683 MycoKeys MycoKeys 119: 67-94 (2025) DOI: 10.3897/mycokeys.119.154387 Research Article Morphology and multigene phylogeny reveals five new species of Hydnaceae (Cantharellales, Basidiomycota) from China Qian Zhou'®, Chengbin Qian'’”®, Chuyun Zhang'2®, Qidong Su*®, Yiliang Li*®©, Shihui Zhang?®, Nian Mu*®, Taimin Xu*®, Hongmin Zhou'®, Changlin Zhao'24© 1 The Key Laboratory of Forest Resources Conservation and Utilization in the Southwest Mountains of China Ministry of Education, Key Laboratory of National Forestry and Grassland Administration on Biodiversity Conservation in Southwest China, Yunnan Provincial Key Laboratory for Conservation and Utilization of In-forest Resource, Southwest Forestry University, Kunming 650224, China 2 College of Forestry, Southwest Forestry University, Kunming 650224, China 3 Yunnan Tongbiguan Provincial Nature Reserve, Mangshi 679319, China 4 Yunnan Key Laboratory of Gastrodia and Fungal Symbiotic Biology, Zhaotong University, Zhaotong, 657000, China Corresponding author: Changlin Zhao (fungi@swfu.edu.cn) OPEN Qrceess Academic editor: Maria PR. Martin Received: 31 March 2025 Accepted: 15 May 2025 Published: 26 June 2025 Citation: Zhou Q, Qian C, Zhang C, Su Q, Li ¥, Zhang S, MUN, Xu T, Zhou H, Zhao C (2025) Morphology and multigene phylogeny reveals five new species of Hydnaceae (Cantharellales, Basidiomycota) from China. MycoKeys 119: 67-94. https://doi.org/10.3897/ mycokeys.119.154387 Copyright: © Qian Zhou et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Wood-inhabiting fungi play a fundamental role in ecosystem processes, particularly in wood degradation and the recycling of organic matter. In this study, a new genus, Clavu- liella gen. nov., and five new species, viz. Burgella albofarinacea sp. nov., B. fissurata sp. nov., Burgoa wumengshanensis sp. nov., Clavuliella sinensis sp. nov., and Sistotrema sin- ense sp. nov., are described from China and illustrated based on morphological charac- teristics and molecular phylogenetic analyses. Sequences of the ITS+nLSU genes were used for the phylogenetic analyses using Maximum Likelihood, Maximum Parsimony, and Bayesian Inference methods. The phylogram of the family Hydnaceae, based on the ITS+nLSU rDNA gene regions, included four genera; Burgella, Burgoa, Clavuliella and Sis- totrema. The topology based on these sequences revealed that Burgella albofarinacea was closely related to B. flavoparmeliae, and B. fissurata was grouped with B. lutea. The taxon Burgoa wumengshanensis was sister to the clade that included B. anomala and B. verzuoliana. The species Sistotrema sinense was grouped closely with S. brinkmannii and S. farinaceum. All new taxa can be readily recognized by their macroscopic and an- atomical characteristics. The five new species, closely related taxa in the phylogenetic tree, and morphologically similar species are discussed. Key words: Asia, biodiversity, molecular systematics, taxonomy, wood-inhabiting fungi Introduction Fungi constitute an integral and valuable part of our natural ecosystem and play an essential ecological role in driving carbon cycling in forest soils, medi- ating mineral nutrition of plants, and alleviating carbon limitations (Chen et al. 2023a, b; Niego et al. 2023; Dong et al. 2024a; Yang et al. 2025). The wood-in- habiting fungal family Hydnaceae Chevall includes many variations of the fruit- ing body types within the order Cantharellales J. Schrot (Cai and Zhao 2023; Gao et al. 2024), in which it comprises many representative wood-inhabiting 67 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae fungi taxa, such as bulbil-shaped, hypochnoid, corticioid, odontoid, poroid, cla- varioid, ramarioid, mucronelloid, cantharelloid, and hydnoid basidiomes with di- verse hymenophoral and cystidial morphology (Uehling et al. 2012; Diederich et al. 2014; Gruhn et al. 2017; Masumoto and Degawa (2020); Lawrey et al. 2020; Bondartseva and Zmitrovich 2023; Cai and Zhao 2023; Gao et al. 2024). The genus Burgella Diederich & Lawrey (Hydnaceae, Cantharellales), typified by B. flavoparmeliae Diederich & Lawrey, is characterized by the following features: conidia cylindrical, conidiophores short, hyphae hyaline, septate, straight, rarely branched or anastomosed. Agglomerations of bulbils gelatinous in appearance, al- most coralloid, composed of irregularly shaped bulbils; bulbils externally and inter- nally composed of irregular, roundish or elongate cells with clamped septa (Died- erich and Lawrey 2007). Based on the MycoBank database (http://www.mycobank. org, accessed on 30 March 2025) and the Index Fungorum (http://www.indexfun- gorum.org, accessed on 30 March 2025), the genus Burgella has 2 registered spe- cies and intraspecies names (Diederich and Lawrey 2007). Previous studies have shown that B. flavoparmeliae and Sistotrema oblongisporum M.P. Christ. & Hauer- slev were the sister group of the genus Multiclavula (Diederich and Lawrey 2007). The species B. flavoparmeliae was only distantly related to the type species of the genus Burgoa Goid., which appeared in a different place in the order Cantharellales, and the research revealed that B. flavoparmeliae should not be included in Burgoa, but instead placed in the new genus Burgella (Diederich and Lawrey 2007). The genus Burgoa, typified by B. verzuoliana Goid. (Hydnaceae, Cantharellales, Agaricomycotina), was established by Goidfnich to accommodate microfungi producing multicellular spore-like structures with differentiated peridial and in- ternal cells, i.e. bulbils. Apart from the production of the bulbils, members of this genus were distinguished by the formation of clamp connections on their myce- lium. This feature showed their affinity to members of Agaricomycotina, but their position within the order Cantharellales was recognised only recently (Diederich and Lawrey 2007; Koukol and Kubatova 2015; Kiyuna et al. 2015). The genus Burgoa is a peculiar microscopic basidiomycete not forming any basidiocarps in its life cycle. So far, this saprotroph has sporadically been found mainly on differ- ent woody substrates but the overall knowledge of its ecology and distribution remains sparse due to its rarity (Koukol and Kubatova 2015). Based on the Myco- Bank database (http://www.mycobank.org, accessed on 30 March 2025) and the Index Fungorum (http://www.indexfungorum.org, accessed on 30 March 2025), the genus Burgoa has 10 registered species and intraspecies names (Diederich and Lawrey 2007; Koukol and Kubatova 2015; Kiyuna et al. 2015). Clavulina J. Schrot. (Hydnaceae, Cantharellales), with Clavulina cristata (Hol- msk.) J. Schr6t. as its type species, was established in 1888 (Schréter 1888; He et al. 2019). In China, 14 Clavulina species have been reported on the basis of morphological and molecular analyses, most of which are found in subtropical regions (Gao et al. 2024). In the present study, the new genus Clavuliella falls within Hydnaceae (Cantharellales) and is closely related to Clavulina. The genus Sistotrema Fr. (Hydnaceae, Cantharellales, Agaricomycetes, Agar- icomycotina, Basidiomycota), typified by S. confluens Pers., is a comparative- ly large genus belonging to the phylum Basidiomycota and is morphologically characterized by resupinate or pileate-stipitate, soft basidiomes, smooth, gran- dinioid, hydnoid, or poroid hymenophore with various characteristic textures (pellicular, membranaceous, or ceraceous), a Monomitic hyphal system with MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 68 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae oily inclusions, urniform basidia, and smooth, thin-walled, basidiospores con- taining cytoplasmic oil droplets (Eriksson et al. 1984; Bernicchia and Gorjon 2010; Cai and Zhao 2023). Based on the MycoBank database (http://www. mycobank.org, accessed on 30 March 2025) and the Index Fungorum (http:// www.indexfungorum.org, accessed 30 March 2025), the genus Sistotrema has 224 registered species and intraspecies names, however the actual number of recognized species is 111 (Eriksson et al. 1984; Bernicchia and Gorjén 2010; Sugawara et al. 2022; Cai and Zhao 2023). In the present study, extensive morphological examinations, combined with analyses of multi-gene sequences data, support the introduction of a new ge- nus and 4 new species of wood-inhabiting fungi. Descriptions and illustrations based on morphological characteristics are presented along with evidence from phylogenetic analyses. Materials and methods Morphology Fresh basidiomata of the fungi growing on angiosperm branches were col- lected from the Dali, Dehong, and Zhaotong of Yunnan Province, and Guiyang of Guizhou Province, P.R. China. Specimens were dried in an electric food de- hydrator at 40 °C (Dong et al. 2024b) then sealed and stored in an envelope and deposited in the Herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, PR. China. Macromorphological descriptions were based on field notes and photos captured in the field and lab. Colour terminol- ogy followed Petersen (1996). Micromorphological data were obtained from the dried specimens when observed under a light microscope following the previous study (Cai and Zhao 2023). The following abbreviations are used for the micro characteristic description: KOH = 5% potassium hydroxide water solution, CB = Cotton Blue, CB- = acyanophilous, IKI = Melzer’s Reagent, IKI- = both inamyloid and indextrinoid, L = mean spore length (arithmetic average for all spores), W = mean spore width (arithmetic average for all spores), Q = vari- ation in the L/W ratios between the specimens studied and n = a/b (number of spores (a) measured from given number (b) of specimens). Molecular phylogeny The EZNA HP Fungal DNA Kit (Omega Biotechnologies Co., Ltd., Kunming, China) was used to extract DNA with some modifications from the dried specimens. The nuclear ribosomal ITS region was amplified with primers ITS5 and ITS4 (White et al. 1990). The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 58 °C for 45 s and 72 °C for 1 min, with a final extension of 72 °C for 10 mins. The nuclear nLSU region was am- plified with primer pair LROR and LR7 (Rehner and Samuels 1994). The PCR pro- cedure for nLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 48 °C for 1 min and 72 °C for 1.5 mins with a final extension of 72 °C for 10 mins. The PCR procedure for ITS and nLSU followed a previous study (Zhao and Wu 2017). All newly generated sequences were depos- ited in NCBI GenBank (https://www.ncbi.nim.nih.gov/genbank/) (Table 1). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 69 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Table 1. Names, specimen numbers, references and corresponding GenBank accession numbers of the taxa used in this study. Bergerella atrofusca Bryoclavula phycophila Species name B. phycophila Burgella albofarinacea B. DoDD Daa dD DD . albofarinacea . albofarinacea . albofarinacea . fissurata . flavoparmeliae flavoparmeliae flavoparmeliae flavoparmeliae flavoparmeliae flavoparmeliae B. lutea Burgella sp. Burgella sp. Burgella sp. Burgoa anomala B. wumengshanensis B. verzuoliana Clavuliella sinensis Clavulina cristata C. iris C. minor C. minor C. parvispora C. parvispora C. samuelsii C. samuelsii C. subrugosa C. subrugosa C. sphaeropedunculata C. sphaeropedunculata Hydnum albidum H. albomagnum H. rufescens Minimedusa obcoronata M. polyspora M. polyspora Multiclavula caput- serpentis M. corynoides M. mucida M. petricola M. vernalis Specimen No. BR Berger 34240 Hiroshi:Bryoclavula4 S-287-FB3 CLZhao 31820 CLZhao 32468 CLZhao 32026 CLZhao 31855 CLZhao 30212 Flakus 23513 Buck 38682 JL192-01 SV1 JL192-01 SV2 JL192-01 SV3 JL192-01 SV4 Etayo 27623 WS34_1_2_A_ As_10000 HHB-19354 HHB-19352 CBS 130.38 CLZhao 33227 CBS 131.38 CLZhao 31231 EL95_97 ML 5135C1 B30912949 B30912949 FCME 27650 FCME 27657 TENN065723 PDD:89881 TENN043395 TN43395 FCME 27661 MEXU 28222 MB11-6024/2 AFTOL-ID 471 MB18-6024/1 CBS 120605 CBS3113:1:6 SH-Ecto-3 KaiR699 Lutzoni 930804-2 AFTOL-ID 1130 NBRC 114399 Lutzoni 930806-1 GenBank accession No. ITS MN902070 0Q791465 LC544109 PQ758751 PQ758754 PQ758753 PQ758752 PQ758749 OR471304 OR471305 OR471306 OR471307 KC336076 LC631658 MW740322 MW740323 AB972780 PQ758755 AB972781 PQ758750 MNO028412 OP738993 OR149156 MH542550 MH542549 JQ638712 GU222317 JQ638711 JN228221 MH542560 MH542557 DQ218305 GQ303278 MW386064 U66440 DQ521417 LC516464 U66439 MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 nLSU MN902070 0Q791464 PQ758759 PQ758762 PQ758761 PQ758760 PQ758757 KC336074 DQ915469 KC336075 PQ758758 AY586648 MN028396 OP737360 OR145333 MN049492 MN049491 JN228221 MK253716 MK253717 AY293186 AY700199 AY293187 GQ303309 MH866167 MG833798 MW369074 U66440 AY885163 LC516465 U66439 Country Austria Japan Japan China China China China China USA Bolivia USA USA USA USA Bolivia Japan New Zealand New Zealand Japan China Italy China Sweden Cyprus China China Mexico Mexico USA New Zealand USA New Zealand Mexico Mexico Thailand USA Panama USA USA China Japan USA Switzerland USA USA References Lawrey et al. (2020) NCBI Database Masumoto and Degawa (2020) Present study Present study Present study Present study Present study Diederich et al. (2014) Diederich et al. (2014) Swenie et al. (2023) Swenie et al. (2023) Swenie et al. (2023) Swenie et al. (2023) Diederich et al. (2014) Unpublished Unpublished Unpublished Kiyuna et al. (2015) Present study Kiyuna et al. (2015) Present study Larsson et al. (2004) Campo et al. (2023) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Gao et al. (2024) Binder et al. (2005) Masumoto and Degawa (2020) Binder et al. (2005) Diederich and Lawrey (2007) Vu et al. (2019) NCBI Database Reschke et al. (2021) Lutzoni (1997) Masumoto and Degawa (2020) Masumoto and Degawa (2020) Lutzoni (1997) 70 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Species name Neoburgoa freyi N. freyi Platygloea disciformis Rogersiomyces malaysianus Sistotrema confluens S. confluens S. adnatum S. adnatum S. alboluteum S. alboluteum S. albopallescens S. athelioides S. brinkmannii S. biggsiae S. chloroporum S. citriforme S. coroniferum S. coroniferum S. coronilla S. efibulatum S. epiphyllum S. eximum S. eximum S. farinaceum S. farinaceum S. flavorhizomorphae S. flavorhizomorphae S. hypogaeum S. luteoviride S. muscicola S. muscicola S. oblongisporum S. octosporum S. octosporum S. pistilliferum S. raduloides S. raduloides S. resinicystidium S. sernanderi S. sernanderi S. sinense S. subconfluens S. subconfluens S. yunnanense S. yunnanense S. brinkmannii Sistotremella perpusilla S. perpusilla Specimen No. JL596-16 EZ4455 AFTOL-ID 710 LE-BIN 3507 FCUG 298 AFTOL-ID 613 FCUG 700 GB700 TAA167982 TAA180259 KHL11070 FCUG 701 NH11412 FCUG 782 TUMH 64399 KHL15898 GB-BN-2 KH Larsson s.n. NH7598 FCUG 1175 CBS A-21517 Thorn429 CBS:531.91 FCUG 659 HK23176 TUMH:64401 TUMH:64402 CBS 394.63 H HK23176 KHL8791 KHL 11721 KHL 14077 FCUG 2822 CBS:126038 EL 28/10 AFTOL-ID 619 LR 44004 FCUG 2188 GB-BN-4 PUL:F24593 CLZhao 24876 Dai 12577 Dai 12578 CLZhao 7357 CLZhao 7395 NH11412 CBS 126048 HFRG EJ210404 GenBank accession No. ITS KX423755 OR471314 DQ234556 KT779285 DQ267125 OR464426 NR178117 KF218962 KF218968 NR155795 MH862275 KF218963 LC642038 LC642040 MH858314 NR158892 MH864053 KF218964 KF218969 MW448599 PQ758748 JX076812 ON817194 ON817195 MH864061 OL828790 MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 nLSU KX423755 OR471068 AY629314 DQ898711 AY647214 DQ898699 OR460895 AY586713 AJ606042 AM259210 DQ898700 AF506473 DQ898697 LC642057 KF218962 AM259215 KF218968 AF506475 DQ898696 AF393076 MH873956 DQ898707 KF218963 LC642059 LC642060 MH869926 AF506474 AJ606040 KF218970 DQ898698 MH875510 KF218964 AY647213 KF218969 DQ898708 AM259219 PQ758756 JX076810 JX076811 ON810362 ON810363 AF506473 MH875516 Country Vietnam Canada USA Poland Canada Canada Sweden Sweden Canada Sweden Canada Japan Sweden Sweden Sweden Sweden Canada Netherlands USA Canada Canada Finland Japan Japan Australia Finland Sweden Finland Sweden Canada USA Spain USA Finland Canada Sweden USA China China China China China Sweden USA UK Thailand USA Panama References Lawrey et al. (2016) Swenie et al. (2023) Sugawara et al. (2022) Psurtseva et al. (2016) Moncalvo et al. (2006) Masumoto and Degawa (2020) Moncalvo et al. (2006) Swenie et al. (2023) Larsson et al. (2004) Nilsson et al. (2006) Nilsson et al. (2006) Moncalvo et al. (2006) Larsson et al. (2004) Moncalvo et al. (2006) Sugawara et al. (2022) Kotiranta and Larsson (2013) Nilsson et al. (2006) Kotiranta and Larsson (2013) Larsson et al. (2004) Moncalvo et al. (2006) NCBI Database Binder and Hibbett (2002) Vu et al. (2019) Moncalvo et al. (2006) Kotiranta and Larsson (2013) Sugawara et al. (2022) Sugawara et al. (2022) Vu et al. (2019) Kotiranta and Larsson (2013) Larsson et al. (2004) Nilsson et al. (2006) Kotiranta and Larsson (2013) Moncalvo et al. (2006) Vu et al. (2019) Kotiranta and Larsson (2013) Masumoto and Degawa (2020) Kotiranta and Larsson (2013) Moncalvo et al. (2006) Nilsson et al. (2006) e NCBI Database | Present study Zhou and Qin (2013) Zhou and Qin (2013) Cai and Zhao (2023) Cai and Zhao (2023) Larsson et al. (2004) Vu et al. (2019) NCBI Database Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae The sequences were aligned in MAFFT version 7 (Katoh et al. 2019) using the G-INS-i strategy. The alignment was adjusted manually using AliView ver- sion 1.27 (Larsson 2014). The sequence alignments were deposited in Tree- Base (https://treebase.org/treebase-web/home.html;jsessionid=4359D21 8F- 4D60336C2A9F7EB7D135CCD) (ID 32177 (Fig. 1)). The sequence alignments were deposited in TreeBase (https://treebase.org/treebase-web/home.html,j- sessionid=4359D218F4D60336C2A9F7EB7D135CCD) (ID 32178 (Fig. 2)). Se- quences of Platygloea disciformis (Fr.) Neuhoff retrieved from GenBank were used as the outgroup in the ITS+nLSU analysis (Figs 1, 2; Sugawara et al. 2022). Maximum Parsimony (MP), Maximum Likelihood (ML), and Bayesian Inference (BI) analyses were applied to the combined three datasets following a previous study (Zhao and Wu 2017). All characters were equally weighted and gaps were treated as missing data. Trees were inferred using the heuristic search option with TBR branch swapping and 1,000 random sequence additions. Max trees were set to 5,000, branches of zero length were collapsed, and all parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1,000 pseudo replicates (Felsenstein 1985). Descriptive tree statistics - tree length (TL), composite consistency index (Cl), composite retention index (RI), composite rescaled consistency index (RC) and composite homoplasy index (HI) — were calculated for each maximum parsimonious tree generated. The combined data- set was also analysed using Maximum Likelihood (ML) in RAXxML-HPC2 through the CIPRES Science Gateway (Miller et al. 2012). Branch support (BS) for the ML analysis was determined by 1,000 bootstrap pseudo replicates. MrModelTest 2.3 (Nylander 2004) was used to determine the best-fit evolu- tion model for each dataset for the purposes of Bayesian Inference (BI) which was performed using MrBayes 3.2.7a with a GTR+I+G model of DNA substitu- tion and a gamma distribution rate variation across sites (Ronquist et al. 2012). A total of four Markov chains were run for two runs from random starting trees for 2 million generations for ITS+nLSU (Fig. 1) and 2 million generations for ITS+nLSU (Fig. 2) with trees and parameters sampled every 1,000 generations. The first quarter of all the generations were discarded as burn-in. A majority rule consensus tree was computed from the remaining trees. Branches were considered as significantly supported if they received a Maximum Likelihood bootstrap support value (BS) of = 70%, a maximum parsimony bootstrap sup- port value (BT) of = 70%, or a Bayesian posterior probability (BPP) of = 0.95. Results Molecular phylogeny Burgella albofarinacea BLASTN homology search using the ITS nucleotide sequence indicated that the sequence had 87% identity with the sequence as OR471304, named Burgella flavoparmeliae from the NCBI culture collec- tion (551/635 bp); the nLSU sequence had 98% identity with the sequence as DQ915469, named B. flavoparmeliae from the NCBI culture collection (1294/1323 bp). Burgella fissurata BLASTN homology search using the ITS nucleotide sequence indicated that the sequence had 88% identity with the sequence as OR471304, named B. flavoparmeliae from the NCBI culture col- lection (553/627 bp); the nLSU sequence had 98% identity with the sequence MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 72 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Multiclavula petricala NBRC 114399 Japan Multiclavula caput-serpentis KaiR699 Panama “Bre Sistotrema brinkmannii NH11412. Sweden 1 Sistotrema Bergerella atrofusca BR Berger 34240 Austiia W ] Bergerella | -/°"" Neoburgoa freyi Bz4455 USAW ! Neoburgoa Multiclavula vernalis Lutzoni 930806-1 USA 1PO/100/1.08 as iticlavula corynoides Lutzoni 930804-2 USA Multiclavula mucida AFTOL-ID 1130 USA | Multiclavula Rogersiomyces malaysianus LE-BIN 3507 Vietnam Rogersiomyces Sistotrema sinense CLZhao 24876 China 9 Irs Sistotrema farinaceum FCUG 659 Canada in Sistotrema coroniferum GB-BN-2 Sweden Sistotrema resinicystidium FCUG 2188 Canada Sistotrema yunnanense CLZhao 7357 China 100/100! -L sistotrema yunnanense CLZhao 7395 China 83 1.007 Burgella sp. HHB-19352 New Zealand c he 61/1.00 Burgella Sp. HHB-19354 New Zealand AN Sistotrema | Burgeila sp. Bolivia Burgella lutea Etayo 27623 Bolivia ; Burgella fissurata CLZhao 30212 China Burgelia sp. WS34_1_2_A_As 10000 Japan -Burgella flavoparmeliae JL192-01 SV3 USA \ ‘Burgella flavoparmeliae JL192-01SV4 USA “ al ‘Burgella flavoparmeliae JL192-01 Sv2 UsA ™ Burgella -Burgella flavoparmeliae JL192-01 SV1 USA Burgella flavoparmeliae Buck 38682 eA ; Burgelia flaveparmeliae Flakus 23513 USA al Burgella albofarinacea CLZhao 31820 China 9% Burgella albofarinacea CLZhao 32468 China Veoburgoa freyi JLS96-16 Switzerland hina Sistotrema subconfluens Dai 12577 China 100/100/1.09 ) Sistotrema confluens AFTOL-ID 613 USA | Sistotrema 100/100/1.01 si stotrema confluens FCUG 298 Canada Hydnum rufescens MB18-6024/1 USA _|-Hydnin albomagnum AFTOL-ID 471 USA | Hydnum ‘Hydnum albidum MB11-6024/2 USA Sistotrema albopallescens KHL11070 Sweden Sistotrema alboluteum TAA167982 Sweden | Sistotrema Sistotrema flaverhizomorphae TUMH:64401 Japan UIST. Sistotrema athelioides FCUG 701 Canada 7 2 TS lesy -FB3 Japan - | Br voclavula [ a ee en we al LY hiBryoclavulad Japan ee) | ee ae _— | 100/100/1.00 =e 98/-/1.00 Minimedusa obcoronata CBS 120605 Thailand || Minimedusa polyspora SH-Ecto-3 China Minimedusa -!17'-1 veinimedusa polyspora CBS 113.16 USA * | 100/100/1.00 Sistotrema hypogaewn CBS 394.63 Australia -/100/ - Sistotrema adnatum FCUG 700 Canada : 4 ae hs Sistotrema sernanderi GB-BN-4 Sweden wae ps Sistotrema efibulatum FCUG 1175 Canada Sistotrema Sistotrema eximum Thom429 USA Sistotrema bigesiae FCUG 782 Canada Sistotrema octosporum FCUG 2822 Canada -/98/1.00 Figure 1. Maximum Parsimony strict consensus tree illustrating the phylogeny of four new species and a new genus within Hydnaceae, based on ITS+nLSU sequences. Branches are labelled with Maximum Likelihood bootstrap values > 70%, parsimony bootstrap values = 50% and Bayesian posterior probabilities = 0.95, respectively. as DQ915469, named B. flavoparmeliae from the NCBI culture collection (1290/1319 bp). Burgoa wumengshanensis BLASTN homology search using the ITS nucleotide sequence indicated that the sequence had 83% identity with the sequence as AB972780, named Burgella flavoparmeliae from the CBS cul- ture collection (532/643 bp). Clavuliella sinensis BLASTN homology search using the ITS nucleotide sequence indicated that the sequence had 88% iden- tity with the sequence as MT196962, named Clavulina castaneipes (G.F. Atk.) Corner. from the NCBI culture collection (602/688 bp); the nLSU sequence had MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 73 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae * Type species . Holotype 98/100/1.00—Sistotrema luteoviride HHK23176 Finland 98/100/1.00 Sistotrema farinaceum HK23176 Finland sale Sistotrema citriforme KHL15898 Sweden ‘Sistotrema pistilliferum EL 28/10 Spain 99/100/1.00[-Sistotrema yunnanense CLZhao 7357 China ‘Sistotrema yunnanense CLZhao 7395 China Sistotrema coroniferum KH Larsson s.n. Sweden ho/1ooods'stotrema coroniferum GB-BN-2 Sweden Sistotrema resinicystidium FCUG 2188 Canada Sistotrema oblongisporum KHL 14077 Sweden ‘Sistotrema brinkmannii NH11412 Sweden as Sistotrema sinense CLZhao 24876 China ‘Sistotrema farinaceum FCUG 659 Canada Sistotrema raduloides AFTOL-ID 619 USA - 99/1 00's; stotrema raduloides AFTOL-ID 619 USA Toorino/iGo See) = 53) ‘Sistotrema muscicola KHL 11721 Finland - /83/- Sistotrema muscicola KHL8791 Sweden Ce istotrema flavorhizomorphae TUMH:64401 Japan NS | —|Sistotrema flavorhizomorphae TUMH:64402 Japan at ‘Sistotrema alboluteum TAA167982 Sweden NY Sistotrema alboluteum TAA 180259 Sweden ‘Sistotrema albopallescens KHL11070 Sweden ‘Sistotrema chloroporum TUMH 64399 Japan - /58/- [Sistotrema efibulatum FCUG 1175 Canada ‘eo/.og Sistotrema sernanderi GB-BN-4 Sweden istotrema eximum CBS:531.912 Canada Sistotrema eximum Thorn429 USA Sistotrema biggsiae FCUG 782 Canada istotrema octosporum CBS:126038 USA 100/96/1.00 'sistotrema octosporum FCUG 2822 Canada _ /100/.- 71/63/1.00[Sistotrema coronilla NH7598 Sweden ‘Sistotrema hypogaeum CBS 394.63 Australia 7097/1.) isistotrema adnatum FCUG 700 Canada 100/99/1.00'sistotrema adnatum GB700 Canada ‘Sistotrema athelioides FCUG 701 Canada Sistotrema epiphyvilum CBS H-21517 Netherlands - /87/0.99 Sistotrema confluens TK-00495 He istotrema subconfluens Dai 12577 China -/99/1.00' si stotrema subconfluens Dai 12578 China Sistotrema sernanderi PUL: F24593 USA Platygloea disciformis AFTOL-ID 710 USA | Outgroup -/-/1.00 81/79/1.00 89/83/1.00 - /65/1.00 - (54! - - /59/0.99. Sistotrema P9/96/1.00 - /81/1.00 50 Figure 2. Maximum parsimony strict consensus tree illustrating the phylogeny of the one new species and related spe- cies in Sistotrema, based on ITS+nLSU sequences. Branches are labelled with Maximum Likelihood bootstrap values > 70%, parsimony bootstrap values > 50% and Bayesian posterior probabilities > 0.95, respectively. 96% identity with the sequence as OM942737, named C. iris from the NCBI cul- ture collection (1281/1338 bp). Sistotrema sinense BLASTN homology search using the ITS nucleotide sequence indicated that the sequence had 81% iden- tity with the sequence as OM100765, named S. coroniferum (Hohn. & Litsch.) Donk from the NCBI culture collection (541/669 bp); the nLSU sequence had 99% identity with the sequence as OR460882, named S. brinkmannii (Bres.) J. Erikss. from the NCBI culture collection (1324/1340 bp). The ITS+nLSU dataset (Fig. 1) comprised sequences from 55 fungal speci- mens representing 73 taxa. The dataset had an aligned length of 3499 charac- ters, of which 2000 characters were constant, 546 were variable and parsimo- ny-uninformative, and 953 were parsimony-informative. Maximum parsimony analysis yielded 6 equally parsimonious trees (TL = 5747, Cl = 0.4047, HI = 0.5953, RI = 0.5604 and RC = 0.2268). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.207165 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 362.5. The phylogram, based on the ITS+nLSU rDNA gene regions (Fig. 1), included five genera, viz. Burgella, MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 74 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Burgoa, Clavuliella and Sistotrema, within the family Hydnaceae (Cantharellales, Agaricomycetes). The phylogenetic tree (Fig. 1) inferred from the ITS and nLSU sequences highlighted that the two new species were grouped into the genus Burgella, in which B. albofarinacea was closely related to B. flavoparmeliae, and B. fissurata was grouped with B. /utea Diederich, Capdet, A.l. Romero & Etayo. The phylogram based on the ITS and nLSU data (Fig. 1) showed that Burgoa wumengshanensis was clustered into the genus Burgoa, in which it was closely related to B. anomala (Hotson) Goid. and B. verzuoliana Goid. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 1), included one new species, viz. Sistotrema sinense, in which it was grouped into the genus Sistotrema. The ITS+nLSU dataset (Fig. 2) comprised sequences from 30 fungal speci- mens representing 40 taxa. The dataset had an aligned length of 2031 characters, of which 1085 characters were constant, 365 were variable and parsimony-unin- formative, and 581 were parsimony-informative. Maximum parsimony analysis yielded 12 equally parsimonious trees (TL = 2690, Cl = 0.5487, HI = 0.4513, RI = 0.5550 and RC = 0.3045). The best model of nucleotide evolution for the ITS+n- LSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.005023 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 813.5. The phylogenetic tree (Fig. 2), inferred from the ITS+nLSU sequences, highlighted that Sistotrema sinense was grouped closely with S. brinkmannii (Bres.) J. Erikss. and S. farinaceum Hallenb. Taxonomy Burgella albofarinacea Q. Zhou & C.L. Zhao, sp. nov. MycoBank No: 857296 Pigs:3=5 Holotype. CHINA * Yunnan Province, Zhaotong, Yiliang County, Longhai Town, Jianfeng mountain, GPS coordinates: 27°76'N, 104°37'E, altitude: 1777 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 26 August 2023, CL Zhao 31820 (SWFC). Etymology. a/bofarinacea (Lat.): refers to the albicans and farinaceous hy- menophore of the type specimens. Description. Basidiomata annual, resupinate, adnate, pellicular, coriaceous, without odor or taste when fresh, up to 11.5 cm long, 2 cm wide, 50-100 um thick. Hymenial surface smooth, farinaceous, white when fresh and drying, cracked. Sterile margin thin, white, thinning out, up to 1 mm wide. Hyphal system monomitic, generative hyphae with clamp connections, some- times with oily contents, colorless, slightly thick-walled, frequently branched, interwoven, 3.5-5.5 um in diameter; IKI-, CB-, tissues unchanged in KOH. Cystidia and cystidioles absent. Basidia suburniform to urniform, slightly thick-walled, with 8 sterigmata and a basal clamp connection, 10.5-22 x 3.5- 7 um; basidioles abundant, in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid to allantoid, colorless, thin-walled, smooth, with oil drop, IKI-, CB-, (4-)4.5-6.5(-7) x 2-4 um, L = 5.34 um, W = 2.83 um, Q = 1.79-1.97 um (n = 120/4). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 75 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae ut BAS Figure 3. Basidiomata of Burgella albofarinacea (holotype). Scale bars: 1 cm (A); 1 mm (B). Additional specimens examined (paratypes). CHINA * Yunnan Province, Zha- otong, Yiliang County, Longan Town, GPS coordinates: 27°73'N, 104°16E, alti- tude: 1550 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 27 August 2023, CLZhao 32026, CLZhao 31855 (SWFC) « Zhaotong, Yiliang County, Xiaocao- ba Town, GPS coordinates: 27°26'N, 104°26'E, altitude: 2225 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 28 August 2023, CLZhao 32468 (SWFC). Burgella fissurata Q. Zhou & C.L. Zhao, sp. nov. MycoBank No: 857297 Figs 6-8 Holotype. CHINA « Yunnan Province: Dehong, Yingjiang County, Tongbiguan provin- cial nature reserve, GPS coordinates: 24°30'N, 097°30E, altitude: 1300 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 19 July 2023, CLZhao 30212 (SWFC). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 76 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 4. Microscopic structures of Burgella albofarinacea (holotype): basidiospores (A), basidia and basidioles (B), a section of hymenium (C). Scale bars: 5 ym (A); 10 um (B, C). Figure 5. Microscopic structures of Burgella albofarinacea (holotype): basidiospores (A), basidia (B), basidioles (C), a section of hymenium (D), generative hyphae (E, F). Scale bars: 10 um (A-F). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 77 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae ue & Pc: = Bike" = Figure 6. Basidiomata of Burgella fissurata (holotype). Scale bars: 1 cm (A); 1 mm (B). Etymology. fissurata (Lat.): refers to the cracking hymenial surface of the type specimens. Description. Basidiomata annual, resupinate, adnate, pruinose, hypochnoid, without odor or taste when fresh, up to 10.2 cm long, 1 cm wide, 50-100 um thick. Hymenial surface smooth, cracked, white when fresh, turning to pale cream upon drying. Sterile margin thin, white, thinning out, up to 1 mm wide. Hyphal system monomitic, generative hyphae with clamp connections, col- orless, thin-walled, frequently branched, interwoven, 2-4 um in diameter; IKI-, CB-, tissues unchanged in KOH. Cystidia umbrella-shaped, colorless, thin-walled, smooth, 5.5-10 x 4-6 um; cystidioles absent. Basidia urniform, with a median constriction, thin-walled, with 4 sterigmata and a basal clamp connection, 6-11.5 x 2-4.5 um; basidi- oles abundant, in shape similar to basidia, but slightly smaller. Basidiospores narrowly ellipsoid, colorless, thin-walled, smooth, IKI-, CB-, (2.5-)3-4 x 1-2 um, L = 3.36 pm, W = 1.63 um, Q = 2.06 um (n = 30/1). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 78 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 7. Microscopic structures of Burgella fissurata (holotype): basidiospores (A), ba- sidia and basidioles (B), umbrella-shaped cystidia (C), a section of hymenium (D). Scale bars: 5 um (A); 10 um (B, C). Figure 8. Microscopic structures of Burgella fissurata (holotype): basidiospores (A), ba- sidia and basidioles; umbrella-shaped cystidia (B), generative hyphae (C), a section of hymenium (D). Scale bars: 10 um (A-D). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 79 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Burgoa wumengshanensis Q. Zhou & C.L. Zhao, sp. nov. MycoBank No: 857298 Figs 9-11 Holotype. CHINA * Yunnan Province: Zhaotong, Yiliang County, Luozehe Town, Lijiaping Village, Wumengshan National Nature Reserve, GPS coordinates: 27°29'N, 103°55E, altitude: 1900 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 19 September 2023, CLZhao 33227 (SWFC). Etymology. wumengshanensis (Lat.): refers to the locality, Wumengshan Na- tional Natural Reserve, of the type specimens. Description. Basidiomata annual, resupinate, adnate, pellicular, pruinose upon drying, without odor or taste when fresh, up to 7.4 cm long, 2.1 cm wide, 40-90 um thick. Hymenial surface smooth, white when fresh, turning to pale cream upon drying. Sterile margin thin, white, up to 1 mm wide. Figure 9. Basidiomata of Burgoa wumengshanensis (holotype). Scale bars: 1 cm (A); 1mm (B) MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 80 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 10. Microscopic structures of Burgoa wumengshanensis (holotype) basidio- spores (A), basidia and basidioles (B), schizopapillate cystidia (C), clavate cystidioles (D), a section of hymenium (E). Scale bars: 5 um (A), 10 um (B, E). Hyphal system monomiitic, generative hyphae with clamp connections, some- times with oily contents, colorless, slightly thick-walled, frequently branched, interwoven, 3.5—-5 um in diameter; IKI-, CB-, tissues unchanged in KOH. Cystidia schizopapillate, colorless, thin-walled, smooth, 30.5-49 x 5.5-8 um; cystidioles clavate, colorless, thin-walled, smooth, 22—29.5 x 5-8 um. Basid- ia urniform, with a median constriction, slightly thick-walled, with 8 sterigma- ta and a basal clamp connection, 12-20 x 3.5-6 um; basidioles abundant, in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, colorless, thin-walled, smooth, with oil drop, IKI-, CB-, (3.5)—4-5.5(-—6) x 2-3.5 um, L = 4.5 um, W = 2.78 um, Q = 1.62 (n = 30/1). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 8] Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 11. Microscopic structures of Burgoa wumengshanensis (holotype) basidio- spores (A), basidia (B), clavate cystidioles (C), schizopapillate cystidia (D), a section of hymenium (E), generative hyphae (F, G). Scale bars: 10 um (A-G). Clavuliella Q. Zhou & C.L. Zhao, gen. nov. MycoBank No: 858532 Type species. Clavuliella sinensis Q. Zhou & C.L. Zhao, sp. nov. Etymology. Clavuliella (Lat.): refers to the related genus Clavulina. Description. Basidiomata annual, coralloid, gregarious to caespitose clus- ters, greyish white to light grey when fresh, turning to dark grey upon drying; with sharply acuminate or cristate tips. Hyphal system monomitic, generative hyphae simple-septa, colorless, slightly thick-walled, frequently branched, in- terwoven. Cystidia and cystidioles absent. Basidia cylindrical, with a median constriction, with 2 sterigmata and a basal simple-septum, with oily contents. Basidiospores subglobose to broadly ellipsoid, colorless, thin-walled, smooth, with oil drop, IKI-, CB-. MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 ry) Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Notes. In our phylogenetical analyses (Fig. 1), Clavuliella was identified as a monophyletic group, typified by C. sinensis. The new genus Clavuliella falls within the family Hydnaceae (Cantharellales) and is closely related to Clavulina. Clavulina is distinguished from Clavuliella by its clavarioid to coralloid, simple or branched basidiomata with amphigenous hymenia, cylindrical to subclavate basidia with two or more cornuted sterigmata (Schréter 1888; Uehling et al. 2012; He et al. 2019; Gao et al. 2024). Clavuliella resembles Clavulina in sharing coralloid basidiomata, subglobose, thin-walled basidiospores. However, Clavuliella differs from Clavulina by cylin- drical basidia, with a median constriction, slightly thick-walled, with 2 sterigma- ta and a basal simple-septum, with oily contents and simple-septa generative hyphae. In this study, Clavuliella originating from the subtropical regions, sug- gests the possibility of discovering new corticioid taxa through further investi- gations and molecular analyses. Clavuliella sinensis Q. Zhou & C.L. Zhao, sp. nov. MycoBank No: 858533 Figs 12-14 Holotype. CHINA * Guizhou Province: Guiyang, Qianlingshan Forest Park, GPS coordinates: 26°36'N, 106°41'E, altitude: 1396 m asl., on the ground, leg. C.L. Zhao, 21 August 2023, CLZhao 31231 (SWFC). Etymology. sinensis (Lat.): refers to the type locality (China). Description. Basidiomata annual, coralloid, gregarious to caespitose clus- ters, 0.6-1.5 cm tall, 0.62-1.6 cm wide, frequently branched 2-3 times, form- ing dichotomous branches at the apices, without odor or taste, soft when fresh, becoming brittle upon drying, usually lacking obvious basal mycelium; greyish white to light grey when fresh, turning to dark grey upon drying; with sharply acuminate or cristate tips. Hyphal system monomitic, generative hyphae simple-septa, colorless, slight- ly thick-walled, frequently branched, interwoven, 4-10.5 um in diameter; IKI-, CB-, tissues unchanged in KOH. Cystidia and cystidioles absent. Basidia cylindrical, with a median constric- tion, slightly thick-walled, with 2 sterigmata and a basal simple-septum, with oily contents, 18.5-43 x 6-9 um; basidioles abundant, in shape similar to ba- sidia, but slightly smaller. Basidiospores subglobose to broadly ellipsoid, colorless, thin-walled, smooth, with oil drop, IKI-, CB-, (7-)7.5-9.5(-10) x 6.5-8(-8.5) um, L = 8.52 um, W = 7.51 um, Q = 1.13 um (n = 30/1). Sistotrema sinense Q. Zhou & C.L. Zhao, sp. nov. MycoBank No: 857299 Figs: 15=1:7 Holotype. CHINA * Yunnan Province: Dali, Weishan County, Qinghua Town, GPS coordinates: 25°01'N, 100°22'E, altitude: 2071.6 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 18 October 2022, CLZhao 24876 (SWFC). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 93 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 12. Basidiomata of Clavuliella sinensis (holotype). Scale bars: 1 cm (A); 1 mm (B). Etymology. sinense (Lat.): refers to the type locality (China). Description. Basidiomata annual, resupinate, adnate, soft coriaceous when fresh, becoming coriaceous upon drying, without odor or taste when fresh, up to 11 cm long, 2.5 cm wide, 50-100 um thick. Hymenial surface smooth, white when fresh, turning to white to incanus upon drying. Sterile margin thin, white, thinning out, up to 1 mm wide. Hyphal system monomitic, generative hyphae with clamp connections, often and characteristically with oil content, colorless, slightly thick-walled, frequently branched, interwoven, 2-4 um in diameter; IKI-, CB-, tissues unchanged in KOH. Cystidia and cystidioles absent. Basidia suburniform to urniform, slightly thick-walled, with 4 sterigmata and a basal clamp connection, 8-13.5 x 3-5 um; basidioles abundant, in shape similar to basidia, but slightly smaller. Basidiospores suballantoid to allantoid, colorless, thin-walled, smooth, IKI-, CB-, 3-4.5(—-5) x (1-)1.5-2.5 um L = 3.8 um, W = 1.72 um, Q = 2.21 um (n= 30/1). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 34 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 13. Microscopic structures of Clavuliella sinensis (holotype): basidiospores (A), basidia and basidioles (B), a section of hymenium (C). Scale bars: 5 um (A); 10 um (B, C). Figure 14. Microscopic structures of Clavuliella sinensis (holotype): basidiospores (A), basidia and basidioles (B), a section of hymenium (C), generative hyphae (D). Scale bars: 10 um (A-D). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 85 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Discussion In recent years, many taxa of wood-inhabiting fungi have been continuously re- ported and recorded all over the world, including in the genus Burgella, Burgoa, and Sistotrema (Diederich et al. 2014; Koukol and Kubatova 2015; Cai and Zhao 2023; Dong et al. 2024b; Yuan et al. 2024; Zhang et al. 2024; Yang et al. 2025). Several previous studies, based on ITS+nLSU sequence data, confirmed phylogenetic rela- tionships, in which the genus Burgella, Burgoa, Clavuliella, and Sistotrema are nest- ed in the order Cantharellales (Diederich et al. 2014; Koukol and Kubatova 2015). Phylogenetically, based on the multiple loci in the family Hydnaceae, four gen- era; Burgella, Burgoa, Clavuliella, and Sistotrema were located in this family (Zhou and Qin 2013; Vu et al. 2019; Masumoto and Degawa 2020; Sugawara et al. 2022; MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 86 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Figure 16. Microscopic structures of Sistotrema sinense (holotype): basidiospores (A), basidia and basidioles (B), a section of hymenium (C). Scale bars: 5 ym (A); 10 um (B, C). Figure 17. Microscopic structures of Sistotrema sinense (holotype): basidiospores (A), a section of hymenium (B). Scale bars: 10 um (A, B). MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 97 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae Campo et al. 2023; Cai and Zhao 2023). In the present study, based on the phy- logram inferred from the ITS+nLSU data (Fig. 1), two new species were grouped into the genus Burgella, in which B. albofarinacea was closely related to B. flavo- parmeliae and B. fissurata was grouped with B. /utea. The phylogram based on inferences from the ITS and nLSU data (Fig. 1) showed that B. wumengshanensis clustered into the genus Burgoa, in which it was grouped closely with B. anomala and B. verzuoliana. The phylogram based on inferences from the ITS and nLSU data (Fig. 1) showed that Clavuliella was identified as a monophyletic group, typ- ified by C. sinensis, in which it was grouped closely with C. minor X.X. Huang & L.H. Qiu, C. cristata, and C. iris Loizides, Bellanger & P-A. Moreau. The phyloge- netic tree (Fig. 2), inferred from the ITS+nLSU sequences, highlighted that Sisto- trema sinense was grouped closely with S. brinkmannii and S. farinaceum. Morphologically, Burgella flavoparmeliae, B. lutea and B. fissurata are similar to B. albofarinacea by having the generative hyphae with septa or with clamp connections. (Diederich and Lawrey 2007). However, B. flavoparmeliae differs from B. albofarinacea by the irregularly shaped, coralloid, orange agglomera- tions of bulbils and generative hyphae with septa or without clamp connec- tions, with both morphologies present on the same hyphae at neighboring sep- ta (Diederich and Lawrey 2007). B. /utea is distinguished from B. fissurata by the superficial, yellow to orange-yellow, roundish bulbils (Diederich et al. 2014). B. fissurata is distinguishable from B. albofarinacea by the pruinose hypochnoid hymenial surface, with umbrella-shaped cystidia, basidia with 4 sterigmata, and its smaller basidia (6-11.5 x 2—4.5 um; Diederich and Lawrey 2007; Koukol and Kubatova 2015; Kiyuna et al. 2015). Burgella flavoparmeliae is separated from B. fissurata by the irregularly shaped, coralloid, orange agglomerations of bulbils and generative hyphae with septa or without clamp connections, both situations present on the same hyphae at neighbouring septa (Diederich and Lawrey 2007). B. /utea is distin- guished from B. fissurata by the superficial, yellow to orange-yellow, roundish bulbils (Diederich et al. 2014). B. albofarinacea differs from B. fissurata by the pellicular, coriaceous hymenial surface, bigger basidia with 8 sterigmata and bigger basidiospores (Cai and Zhao 2023). Morphologically, Burgoa anomala and B. verzuoliana are similar to B. wu- mengshanensis by having the generative hyphae with clamp connections (Koukol and Kubdatova 2015). However, B. anomala is distinguishable from B. wumengshanensis by having spherical bulbils, and hyaline to pale brown gener- ative hyphae, thin-walled, thinner (2-5 um) in diameter (Koukol and Kubatova 2015); B. verzuoliana is distinguished from B. wumengshanensis by having spherical bulbils (Diederich and Lawrey 2007). Morphologically, Clavulina cristata, C. griseoviolacea Yue Gao, Hao Zhou, & C.L. Hou, and C. pallida Yue Gao, Hao Zhou & C.L. Hou are similar to Clavuliella sinensis by having clavarioid to coralloid basidiomata and guttulate basidio- spores (Uehling et al. 2012; Gao et al. 2024). However, Clavulina cristata is separated from Clavuliella sinensis by having cylindrical to subclavate basidia with two or more cornuted sterigmata (Gao et al. 2024); Clavulina griseovi- olacea differs from Clavuliella sinensis by having gray to dark grayish violet basidiomata with a white stipe, hyphae with clamp connections, and smaller basidiospores (6.5-8.0 x 6.2—7.2 um; Crous et al. 2014); Clavulina pallida is distinguishable from Clavuliella sinensis by having solitary or scattered basidi- MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 88 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae omata, generative hyphae clamp connections, and longer basidia (34.2—48.5 x 4.8-6.3 um; Gao et al. 2024). Morphologically, Sistotrema diademiferum (Bourdot & Galzin) Donk, S. coroniferum (Hohn. & Litsch.) Donk and S. hispanicum M. Duefias, Ryvarden & Telleria are similar to S. sinense by having the urniform basidia and basal hyphae with clamp connections (Bernicchia and Gorjén 2010). However, S. dia- demiferum is separated from S. sinense by the smooth, porulose hymenophore, larger basidia with 6 sterigmata (15-20 x 5-7 um), and ovoid to subglobose basidiospores (Bernicchia and Gorjon 2010). S. coroniferum is distinguishable from S. sinense by the smooth hymenophore, with gloeocystidia, basidia with 6 sterigmata, and longer subcylindrical basidiospores (5-6 x 2—2.5 um; Bernic- chia and Gorjon 2010). S. hispanicum differs from S. sinense by the whitish to yellow hymenial surface and bigger narrowly ellipsoid to subreniform basidio- spores (5.5—-6 x 3-4 um; Bernicchia and Gorjén 2010). As wood-inhabiting fungi efficiently degrade lignocellulose in wood, they play a crucial ecological role in material recycling and energy flow in forest ecosys- tems, as well as playing a major economic role (Sugawara et al. 2022; Zhang et al. 2022; Bondartseva and Zmitrovich 2023; Campo et al. 2023; Cui et al. 2019; Gao et al. 2024; Liu et al. 2023a, b; Sun et al. 2020; Sun et al. 2022; Ji et al. 2022). Wood-inhabiting fungi are an extensively studied group of Basidiomycota, but their diversity is still not well known in China, and many recently described taxa in this ecological group have been discovered from China (Sugawara et al. 2022; Cai and Zhao 2023; Wang et al. 2023; Wang et al. 2024; Wu et al. 2022; Yuan et al. 2023; Zhao et al. 2024). Four new species and a new genus, from the Yunnan and Guizhou Provinces of China, serve as examples of the understudied fungal diversity present in the PR. of China.. On a wider scale, this study enriches our knowledge on the diversity of wood-inhabiting fungi worldwide. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding The research was supported by the National Natural Science Foundation of China (Proj- ect Nos. 32170004, U2102220) and Forestry and Grass Science and Technology Inno- vation Joint Project of Yunnan Province (Project No. 202404CB090008), the High-lev- el Talents Program of Yunnan Province (YNQR-QNRC-2018-111), Forestry Innovation Programs of Southwest Forestry University (Grant No: LXXK-2023Z07), and the Yunnan Province College Students Innovation and Entrepreneurship Training Program (Project No. S202410677089). Author contributions Conceptualization: CZ. Methodology: CZ and QZ. Software: CZ, QZ, and HZ. Validation: CZ and QZ. Formal analysis: CZ and QZ. Investigation: CZ, CQ, CZ, QS, SZ, NM, TX, and QZ. Resources: CZ. Writing — original draft preparation: CZ, CQ, CZ, QS, YL, SZ, NM, TX, MycoKeys 119: 67-94 (2025), DOI: 10.3897/mycokeys.119.154387 89 Qian Zhou et al.: Morphology and multigene phylogeny revealed five species of Hydnaceae HZ, and QZ. Writing — review and editing: CZ and QZ. Visualization: CZ and QZ. Super- vision: CZ and QZ. Project administration: CZ. Funding acquisition: CZ. All authors have read and agreed to the published version of the manuscript. Author ORCIDs Qian Zhou ® https://orcid.org/0009-0007-0845-2115 Chengbin Qian © https://orcid.org/0009-0003-5329-1016 Chuyun Zhang ® https://orcid.org/0009-0008-8239-8492 Qidong Su © https://orcid.org/0009-0000-1 521-7606 Yiliang Li © https://orcid.org/0009-0009-9599-256X Shihui Zhang ® https://orcid.org/0009-0008-3309-4214 Nian Mu © https://orcid.org/0009-0009-1316-8658 Taimin Xu © https://orcid.org/0000-0002-5230-4140 Hongmin Zhou © https://orcid.org/0000-0002-0724-5815 Changlin Zhao © https://orcid.org/0000-0002-8668-1075 Data availability All of the data that support the findings of this study are available in the main text. References Bernicchia A, Gorjon SP (2010) Fungi Europaei 12: Corticiaceae s.|. Edizioni Candusso, Alassio, Italy, 1008 pp. Binder M, Hibbett DS (2002) Higher-level phylogenetic relationships of Homobasidiomy- cetes (mushroom-forming fungi) inferred from four rDNA regions. 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