264221 — —
F. subtropica T Dai 19957 China PP732565 PP732634 — —
F. terminalianae T MUBL1107 India PQ098033 PP390501 PV638739 PV638733
F. terminalianae VM2B India PQ098034 PQ113745 PV638740 PV638734
F. torulosa JV 1405/2 Czech Republic KX961106 KY189106 MH636405 MN159392
F. viticola He 2081 United States MN121829 MN121770 — —
F. wahlbergii JV 1312/20-Kout Spain MN81672 MG008462 — —
F. yunnanensis Cui 8182 China MH050756 MN810029 — MN848789
Outgroup
Coniferiporia weirii CFS 504 Canada AY829341 AY829345 — —
Phellinidium fragrans CBS 202.90 Canada AY558619 AY059027 — —
Results Molecular phylogeny

In total, 22 new sequences of ITS, nrLSU, rpb2, and tef1-α generated in this study were submitted to GenBank, and the accession numbers are listed in Table 1. In addition, ITS, nrLSU, rpb2, and tef1-α sequences of 74 allied taxa (65 ITS, 72 nrLSU, 28 rpb2, and 39 tef1-α) were retrieved from GenBank (Table 1), along with the outgroups Coniferiporia weirii (CFS504) and Phellinidium fragrans (CBS 202.90). The concatenated multiple sequence alignment was 3,431 bases long, of which 1,854 were constant, 1,468 were variable, and 1,130 (33%) were parsimony informative. The ITS region comprised 879 bases, nrLSU 1,385 bases, rpb2 620 bases, and tef1-α 544 bases. Maximum likelihood (ML) and Bayesian inference (BI) analyses generated nearly identical tree topologies, with little variation in statistical support. Therefore, only the ML tree is shown (Fig. 1).

10.3897/mycokeys.125.168173.figure1 39D0567D-303F-5C64-B69D-40538F2C880C

Phylogram generated from maximum likelihood analysis based on combined ITS, nrLSU, rpb2, and tef1-α sequence data of species of Fuscoporia with the outgroups Phellinidium fragrans (CBS 202.90) and Coniferiporia weirii (CFS504). Bootstrap support values ≥ 60% for ML and Bayesian posterior probabilities ≥ 0.80 are indicated above the nodes. Species complexes are highlighted, and newly generated sequences are shown in bold red, blue, and pink; type specimens are tagged with “T.”

https://binary.pensoft.net/fig/1468445

The phylogenetic analyses revealed six major clades representing six species complexes in Fuscoporia, which were consistent with earlier reports (Fig. 1) (Chen et al. 2020, 2023a, b). Our Indian specimens were recovered in the F. torulosa complex, which received robust phylogenetic support in both ML and BI analyses (92 ML/1.00 BI). Fuscoporia sirumalaiensis formed a sister clade to F. rhabarbarina; F. indica formed a sister taxon to F. sirumalaiensis, F. rhabarbarina, F. licnoides, and F. callimorpha, while F. terminalianae formed a distinct lineage within the F. torulosa complex. The newly identified species are characterized by effused-reflexed to pileate basidiocarps, smaller pores, septate skeletal hyphae, the presence of hymenial setae, the absence of mycelial setae, the presence of cystidioles, and broadly ellipsoid to subglobose basidiospores.

 Taxonomy Fungi Hymenochaetales Hymenochaetaceae 666E4A48-EAC3-5CEE-958C-DD6DF86C6AC1 Fuscoporia indica MB 859271 M. Kaliyaperumal, S. Gunaseelan & K. Kezo sp. nov. Fig. 2 Diagnosis.

Fuscoporia indica is diagnosed by its annual, applanate, dimidiate basidiome with smooth to glabrous, indistinctly zonate pilear surface, homogenous context, the presence of cystidioles, and ellipsoidal, acyanophilic basidiospores.

 Holotype.

India • Tamil Nadu, Salem District, Yercaud, on hardwood, 23 January 2018, S. Gunaseelan, HRS-15A (holotype MUBL1104). GenBank: PP390498 (nrLSU); PV638735 (rpb2); PV638743 (tef1-α).

 Etymology.

Refers to the type locality, “India.”

 Description.

Basidiomes annual, pileate, applanate, sessile to imbricate, soft, to light corky when dry. Pilei dimidiate, convex, projecting up to 4.5 cm long, 7 cm broad, and 1 cm thick at the base. Pileal surface yellowish brown (5D8), brown (6E6) to dark brown (6F8), smooth to glabrous, indistinctly zonate. Margin brown (6E5), acute, margin sterile, light brown (6D8), 1 mm in thickness. Pore surface light brown (6D8) to dark brown (6F8). Pores round to angular, 6–8 per mm. Marginal setae absent. Context light brown (6D8), homogenous, up to 3 mm in thickness. Tube layer yellowish-brown (6D8) to brownish-yellow (5C7), hard and corky, with tubes up to 2 mm long.

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

Context. Generative hyphae, hyaline to pale yellow, thin to slightly thick-walled, branched, frequently septate, 2–4 µm; skeletal hyphae dominant, rust-brown, thick-walled with a medium to wide lumen, unbranched, rarely septate, more or less straight, and regularly arranged, 2.2–3.5 µm.

Tubes. Generative hyphae, dominant at the dissepiment edges and subhymenium, thin-walled, frequently branched and septate, hyaline to pale yellow, 1.8–3.4 µm, some encrusted at dissepiment edges and in hymenium; skeletal hyphae dominant, thick-walled with a medium to wide lumen, more or less straight, subparallel along the tubes, yellow to golden yellow, 2–3.2 µm. Hymenial setae subulate to ventricose, acute to acuminate at the apex, encrusted, mostly originating from tramal hyphae, dark brown, thick-walled, 8−39 × 5−8 µm; Cystidioles hyaline to pale yellow in water, fusoid to subulate, rare, tapering at the end, 6–42 × 2.8–7 µm. Basidioles clavate to broadly clavate 6.5–18 × 2.5–6.5 µm. Basidia broadly clavate, hyaline, four sterigmata, 7–17 × 2.8–6.7 µm. Basidiospores ellipsoid, hyaline, thin-walled, smooth, CB−, IKI−, (3.2–)3.5–4.5(–4.8) × (2.2–)2.5–2.9(–3.2) µm, Q=1.5, Q range = 1.3–1.7, (n = 30/2).

10.3897/mycokeys.125.168173.figure2 5C54793A-1A01-5432-BEB7-778D5D9A4492

Fuscoporia indica (holotype MUBL1104). a. Basidiomes; b. Pore surface; c. Transverse section of a basidiome; d. Dissepiment edges and skeletal hyphae; e. Cystidioles; f. Hymenial setae; g. Basidioles; h. Basidia; i. Basidiospores. Scale bars: 5 cm (a, b); 2 cm (c); 5 μm (d–i).

https://binary.pensoft.net/fig/1468446 Distribution.

India (Tamil Nadu).

 Additional specimen examined (paratype).

India • Tamil Nadu, Salem District, Yercaud (11°79'63"N, 78°21'20"E), on hardwood, 23 January 2018, S. Gunaseelan, HRS-15B. GenBank: PQ113747 (nrLSU); PV638736 (rpb2); PV638744 (tef1-α).

 Notes.

The phenetic characters of F. indica and F. licnoides are similar, both having a concentrically zonate, glabrous pileal surface, homogeneous context, and acute margin. However, F. indica differs in having imbricate, dimidiate, convex pilei and smaller basidiospores (F. indica 3.2–4.8 × 2.2–3.2 μm vs. F. licnoides 4–5 × 2.5–3.5 μm) (Oliveira and Gibertoni 2023). Fuscoporia indica differs from F. callimorpha in having an imbricate basidiome, convex pilei, smaller pores, and larger hymenial setae (Dai 2010). Fuscoporia indica, F. senex, and F. torulosa share similar features such as applanate, dimidiate basidiomes with a dimitic hyphal system, the presence of cystidioles, and cyanophilic basidiospores, but F. indica differs in having a glabrous, azonate pilear surface and larger basidiospores. Fuscoporia indica also differs from F. rhabarbarina by the absence of a crust above the context at maturity and, microscopically, by larger basidiospores. In contrast, F. rhabarbarina is identified by the presence of a crust above the context in older specimens and smaller spores (3.2–4.2 × 2–2.5 μm) (Dai 2010).

 Fungi Hymenochaetales Hymenochaetaceae CFCC5B7B-A5D2-56B4-ADDF-210A983564FD Fuscoporia sirumalaiensis MB 859272 E. Arumugam, S. Gunaseelan & M. Kaliyaperumal sp. nov. Fig. 3 Diagnosis.

Morphologically, Fuscoporia sirumalaiensis is unique with a perennial, imbricate to pileate, dimidiate basidiome, smooth to glabrous, concentrically zonate pilear surface, the presence of cystidioles, hymenial setae, and ellipsoidal basidiospores.

 Holotype.

India • Tamil Nadu, Dindigul District, Sirumalai (10°28'23"N, 78°01'54"E), on hardwood, 31 Dec. 2022, M. Kaliyaperumal, SRM09 (holotype MUBL1106). GenBank: PP390500 (nrLSU); PQ098037 (ITS); PV638737 (rpb2); PV638741 (tef1-α).

 Etymology.

Refers to the type locality “Sirumalai,” India.

 Description.

Basidiomes biennial to perennial, imbricate to pileate, sessile, applanate. Pilei dimidiate, projecting up to 5.2 cm long, 7 cm wide, and 2 cm thick near the attachment. Pilear surface dark brown (6F7) to greyish brown (6E3), smooth to glabrous, concentrically zonate. Margin brown (6E8), acute, up to 1 mm. Context light brown (6D8) to brown (6E8), duplex, 0.4 cm. Pore surface, light brown (6D6) to brown (6E7). Pores circular, 5–8 per mm. Tubes light brown (6D8) to brown (6E8), up to 1 cm long, each stratum up to 0.2 cm.

Hyphal system. 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 septate, 2–5 µm; skeletal hyphae rust-brown, thick-walled with a narrow to wide lumen, unbranched, aseptate, 2.5–5 µm.

Trama. Generative hyphae hyaline, thin to thick-walled, mostly present at dissepiment edges and subhymenium, frequently branched with simple septate, a few are encrusted at dissepiment edges and in the hymenium, 2.5–5.2 µm; skeletal hyphae dominant, yellowish brown, thick-walled with a medium to wide lumen, aseptate, subparallel along the tubes, 2.5–5.6 µm. Hymenial setae subulate to ventricose, indistinctly encrusted, dark brown, 20−45 × 5.2−9 µm. Cystidioles hyaline, fusoid to subulate, infrequent, tapering at the end, 9–28 × 2.5–7.2 µm. Basidioles broadly clavate 6.5–18 × 3.5–6.2 µm. Basidia broadly clavate, hyaline, four sterigmata, 6–17 × 4–6.5 µm. Basidiospores ellipsoid, hyaline, thin-walled, smooth, CB ̄, IKI ̄, (2.8–)3.1–3.6(–3.8) × (1.8–)2.1–2.6(–2.8) µm, Q=1.5, Q range = 1.2–1.6, (n = 30/2).

10.3897/mycokeys.125.168173.figure3 F3B1D84E-8713-5928-972D-F3631B54A3C0

Fuscoporia sirumalaiensis (holotype MUBL1106). a. Basidiomes; b. Pore surface; c. Transverse section of a basidiome (arrows indicate the stratified tube layers); d. Dissepiment edges and skeletal hyphae; e. Hymenial setae; f. Cystidioles; g. Basidioles; h. Basidia; i. Basidiospores. Scale bars: 2 cm (a–c); 5 μm (d–i).

https://binary.pensoft.net/fig/1468447 Distribution.

India (Tamil Nadu).

 Additional specimen examined.

India • Tamil Nadu, Dindigul District, Sirumalai (10°27'73"N, 78°01'57"E), on hardwood, 31 December 2022, M. Kaliyaperumal, SRM21. GenBank: PQ113746 (nrLSU); PQ098038 (ITS); PV638738 (rpb2); PV638742 (tef1-α).

 Notes.

Multigene phylogenetic analyses revealed that Fuscoporia sirumalaiensis forms a sister clade to F. rhabarbarina (83% ML/1 BI) within the F. torulosa complex (Chen et al. 2023a, b). Fuscoporia sirumalaiensis and F. rhabarbarina are similar only in their concentrically zonate brown basidiomes, but the two species differ greatly in other morphological characteristics such as the size of cystidioles, hymenial setae, and basidiospores (Dai 2010). The phenetic features of F. sirumalaiensis vary significantly from those of F. callimorpha, F. dhofarensis, and F. licnoides (Dai 2010; Oliveira and Gibertoni 2023). Fuscoporia sirumalaiensis differs from F. indica in having larger hymenial setae, differences in cystidiole size, and smaller basidiospores (F. sirumalaiensis (2.8–)3.1–3.6(–3.8) × (1.8–)2.1–2.6(–2.8) μm vs. F. indica (3.2–)3.5–4.5(–4.8) × (2.2–)2.5–2.9(–3.2) μm) (present study).

 Fungi Hymenochaetales Hymenochaetaceae 4E9D40F2-3415-5E87-BB22-D1F6B92034B9 Fuscoporia terminalianae MB 859273 E. Arumugam, S. Gunaseelan & M. Kaliyaperumal sp. nov. Fig. 4 Diagnosis.

Morphologically, F. terminalianae can be diagnosed by an annual, resupinate to effused, often pileate basidiome, infrequently warted, and widely zonate pilear surface with homogeneous context. Microscopically, the presence of cystidioles and ellipsoidal, acyanophilic basidiospores are characteristic features of this species.

 Holotype.

India • Tamil Nadu, Kallakurichi District, Vellimalai (11°86'16"N, 78°70'16"E), on dead wood (Terminalia bellerica), 30 Oct. 2019, M. Kaliyaperumal, VM2A (holotype MUBL1107). GenBank: PP390501 (nrLSU); PQ098033 (ITS); PV638733 (rpb2); PV638739 (tef1-α).

 Etymology.

Refers to the host tree Terminalia bellerica on which the fungus was collected.

 Description.

Basidiome annual, resupinate to effused, reflexed, pileate, applanate, sessile, fused to imbricate, soft when fresh, corky when dry. Pilei dimidiate, projecting up to 2.8 cm long, 5.4 cm wide, and 3.2 cm thick at the base. Pileal surface light brown (6D5), brown (6E8) to dark brown (6F8), smooth, widely zonate, infrequently warted. Margin brown (6E8), velutinate, obtuse to acute, up to 3 mm in thickness. Marginal setae absent. Pore surface brown (6E7) to dark brown (6F8), glancing. Pores round to angular, 6–9 per mm. Context light brown (6D8), zonate, homogenous, up to 3 mm in thickness. Tube layer golden brown (5D7) to light brown (6D8), hard and corky, with tubes up to 2 mm long.

10.3897/mycokeys.125.168173.figure4 268B7569-9D42-5A5A-AF4C-5DB9D844398B

Fuscoporia terminalianae (holotype MUBL1107). a. Basidiomes; b. Habitat; c. Pore surface; d. Transverse section of a basidiome; e. Dissepiment edges and skeletal hyphae; f, g. Hymenial setae; h. Cystidioles; i. Basidioles; j. Basidia; k. Basidiospores. Scale bars: 2 cm (a–d); 5 μm (e–k).

https://binary.pensoft.net/fig/1468448

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

Context. Generative hyphae hyaline to golden yellow, thin to thick-walled, branched, frequently septate, 2–5 µm; skeletal hyphae dominant, rust-brown, thick-walled with a medium to wide lumen, unbranched, occasionally septate, 2.5–4.8 µm.

Tubes. Generative hyphae mostly present at dissepiment edges and subhymenium, hyaline to pale yellow, thin to thick-walled, frequently branched and simple septate, 1.5–5.2 mm, most of the hyphae at dissepiment edges and hymenium are encrusted with simple crystals, skeletal hyphae dominant, yellowish brown, thick-walled with a medium to wide lumen, frequently septate, more or less straight, subparallel along the tubes, 2.5–5 mm. Hymenial setae ventricose to subulate, mostly originating from tramal hyphae, dark brown, thick-walled, 13−45 × 4−7 µm. Cystidioles hyaline, fusoid to subulate, rare, tapering at the end, 9–32 × 4.5–7 µm. Basidioles broadly clavate 5–13 × 2.5–6 µm. Basidia broadly clavate, hyaline, four sterigmata, 5.2–15 × 2.9–6 µm. Basidiospores ellipsoid, hyaline, thin-walled, smooth, CB ̄, IKI ̄, (3–)3.3–4.3(–4.5) × (2.5–)2.8–3 µm, Q=1.48, Q range=1.2–1.7, (n = 30/2).

 Additional specimen examined.

India • Tamil Nadu, Kallakurichi District, Vellimalai (11°87'77"N, 78°69'52"E), on dead wood (Terminalia bellerica), 30 Oct. 2019, M. Kaliyaperumal, VM2B. GenBank: PQ113745 (nrLSU); PQ098034 (ITS); PV638734 (rpb2); PV638740 (tef1-α).

 Notes.

In multigene phylogenetic analyses, F. terminalianae forms an isolated lineage within the F. torulosa complex. Fuscoporia rhabarbarina is distinguished from F. terminalianae by its glabrous, sulcate pilear surface, number of pores, and basidiospore size; in contrast, F. terminalianae has a smooth, infrequently warted, widely zonate pilear surface (Dai 2010). Fuscoporia licnoides and F. terminalianae are similar in having a zonate basidiome and homogeneous context, but the latter differs in having an effused-reflexed basidiome, larger setae, and smaller basidiospores (F. terminalianae 3–4.5 × 2.5–3 μm vs. F. licnoides 4–5 × 2.5–3.5 μm) (Oliveira and Gibertoni 2023). Fuscoporia terminalianae, F. senex, and F. torulosa are similar in having applanate, dimidiate basidiomes with narrowly zonate pilear surfaces and dimitic hyphal systems, as well as the presence of cystidioles and cyanophilic basidiospores, while F. terminalianae differs in having a warted pilear surface and smaller basidiospores (Dai 2010).

 Discussion

The present study contributes to the taxonomy of Indian representatives of the genus Fuscoporia by describing new taxa using an integrative approach that combines morphological observations with multimarker phylogenetic analyses. Compared with information available from classical taxonomy, phylogenetic analyses have been significant in resolving species limits and clarifying evolutionary relationships, enabling a more comprehensive classification of the taxa.

Chen et al. (2020) used phylogenetic analysis to reveal six major complexes within Fuscoporia. These were (i) the F. contigua group (resupinate basidiome, moderately large pores, the presence of mycelial setae and hymenial setae, and ellipsoid to broadly ellipsoid basidiospores), (ii) the F. ferrea group (resupinate basidiome, aseptate skeletal hyphae, absence of mycelial setae, the presence of hymenial setae and cystidioles, and cylindric basidiospores), (iii) the F. ferruginosa group (resupinate basidiome, relatively small pores, entire dissepiments, straight hymenial setae, the presence of mycelial setae, and ellipsoid basidiospores), (iv) the F. gilva group (effused-reflexed to pileate basidiome, indistinctly concentrically sulcate zones, hispid to rugose or nodulose pilear surface, lacerate dissepiments, the presence of cystidioles, and ellipsoid to cylindric basidiospores), (v) the F. torulosa group (resupinate, effused-reflexed to pileate basidiome, smaller pores, septate skeletal hyphae, straight or hooked hymenial setae, absence of mycelial setae, the presence of cystidioles, and broadly ellipsoid to subglobose basidiospores), and (vi) the F. viticola group (resupinate to effused-reflexed basidiome, moderately large pores, absence of mycelial setae, narrowly subulate and long hymenial setae, and cylindric, long basidiospores) (Chen et al. 2019, 2020).

Over the past decade, there has been significant progress in understanding the diversity and geographical distribution of Fuscoporia species (Dai 2010; Raymundo et al. 2013; Chen et al. 2019, 2020, 2023a, b; Hussain et al. 2022; Oliveira and Gibertoni 2023; Crous et al. 2024). India is characterized by rich vegetation and several biodiversity hotspots; nevertheless, limited attention has been devoted to the diversity of hymenochaetoid fungi. The few conventional taxonomic studies undertaken were confined to the northern parts of the country (Singh et al. 2014; Kumari et al. 2021). As described herein, we sampled and delimited three new species of Fuscoporia based on morpho-taxonomic characters and phylogenetic relationships ascertained from a concatenated ITS+nrLSU+rpb2+tef1-α sequence dataset. The three species—F. indica, F. sirumalaiensis, and F. terminalianae—were recovered in the F. torulosa complex (92% ML/1 BPP) (Fig. 1). They show significant variation in morpho-microscopic characters, and the grouping is consistent with the literature (Chen et al. 2020, 2023a, b). Recently, our team reported F. naditirana from the Eastern Ghats of India, a species recovered within the F. gilva complex (Crous et al. 2024).

Phylogenetically, the newly described species were recovered within the F. torulosa complex, which accommodates species characterized by resupinate, effused-reflexed to pileate basidiomes with septate skeletal hyphae, the presence of cystidioles, straight or hooked hymenial setae, and broadly ellipsoid to subglobose basidiospores (Chen et al. 2020, 2023a, b). Morphologically, F. terminalianae differs from F. indica and F. sirumalaiensis by having resupinate to imbricate and/or pileate basidiomes with infrequently warted pilear surfaces, obtuse margins, and larger basidiospores (3–4.5 × 2.5–3 μm). Likewise, F. sirumalaiensis differs from F. indica by its basidiome with a smooth to glabrous, narrowly zonate pilear surface, duplex context, larger pores (5–8/mm), and smaller basidiospores (2.8–3.8 × 1.8–2.8 μm).

Nevertheless, F. indica, F. sirumalaiensis, and F. terminalianae are phylogenetically and morphologically distinct from close allies and other species in the F. torulosa complex—F. australasica, F. callimorpha, F. chrysea, F. dhofarensis, F. eucalypti, F. hawaiiana, F. licnoides, F. montana, F. marquesiana, F. rhabarbarina, F. senex, F. roseocinerea, F. shoreae, F. torulosa, and F. wahlbergii (Chen et al. 2020, 2023a, b). The Indian species of Fuscoporia described herein have pileate basidiomes and lack hyphidia, whereas F. montana is characterized by completely resupinate basidiomes, occasionally with swollen edges, and the presence of hyphidia (Dai 2010; Chen et al. 2020). Likewise, F. chrysea differs from the three Indian species of Fuscoporia in having resupinate to effused-reflexed basidiomes that rarely form a narrow pileus or pseudo-pileus (Raymundo et al. 2013). Despite sharing pileate, applanate to dimidiate basidiomes, F. indica, F. sirumalaiensis, and F. terminalianae differ from F. australasica, F. eucalypti, F. marquesiana, and F. wahlbergii in having straight hymenial setae with acute tips, whereas the latter species are reported to have hooked hymenial setae (Dai 2010; Chen et al. 2020; Yuan et al. 2020). The absence of setae makes F. shoreae distinct from the Indian Fuscoporia species (Chen et al. 2020).

 Conclusion

Considering the significance of Fuscoporia in pharmacognosy and ecology, as well as the taxonomic disarray within the Hymenochaetaceae, effective species identification is crucial for future research. Taxonomic classification within the genus has remained challenging because of overlapping morphological traits and the limited resolution of the routinely used ITS region. In India, species of Fuscoporia were historically misclassified as Phellinus based solely on morphological characteristics. Our study highlights the importance of employing multimarker phylogenetic analysis in conjunction with detailed morphological observations to delineate the three novel species within the F. torulosa complex collected from the Eastern Ghats of Tamil Nadu. Furthermore, additional sampling from the understudied regions of Tamil Nadu, particularly the Western Ghats, is likely to enhance our understanding of the taxonomic complexity and species diversity of Fuscoporia.

 Acknowledgements

Malarvizhi Kaliyaperumal and Sugantha Gunaseelan thank EMR-SERB, DST (EMR/2016/003078), and the Government of India for financial assistance. MK and GS are grateful to the Principal Chief Conservator of Forests, Tamil Nadu Forest Department, for granting permission (E2/20458/2017) and for their assistance and support during field visits in the Eastern Ghats of Tamil Nadu. MK and Elangovan Arumugam acknowledge the Tamil Nadu State Council for Higher Education (RGP/2019-20/MU/HECP-0040) for financial aid. Samantha C. Karunarathan thanks the National Natural Science Foundation of China (No. 32260004), the High-Level Talent Recruitment Plan of Yunnan Province (High-End Foreign Experts Program), and the Key Laboratory of the Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River for their support. The authors extend their appreciation to the Ongoing Research Funding Program (ORF-Ctr-2025-6), King Saud University, Riyadh, Saudi Arabia. NS and JK thank Chiang Mai University for their support. Prof. Steven L. Stephenson, Department of Biological Sciences, University of Arkansas, USA, is thanked for English editing.

 Additional information Conflict of interest

The authors have declared that no competing interests exist.

 Ethical statement

No ethical statement was reported.

 Use of AI

No use of AI was reported.

 Funding

This research was funded by EMR-SERB, DST (EMR/2016/003078), Government of India; the Tamil Nadu State Council for Higher Education (RGP/2019-20/MU/HECP-0040); and the Ongoing Research Funding Program (ORF-Ctr-2025-6), King Saud University, Riyadh, Saudi Arabia.

 Author contributions

Conceptualization: MK, SG, KK, EA; Data curation: MK, SG, KK, EA; Formal analysis: MK; Funding acquisition: MK, AF, TA, AE, EC, SK; Investigation: MK, SG, KK, EA; Methodology: MK, KK, SG, EA; Project administration: MK; Resources: MK, SG, KK, EA; Software: MK, KK; Supervision: MK; Validation: MK, SG; Visualization: MK; Writing—original draft: MK, SG, KK, EA; Writing—review and editing: MK, KK, SK.

 Author ORCIDs

Sugantha Gunaseelan https://orcid.org/0000-0001-7089-2292

Kezhocuyi Kezo https://orcid.org/0000-0002-3723-0462

Elangovan Arumugam https://orcid.org/0009-0003-9720-2978

Samantha C. Karunarathna https://orcid.org/0000-0001-7080-0781

Abdallah M. Elgorban https://orcid.org/0000-0003-3664-7853

Jaturong Kumla https://orcid.org/0000-0002-3673-6541

Nakarin Suwannarach https://orcid.org/0000-0002-2653-1913

Ekachai Chukeatirote https://orcid.org/0000-0002-9968-5841

Malarvizhi Kaliyaperumal https://orcid.org/0000-0002-1218-3778

 Data availability

All holotype and paratype collections of the new species are deposited at the Madras University Botany Laboratory (MUBL), Centre for Advanced Studies in Botany, University of Madras, Chennai 600025, Tamil Nadu, India. The sequences generated during this study are deposited in NCBI GenBank. The ITS, nrLSU, partial rpb2, and tef1-α alignment is deposited in FigShare.

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