<8) MycoKeys MycoKeys 120: 157-172 (2025) DOI: 10.3897/mycokeys.120.138950 Research Article Two new species of Phaeoclavulina (Gomphaceae, Gomphales) from North China based on morphological and phylogenetic analysis Xin Tong™®, Yue Gao2™®, Hai-Qi Wang2®, Hao Zhou2®, Cheng-Lin Hou2® 1 Department of Life Sciences, Natural History Museum of China, Tiangiaonandajie 126, Dongcheng, 100050, Beijing, China 2 College of Life Science, Capital Normal University, Haidian, 100048, Beijing, China Corresponding author: Cheng-Lin Hou (chenglin-hou@cnu.edu.cn) OPEN Qrceess Academic editor: Thorsten Lumbsch Received: 10 October 2024 Accepted: 2 July 2025 Published: 29 July 2025 Citation: Tong X, Gao Y, Wang H-Q, Zhou H, Hou C-L (2025) Two new species of Phaeoclavulina (Gomphaceae, Gomphales) from North China based on morphological and phylogenetic analysis. Mycokeys 120: 157-172. https://doi. org/10.3897/mycokeys. 120.138950 Copyright: © Xin Tong 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 Two new species, Phaeoclavulina aurea and P. fulva, were discovered and described from North China. Identification was based on morphological observations combined with phylogenetic analysis of nrlTS-nrLSU sequences. Phaeoclavulina aurea is charac- terized by its pale yellow to golden yellow basidiomata, brownish black coloration with dark tips at maturity, and basidiospores bearing truncate spines. Phaeoclavulina fulva is distinguished by its dirty orange basidiomata, pale yellow stipe surface, and basid- iospores ornamented with rounded warts. This study enriches the species diversity of Phaeoclavulina in North China. Key words: Clavarioid fungi, new taxa, systematics, taxonomy Introduction Phaeoclavulina Brinkmann (1897: 197) (Gomphaceae, Gomphales), with P macrospora Brinkmann (1897: 198) as the type species, was established in 1897 (Brinkmann 1897). It includes gomphoid and ramarioid forms character- ized by ramarioid, unipileate, or merismatoid basidiomata. When pileate, ba- sidiomata are glabrous or subtomentose and infundibuliform or flabelliform (Gonzalez-Avila et al. 2020). The genus exhibits a diverse palette of colors on the pileus and branch surfaces, varying from white, brown-green, pale to sordid olivaceous, violet, brown-yellow, and red cinnamon, through to gray, brick red, and pale to dark orange-yellow, even blue-green. Spores show the presence of clamps and are echinulate to verrucose, subreticulate, or reticulate (Giachini 2004; Giachini and Castellano 2011). Some Phaeoclavulina species are ecto- mycorrhizal. For example, P. abietina (Pers.) Giachini (2011: 189) is associated with Betula, Pinus, and Pseudotsuga (Herrera et al. 2002; Norvell and Exeter 2004); P. flaccida (Fr.) Giachini (2011: 192) with Pinus and Quercus (Kim et al. 2003); P. zippelii (Lév.) Overeem (1923: 262) with Acacia, Casuarina, and Eu- calyptus (Sims et al. 1997); and P. cyanocephala (Berk. & M.A. Curtis) Giachini (2011: 191) with Abies (Estrada-Torres 1994). * These authors contributed equally to this work. Tae Xin Tong et al.: New species of Phaeoclavulina from China Initially, Brinkmann (1897) placed Phaeoclavulina within the family Clavariace- ae, along with other genera, viz. Clavaria Vaill. ex L. (1753: 1182), Clavariella P. Karst. (1881: 21), Clavulina J. Schrét. (1888: 442), and Typhula (Pers.) Fr. (1818: 296). Overeem (1923) recognized Phaeoclavulina as a distinct genus and assigned one species, P zippelii, to it, which had previously been classified under Clavaria based on macromorphological characteristics. Despite this, the recognition of Phaeoclavulina did not gain widespread support among mycologists, and species of this genus were often accommodated with other ramarioid fungi having ochre spores in Ramaria Fr. ex Bonord. (1851: 166) (Gonzalez-Avila et al. 2020). Corner (1970) went further by placing the ramarioid species of Phaeoclavulina within Ra- maria subgen. Echinoramaria Corner (1970: 238), a classification later adopted by other authors such as Marr and Stuntz (1973) and Petersen (1981). In recent research, Phaeoclavulina was once again recognized as a valid genus based on morphological, molecular, and phylogenetic data (Giachini 2004; Giachini et al. 2010; Giachini and Castellano 2011). It now includes not only the original species but also some gomphoid species with spiny, verrucose, subreticulate, or reticulate spores and a terrestrial and/or lignicolous substrate affinity (Gonzdlez-Avila et al. 2020). Currently, approximately 57 species of Phaeoclavulina have been de- scribed from temperate and tropical ecosystems; however, it is likely more abun- dant in the tropics and subtropics (Gonzalez-Avila et al. 2020; Liu et al. 2022). In China, 22 Phaeoclavulina species have been reported based on morpho- logical features and phylogenetic analyses, viz. P abietina, P aeruginea P. Zhang (2022: 31), P bicolor P. Zhang & W.H. Liu (2024: 6), P campestris (K. Yokoy. & Sagara) Giachini (2011: 190), P capucina (Pat.) Giachini (2011: 190), P. cinna- momea W.Q. Qin (2022: 32), P cokeri (R.H. Petersen) Giachini (2011: 190), P. cur- ta (Fr.) Giachini (2011: 190), P cyanocephala, P. decolor (Berk. & M.A. Curtis) Giachini (2011: 191), P echinoflava P. Zhang & W.H. Liu (2024: 7), RP eumorpha (P. Karst.) Giachini (2011: 191), P. flaccida, P grandis (Corner) Giachini (2011: 193), P jilinensis P. Zhang & W.H. Liu (2024: 9), P longicaulis (Peck) Giachini (2011: 193), P macrospora, P mutabilis (Schild & R.H. Petersen) Giachini (2011: 194), P. sikkimia (S.S. Rattan & Khurana) Giachini (2011: 194), P. viridis (Pat.) Giachini (2011: 195), P yunnanensis W.H. Lu, D.G. Zheng, Karun. & Tibpromma (2024: 113), and P zippelii (Liu et al. 2022; Deng et al. 2024; Zheng et al. 2024). Despite numerous prior reports from China, there remains significant potential for the discovery of new species, particularly in North China. Recently, several Phaeoclavulina-like samples were collected during an inves- tigation of the Yanshan Mountains (39°40'-41°20' N, 115°-119°47' E) in North China, a warm temperate region. In this study, two new species are described and illustrated. The nuclear ribosomal internal transcribed spacer (nrITS) and the large subunit of nuclear ribosomal RNA (nrLSU) were sequenced from dried basidiomata of each species for phylogenetic analysis. Materials and methods Collecting and site description The specimens were collected from Beijing and Tianjin, North China, between 2019 and 2023, and important collection data were recorded (Rathnayaka et al. 2024). These regions have a warm temperate continental monsoon climate, MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 158 Xin Tong et al.: New species of Phaeoclavulina from China which supports a diverse assemblage of plant species (Zhou et al. 2022). De- ciduous broad-leaved forest and mixed coniferous and broad-leaved forest are the dominant vegetation types in the area. Notable plant species include Abies nephrolepis (Trautv.) Maxim., Betula platyphylla Suk., Pinus tabuliformis Carr., Populus tomentosa Carriére, and Quercus mongolica Fisch. ex Ledeb. (Wang et al. 2021; Zhou et al. 2022). The annual precipitation is approximately 700 mm (Zhou et al. 2022), and the elevation ranges from 200 to 2200 m. The collected specimens were dehydrated using an electric dryer (Dorrex) at 50 °C and then deposited in the Herbarium of the College of Life Sciences, Capital Normal Uni- versity, Beijing, China (BUTC), and the Herbarium of the Natural History Muse- um of China, Beijing, China (NNHMC). Morphological observation Macroscopic characteristics of the specimens were recorded, including basid- iomata color, size, branching pattern, stipe color, morphology, and dimensions. Microscopic features were analyzed by examining thin sections mounted in 3% potassium hydroxide (KOH) or sterilized water. The morphology and dimensions of microscopic structures were observed and recorded using a light microscope (Olympus DP71, Tokyo, Japan). In the description of basidiospores, the abbrevi- ation n/m/p indicates that n basidiospores were measured from m basidiomata of p collections. Measurements and Q values are presented in the format (a) b-c(d), where “a” represents the minimum value, “b-c” the 10% to 90% range, and “d” the maximum value. Q represents the ratio of basidiospore length to width in side view (Liu et al. 2022), and Q_ represents the average Q value of all measured basidiospores + the sample standard deviation (Liu et al. 2022). Nomenclatural details were submitted to MycoBank. Color terms followed the designations provided by the website ColorHexa (https://www.colorhexa.com). DNA extraction, PCR amplification, and sequencing DNA extraction was carried out using the M5 Plant Genomic DNA Kit (Mei5 Bio- technology Co., Ltd., China). The extracted DNA was dissolved in 1x TE buffer and stored at —20 °C for later use. PCRs were performed using a Bio-Rad S1000 thermal cycler (Bio-Rad Laboratories, Inc., USA). The primer set ITS5/ITS4 (Vil- galys and Hester 1990; White et al. 1990) was used to amplify the nrlTS region, and LROR/LR&5 (Vilgalys and Hester 1990) was used for the nrLSU region. PCRs were conducted in a 25 uL reaction volume containing 2 uL of DNA template, 1 uL of each primer (10 uM), 12.5 uL of 2x Master Mix (Mei5 Biotechnology Co., Ltd., China), and 8 uL of ddH,O. PCR amplification conditions for nrlTS fol- lowed Wannathes et al. (2018), Liu et al. (2022), and Gao et al. (2024), and the conditions for nrLSU followed Yan et al. (2020) and Sui et al. (2023). All DNA sequences were generated by Sangon Biotech (Shanghai) Co., Ltd. Molecular phylogenetic analyses The newly obtained sequences were submitted to NCBI (https://www.ncbi. nlm.nih.gov). The nrlTS and nrLSU sequences were aligned with selected se- quences from GenBank and previously published literature (Deng et al. 2024). MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 159 Xin Tong et al.: New species of Phaeoclavulina from China All sequences used are listed in Table 1. The raw reads of the DNA sequenc- es were processed to obtain consensus sequences using SeqMan 7.1.0 in the DNASTAR Lasergene Core Suite software (DNASTAR Inc., Madison, WI, USA). Sequence alignment was performed using MAFFT 6 (Katoh and Toh 2010), and manual trimming was carried out in MEGA 6 (Tamura et al. 2013). For phyloge- netic analyses, newly obtained sequences and additional reference sequences of Phaeoclavulina and Ramaria species were included in the nrlTS and nrLSU dataset (Table 1), with Gomphus clavatus (Pers.) Gray designated as the out- group following Wannathes et al. (2018). To estimate maximum likelihood (ML) gene trees, RAXML 7.4.2 Black Box software was used (Stamatakis 2006; Stamatakis et al. 2008; Zhou and Hou 2019; Zhou et al. 2021), employing a GTRGAMMAI site substitution model (Guindon et al. 2010). Branch support was assessed with 1000 bootstrap (BS) replicates (Hillis and Bull 1993). Bayesian Inference (BI) analysis was conduct- ed using MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003), with a Markov chain Monte Carlo (MCMC) algorithm (Rannala and Yang 1996). The best-fit substitu- tion model was determined using MrModeltest 2.3 (Zhou and Hou 2019; Zhou et al. 2021, 2022). The selected models were SYM + G for nrlTS and SYM+1+G for nrLSU. Two MCMC chains were run for 10,000,000 generations, terminating when the average standard deviation of split frequencies fell below 0.01. Trees were sampled every 1000 generations, with the first 25% discarded as burn-in. Significant Bayesian posterior probabilities (pp) were calculated for branches in the resulting majority-rule consensus trees. The analyses yielded relatively stable topologies, and clades with high pp values reflected the phylogenetic relationships among species (Posada and Crandall 1998). Results Molecular phylogeny A total of nine sequences, including five for nrlTS and four for nrLSU, were new- ly generated in this study. The nrlTS and nrLSU datasets were compiled to in- vestigate the phylogenetic position of the new species in Phaeoclavulina. The combined nrlTS—-nrLSU dataset included 146 sequences (69 for nrlTS and 77 for nrLSU), representing 102 samples. The concatenated alignment contained 840 characters, including gaps. Maximum Likelihood (ML) and Bayesian Infer- ence (BI) analyses yielded highly similar tree topologies; therefore, only the tree inferred from the ML analysis is shown (Fig. 1). The tree topology is consistent with that of Zheng et al. (2024). In the phy- logenetic reconstruction, each of the two new species, P aurea and P fulva, formed a distinct monophyletic lineage within the Phaeoclavulina clade (Fig. 1). The specimens of P aurea (BUM 344955 and BUTC L007) and P fulva (BUTC C274 and BUTC ZH1138) were identified as the sister species of P yunnanensis, with strong statistical support (MLB = 100%, BPP = 1.00). Phaeoclavulina aurea (BJM 344955 and BJTC L007) formed a clade with P minutispora and P. decur- rens, although this relationship was supported by a relatively lower value (MLB = 85%, BPP = 0.83). In conclusion, the ML and BI analyses of the nrlTS—nrLSU dataset support the recognition of two new species of Phaeoclavulina from China, viz. P aurea and P fulva. MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 160 Xin Tong et al.: New species of Phaeoclavulina from China Table 1. Specimens used in phylogenetic analysis and their GenBank accession numbers. Newly generated sequences are shown in bold. Voucher numbers marked with “T” indicate holotypes. GenBank Number Taxonomy Voucher/strain Location References P carovinacea Franchi et al. (2022) Pes P. clavarioides Kiiz et al. (2019) LR723645 MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 KX008988 EU118628 KY510818 JX310378 JX310379 ON262784 ON262785 NR_176722 MT055964 MT055916 JQ408231 JQ408234 PQ287856 PQ287853 PP809798 PP809799 NR_176719 MT055918 NR_177141 MT055920 ON262786 0Q703628 LR723646 MH322666 NR_176718 AJ408359 KX236126 KT339249 0Q586352 PP809801 PP809800 OM152300 AJ408390 AJ408371 MT055926 MW115424 MW115426 PQ287852 161 Xin Tong et al.: New species of Phaeoclavulina from China Taxonomy P. fulva P fulva P gigantea P grandis P. guyanensis P insignis P jilinensis P jilinensis P jilinensis P liliputiana P liliputiana P liliputiana P liliputiana P longicaulis P macrospora P. minutispora P. minutispora P murrillii P mutabilis P myceliosa P. nigricans P ochracea P. ochracea P ochraceovirens P pancaribbea P. pseudozippelii P pseudozippelii P quercusilicis P roellinii P. subclaviformis P. subdecurrens P tropicalis P yunnanensis P yunnanensis P zealandica P zippelii Ramaria admiratia R. admiratia R. aurantiisiccescens R. aurantiisiccescens R. botrytis R. botrytis R. calvodistalis R. calvodistalis R. cf. celerivirescens R. foetida R. gracilis R. gracilis R. largentii R. largentii R. luteovernalis R. luteovernalis Voucher/strain References GenBank Number ~eirezHoois china —=SSSCresent study wHNUTOSD8 (7) Deng et al. (2024) PP800481 3281 Gonzalez et al. (2020) MT214488 3533 Gonzalez et al. (2020) MT214489 5266 (1) Gonzélez et al. (2020) MT214490 awe i961 tay | Foret (202) ~aieasa82 Spain| ‘Marnetak 020) AvTos70F828646 | Conodo | Unpublnhed “waungia7994—Spein | Unpubnhed PRM:945446 Czech Republic Kifz et al. (2019) pro7siss07 | «ich GoTo) STAG posses |SSCUnptbohed |S TTO re 1aas0(T) USA Petersen ove) TENN 005) USA | Peterseneta ore) neko72 USA) Urptbnhed wove 20652 tay Fanehieta @015) ~wove%e6s7 (7) | tay =| SCanehiet al (205) MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 nrITS PQ287854 PQ287855 PP809802 PP809803 PP809804 MT452510 0Q749903 0Q729761 MH322683 MN992499 MT055970 MT055924 MG214660 MG214661 AJ408382 LR723648 MT055930 0Q755411 0Q755412 KJ416133 EU669247 JX310388 NR_189799 OP177708 KJ416132 JQ408243 JX310399 KY354745 KP658130 KT357476 KT357471 162 Xin Tong et al.: New species of Phaeoclavulina from China 100/1,00) P2aeoclavulina yunnanensis HKAS 127150 China Phaeoclavulina yunnane: 100/100) : Phaeoclavulina fulva | 98/0.97| Phaeoclavulina fulva BJTC 33/- Phaeoclavulina fulva BJTC 00/1.00) P2aeoclavulina carovinacea AMB 18533 Italy (T) Phaeoclavulina carovinacea AMB 18534 Italy Phaeoclavulina nigricans AMB 18589 Italy 99/1.00 97/1.00 is HKAS 128154 China Pens Phaeoclavulina abietina OSC 140661 USA Phaeoclavulina abietina OSC 112178 USA 100/1.00} Phaeoclavulina abietina OSC 134649 USA 9410.97 Phaeoclavulina abietina U 066 USA Phaeoclavulina minutispora GT21030 Belgium 100/1.00| Phaeoclavulina minutispora LD5028 Belgium 100/1.00} 8b/- Phaeoclavulina decurrens 257517179 Netherlands 100/1.00) Phaeoclavulina aurea BJTC L007 China Phaeoclavulina aurea BJM 344955 China 100/1.00) P2aeoclavulina alboapiculata AMB 18590 Italy (T) 97/- Phaeoclavulina alboapiculata AMB 185835 Italy 100/--— Phaeoclavulina curta MA-Fungi 48081 Spain Phaeoclavulina curta UBC F32034 Canada Phaeoclavulina flaccida MA-Fungi 48020 Spain Phaeoclavulina flaccida AMB 18544 Italy Phaeoclavulina flaccida MA-Fungi 48076 Spain Phaeoclavulina flaccida AMB 18645 Italy Phaeoclavulina flaccida AMB 18643 Italy Phaeoclavulina ochracea AMB 18542 Italy 100/1.00} 99/1.00 98/0.96 Phaeoclavulina jilinensis MHHNU9149 China 100/1.00! Phaeoclavulina jilinensis MHHNU9164 China 100/1.00} 100/-— Phaeoclavulina coniferarum AMB 18531 Italy (T) P7/1.00) Phaeoclavulina macrospora AMB 18614 Italy Phaeoclavulina ochraceovirens OSC23475 USA Phaeoclavulina subdecurrens AMB 18548 Italy Phaeoclavulina roellinii PRM:945446 Czech Republic Phaeoclavulina jilinensis MHHNU10504 China (T) Phaeoclavulina viridis OSC97708 USA 99/- Phaeoclavulina echinoflava HKAS 45984 China (T) 100/1.00} Phaeoclavulina echinoflava_HKAS 45992 China Phaeoclavulina murrillii AH:48382 Spain Phaeoclavulina cokeri TENN36030 USA Phaeoclavulina arcosuensis AMB 18532 Italy 100/1.00) Phaeoclavulina pseudozippelii BBH 43576 Thailand 97/- 98/1.00 97/0.99 Phaeoclavulina pseudozippelii BBH 43575 Thailand (T) Phaeoclavulina cyanocephala TENN 37827 USA 100/1.00) P2aeoclavulina aeruginea MHHNU6887 China Phaeoclavulina aeruginea MHHNU8909 China (T) Phaeoclavulina cyanocephala TH 9064 Guyana Phaeoclavulina apiahyna LPS13259 USA Phaeoclavulina gigantea FH109 USA 100/1.00 Phaeoclavulina insignis FH104 USA 82/- Phaeoclavulina africana TENN39621 USA Phaeoclavulina cinnamomea MHHNU10376 China (T) 100/-) Phaeoclavulina eumorpha GLM:GLM-F 116666 Germany 86/- Phaeoclavulina myceliosa ANT057-QFB28646 Canada 100/1.00r Phaeoclavulina caroviridula AMB 18535 Italy (T) Phaeoclavulina caroviridula AMB 18536 Italy MT055920 Phaeoclavulina grandis BR079158-06 USA Phaeoclavulina clavarioides ERD:9641 Portugal 100/1.00] Phaeoclavulina clavarioides PRM-945440 Czech Republic 96/- Phaeoclavulina quercusilicis MA-Fungi 47984 Spain 95/- Phaeoclavulina argentea AGK 036 USA Phaeoclavulina argentea AGK 042 USA Phaeoclavulina guyanensis FH84 USA 100/1.00) P2aeoclavulina bicolor MHHNU10702 China (T) 100/1.00) Phaeoclavulina bicolor MHHNU10703 China 91/- Phaeoclavulina longicaulis TENN33826 USA Phaeoclavulina pancaribbea TENN31836 USA Phaeoclavulina liliputiana 3281 Mexico Phaeoclavulina liliputiana 3563 Mexico 100/0.95) Phaeoclavulina liliputiana 3533 Mexico 97/- Phaeoclavulina liliputiana 3266 Mexico (T) 100/0.95,- Phaeoclavulina corrugata SJ99002 89/- Phaeoclavulina curta OSC8711 USA Phaeoclavulina cokeri MA:Fungi:79893 Spain 88/-) Phaeoclavulina tropicalis NY551 USA 82/4 Phaeoclavulina decolor FH1 USA Phaeoclavulina capucina GH 288 USA Phaeoclavulina mutabilis TENN3 9893 USA 97/0.99 99/- Phaeoclavulina campoi LPS39622 USA 96/-) Phaeoclavulina angustata BPI2 Phaeoclavulina zealandica PDD43383 85/- Phaeoclavulina zippelii FH2 USA Phaeoclavulina ochracea 81 USA 100/1.00) Ramaria gracilis OSC 134659 USA Ramaria gracilis OSC 112168 USA 78/- 100/- Ramaria aurantiisiccescens OSC 65703 USA 98/-| | Ramaria aurantiisiccescens OSC 130871 USA 99/1.00 Ramaria calvodistalis TENN 69095 USA (T) 100/- Ramaria calvodistalis TENN 69095 USA Ramaria foetida AGK 058 USA Ramaria admiratia TENN 69114 USA 92/- 99/1.007— Ramaria largentii OSC 67012 USA 100/1.00 Ramaria largentii OSC 140726 USA 100/1.00) Ramaria luteovernalis MCVE 28662 Italy 91/ Ramaria luteovernalis MCVE 28637 Italy (T) Ramaria admiratia TFB14450 USA (T) Ramaria cf. celerivirescens AGK 072 USA Phaeoclavulina subclaviformis BRO79159-07 Ramaria botrytis AMB 18201 (T) Ramaria botrytis GM19124 Argentina 92/--— Gomphus clavatus EL 64/03 Sweden Gomphus clavatus GO71 USA Gomphus clavatus LL 115 China 90/- 99/0.99 0.04 Figure 1. Phylogenetic tree generated from an ML analysis based on nrlTS-nrLSU sequences. Numbers representing maximum likelihood bootstrap support (MLBS = 75%, left) and significant Bayesian posterior probability (BPP = 0.95, right) are indicated above the nodes. Novel sequences are printed in bold. Voucher specimens and localities where the specimens were collected are provided behind the species names. MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 163 Xin Tong et al.: New species of Phaeoclavulina from China Taxonomy Phaeoclavulina aurea Y. Gao, X. Tong, & C.L. Hou, sp. nov. MycoBank No: 855628 Figs 2A-C, 3 Diagnosis. Phaeoclavulina aurea differs from the known species of Phaeoclavu- lina in its pale yellow to golden yellow basidiomata, tips brownish black when age, truncate spines basidiospores 4.7-6.3 x 3.0-3.7 um, basidia 25-50 x 4.6-7.4 um, clamp connections present. Etymology. The epithet “aurea” refers to the yellow to golden yellow basidiomata. Type. CHINA * Beijing, Yanging District, Songshan, 40°30'28'N, 115°54'E, elev. 869 m, 16 Aug. 2023, H. Zhou, Y. Gao & X. Tong (BUM 344955). GenBank nrlTS: PQ287856, nrLSU: PQ287860. Description. Basidiomata coralloid, solitary or scattered; individual basidi- omata 3-8 cm tall, 2—5.5 cm wide across branches. Stipe 0.4—0.9 cm tall, 0.3- 0.5 cm wide, subclavate to flattened, snow-white rhizomorphic strands near the ground, branches di- or tri-dichotomous, generally 3-4 times, cylindrical, pale yellow (#ffdd9a) to golden yellow (#ffc44d); as maturity deepens, tips blunt and short with brownish black (#473826), stipe surface white (#ffffff) to pale yellow (#ffdd9a). No color change when bruised. Odor and taste not recorded. Basidiospores [70/2/2] (4.4-)4.7-6.3(- 6.7) x3.0-3.7(- 4.2) um, Q = 1.4- 1.9(- 2.1), Q_ = 1.66 + 0.16, ellipsoid, pale yellow to golden yellow in KOH, with thick wall and cyanophilic ornamentation in cotton blue; when seen with SEM, conical, truncate spines up to 0.6 um high; with oleiferous guttule contents. Hi- lar appendixes acuminate (up to 1.1 umin length). Basidia 25-50 x 4.6—7.4 um, clavate, 3-4 sterigmata occur per basidium, 3.6-6.0 um long, and cornute, clamped. Basidioles abundant, subclavate to subcylindrical. Tramal hyphae in the stipe smooth, thin-walled, hyaline in KOH, 2.3-—5.8 um wide; tramal hyphae in branch with hyaline and thin-walled, 1.8-4.0 um wide; clamp connections abundant, H-connections present. Hyphal system monomitic. Cystidia absent. Habit, habitat, and distribution. Solitary or gregarious caespitose in humus lay- ers on soils in mixed coniferous and broad-leaved forests, associated with Pinus L. Basidiomata generally occur in August; currently known only from Beijing, China. Additional specimens examined. CHINA * Beijing, Huairou District, Hongluo Temple, 40°22'26"N, 116°37'26'E, elev. 153 m, 13 Aug. 2019, J.Q. Li, X.Y. Shen & R.T. Zhang (BUTC L007). Notes. Phaeoclavulina aurea is morphologically similar to P cinnamomea, which was originally described from China. However, P cinnamomea produced cinnamon-to-salmon-orange basidiomata and relatively large basidiospores (12- 15 x 5-7 um) and basidia (40-60 x 8-12 um) compared P aurea (Liu et al. 2022). Phaeoclavulina angustata (Lév.) Giachini (2011: 189) also has crowded branch- es, but the color of that is pale pink, and all parts soon turned sordid vinaceous on bruising, and the cyanophilic spines of basidiospores 1-3.5 um tall (Giachi- ni 2004). Phaeoclavulina glaucoaromatica (R.H. Petersen) Giachini (2011: 193) has similar basidiomata with P aurea, which doesn't have molecular data. But P glaucoaromatica differs from P aurea by its branches becoming watery where handled and then changing color to olive-green; basidiospores are larger (8.2- 11.1 x 4.4—5.5 um) (Petersen 1981). Phaeoclavulina vinaceipes (Schild) Giachini MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 164 Xin Tong et al.: New species of Phaeoclavulina from China Figure 2. Morphology of Basidiomata. A-C. Phaeoclavulina aurea (A, B. BUM 344955 holotype; C. BUTC L007); D-F. Phaeoclavulina fulva (D. BJTC C274 holotype; E. BITC ZH0015; F. BITC ZH1138). Scale bars: 1 cm. Figure 3. Microscopic characteristics of Phaeoclavulina aurea: A. Basidiospores; B. Basidia; C. Tramal hyphae. Scale bars: 20 um (A); 10 um (B, C). MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 165 Xin Tong et al.: New species of Phaeoclavulina from China (2011: 195) has the same branch color as P. aurea, which also doesn't have mo- lecular data. But P. vinaceipes’s stipe context bruising violet when exposed or handled, per basidium occur 4 sterigmata, and basidiospores are wider (4.8-8 x 2.7-3.8 um) (Schild 1990). Phaeoclavulina aurea is phylogenetically close to P. abietina, P alboapiculata Franchi & M. Marchetti (2020: 1), and P: minutispo- ra Franchi & M. Marchetti (2020: 3) (Fig. 1), but the stipes of P abietina quickly turned deep blue-green when handled or confined (Gonzalez-Avila et al. 2020); the branches of P alboapiculata are with white tips when young, and the white context gradually turns to wine when exposed to the air, and it has smaller basidiospores (4.1-5.5 x 2.5-3.4 um) and basidia (22-30 x 5.0-6.5 um) than P aurea (Index Fungorum 2024); P minutispora can be distinguished by its basidiomata surface, which gradually shows dark brown pigmentation when touched, white flesh turns maroon rapidly when exposed to air, and producing relatively small basidiospores (3.7-5.0 x 2.5-3.6 um) and basidia (22-30 x 5-6.5 um) (Index Fungorum 2024). Phaeoclavulina fulva Y. Gao, X. Tong, & C.L. Hou, sp. nov. MycoBank No: 855499 Figs 2D-F, 4 Diagnosis. Phaeoclavulina fulva differs from the known species of Phaeoclavu- lina in its dirty orange basidiomata, stipe surface pale yellow, color lightens to- wards tips, rounded warts basidiospores 4.8-7.8 x2.8-3.8 tm, basidia 22-43 x 4.3-7.3 um. Etymology. The epithet “fu/vus” refers to the dirty orange to yellowish-brown basidiomata. Figure 4. Microscopic characteristics of Phaeoclavulina fulva: A. Basidiospores; B. Basidia; C. Tramal hyphae. Scale bars: 20 um (A); 10 um (B, C). MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 166 Xin Tong et al.: New species of Phaeoclavulina from China Type. CHINA * Tianjin, Jizhou District, Limutai Scenic Spot, 40°12'47'N, 117°2'7'E, elev. 658 m, 21 Aug. 2020, G.Q. Cheng, R.T. Zhang & C.L. Hou (BUTC C274). GenBank nrITS: PQ287852, nrLSU: PQ287857. Description. Basidiomata coralloid, solitary or scattered; individual basidi- omata 4.2-12.5 cm tall, 4.5-10 cm wide across branches. Stipe 1.2-3.8 cm tall, 0.3-0.5 cm wide, subclavate to flattened, snow-white rhizomorphic strands near the ground, branches dichotomous, generally 3-4 times, subcylindrical, dirty orange (#d5995b) to yellowish-brown (#a27f16), tips slightly acute, stipe surface pale yellow (#fff7dc), color lightens towards tips. No color change when bruised. Odor and taste not recorded. Basidiospores [90/4/3] (4.3-)4.8-7.8 x (2.5-)2.8-3.8(- 4.2) um, Q = 1.5- 2.2(- 2.37), Q, = 1.89 + 0.19, elongate-ellipsoid, pale yellow to golden yellow in KOH, with thick wall and cyanophilic ornamentation in cotton blue; when seen with SEM, conical, truncate warts up to 0.6 um high; with oleiferous guttule con- tents. Hilar appendixes acuminate (up to 1.0 um in length). Basidia 22-43 x 4.3- 7.3 um, clavate, 2-4 sterigmata occur per basidium, 1.8-7.0 um long, and corn- ute, clamped. Basidioles abundant, subclavate to subcylindrical. Tramal hyphae in the stipe smooth, thin-walled, hyaline in KOH, 2.2—7.5 um wide; tramal hyphae in branch with hyaline and thin-walled, 2.5-6.5 um wide; clamp connections abundant, H-connections present. Hyphal system monomitic. Cystidia absent. Habit, habitat, and distribution. Solitary or gregarious caespitose in humus layers on soils in evergreen coniferous forests and deciduous broad-leaved for- ests, associated with Pinus L. and Populus L. Basidiomata generally occur from August; currently known only from Tianjin and Beijing, China. Additional specimens examined. CHINA * Beijing, Huairou District, Hongluo Temple, 40°22'34"N, 116°37'28"E, elev. 109 m, 13 Aug. 2019, H. Zhou, G.Q. Cheng & C.L. Hou (BUTC ZH0015); CHINA * Beijing, Huairou District, Erdaogou, 40°52'36'N, 116°31'23"E, elev. 780 m, 24 Aug. 2020, H. Zhou, X.Y. Shen & X.B. Huang (BUTC ZH1138). Notes. Phaeoclavulina fulva generally shares similar branching with P. sik- kimia, P. cokeri, and P. articulotela (R.H. Petersen) Giachini (2011: 190). Phaeo- clavulina sikkimia and P articulotela lack molecular data (Petersen 1981). How- ever, P. sikkimia differs from P fulva in its relatively large basidiospores (6.5—9 x 4.5-6 um) and basidia (60-70 x 7.5—9 um) and in the presence of cyanophilic warts up to 2 um tall (Giachini 2004). P cokeri has cinnamon- to pink-yellow branches that become orange or dark red-brown, with pale orange tips; it also features strongly cyanophilic basidiospore spines 1-2 um tall and relatively large basidiospores (9-16 x 4—7.5 um) and basidia (45-80 x 7.5-12 um) (Gia- chini 2004). P articulotela has orange to dark orange basidiomata and relatively large basidiospores (6—9.5 x 3-5 um) (Giachini 2004). Phylogenetically, P fulva is related to P. abietina, P carovinacea Franchi & M. Marchetti (2020: 2), P. nigri- cans E. Campo, Franchi & M. Marchetti (2020: 4), and P. yunnanensis in the anal- yses of the multilocus datasets, respectively (Fig. 1). However, P. abietina has a stipe that is olive-ochraceous to dull ocher upward, quickly turning deep blue- green when handled or confined; its branches are yellow-ocher to dull ocher when fresh or somewhat greenish-ocher (Gonzalez-Avila et al. 2020). Phaeo- clavulina carovinacea differs from P fulva in its white flesh, which turns reddish brown when exposed to air (Index Fungorum 2024). Phaeoclavulina nigricans has basidiomata whose surface gradually develops a burgundy coloration when MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 167 Xin Tong et al.: New species of Phaeoclavulina from China touched, and its light cream flesh turns black rapidly upon exposure to air (Index Fungorum 2024). Phaeoclavulina yunnanensis is characterized by its yellow ba- sidiomata, with the tips showing a slight greenish-grey tint, and relatively small basidia (20-40 x 3-5 um) (Zheng et al. 2024). The species also differs from the other new species described in this study, Phaeoclavulina aurea, in both ba- sidiomata color and basidiospore morphology. Phaeoclavulina aurea has pale yellow to golden yellow basidiomata and basidiospores with truncate spines. Discussion The ornamentation of spores holds paramount significance in differentiating species within Phaeoclavulina. Previous studies have classified spore ornamen- tation in R. subg. Echinoramaria into five distinct types (Petersen 1981; Villegas et al. 2005). Volcanic spines, a distinctive form of spore ornamentation, are also evident in distinguishing Phaeoclavulina species, such as P. pancaribbea (R.H. Petersen) Giachini (2011: 194) (Petersen 1981; Giachini and Castellano 2011) and P. zealandica (R.H. Petersen) Giachini (2011: 195) (Petersen 1988; Giachini and Castellano 2011). The new species P aurea and P fulva described in this study also possess truncate warts or spines. However, it is noteworthy that both P pancaribbea and P zealandica have larger spores than the newly discovered P aurea and P fulva. Some species within Phaeoclavulina exhibit host specificity. For example, P. yunnanensis has only been found under Quercus sp. (Zheng et al. 2024), P. cyanocephala is exclusively associated with Abies sp. (Estrada-Torres 2007), and the new species P aurea discovered in this study has only been found un- der Pinus sp. This host specificity may serve as one of the diagnostic character- istics for species identification within Phaeoclavulina. Additionally, numerous species remain undiscovered globally, particularly in understudied forest eco- systems with favorable environmental conditions (Exeter et al. 2006), and many endemic taxa are still awaiting documentation. Therefore, further exploration of these habitats, combined with molecular and ecological studies, is essential to uncover the full diversity and host associations of Phaeoclavulina species. Previous studies have indicated that clavarioid fungi, including Phaeoclavu- lina, are highly diverse and widely distributed across the globe (Corner 1950). However, to date, only 22 species of Phaeoclavulina have been described from China, despite the country’s vast territory, complex climate, diverse habitats, and rich species resources, all of which contribute to its exceptionally high fun- gal diversity (Miller et al. 1991). This gap may be attributed to the limited num- ber of collections. In this study, two new species of Phaeoclavulina from North China were described using nrlTS-nrLSU phylogenetic analyses (Fig. 1) and macrofungal morphological examination, thereby contributing to the current understanding of the species diversity within this genus. Moreover, specimen collection and field investigation were conducted only in August, suggesting that additional Phaeoclavulina species may yet be discovered in this study area. Acknowledgments Special thanks are due to Dr. Li Fan and Dr. Qian Chen for their suggestions on the manuscript. MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 168 Xin Tong et al.: New species of Phaeoclavulina from China Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of Al No use of Al was reported. Funding This study was financed by the BUAST Budding Talent Program (23CE-BGS-06), the National Natural Science Foundation of China (No. 32270012), and the Biodiversi- ty Survey and Assessment Project of the Ministry of Ecology and Environment, China (2019HJ2096001006). Author contributions All authors have contributed equally. Author ORCIDs Xin Tong ® https://orcid.org/0000-0002-2091-6636 Yue Gao © https://orcid.org/0009-0001-0831-3656 Hai-Qi Wang ® https://orcid.org/0009-0004-3388-6263 Hao Zhou © https://orcid.org/0000-0002-4869-2187 Cheng-Lin Hou © https://orcid.org/0000-0001-81 62-5560 Data availability All of the data that support the findings of this study are available in the main text. References Brinkmann W (1897) Vorarbeiten zu einer Pilzflora Westfalen. Jahresbericht des West- falischen Provinzial-Vereins fiir Wissenschaft und Kunst 25: 195-207. Corner EJH (1950) A Monograph of Clavaria and allied Genera. Oxford University Press, London, 740 pp. Corner EJH (1970) Supplement to “A monograph of Clavaria and allied genera”. Beihefte zur Nova Hedwigia 33: 1-299. Deng PT, Liu WH, Ge ZW, Zhang P (2024) Three new ramarioid species of Phaeoclavu- lina (Gomphaceae, Gomphales) from China. MycoKeys 108: 1-14. https://doi. org/10.3897/mycokeys.108.128716 Estrada-Torres A (1994) La Familia Gomphaceae (Aphyllophorales: Fungi) en el Estado de Tlaxcala. Tesis doctorado (Doctorado en Ciencias (Biologia) ENCB, IPN. México D. F., 186 pp. Estrada-Torres A (2007) Hongos ectomicorrizdgenos y myxomycetes del Parque Nacional Lagunas de Montebello, Chiapas. Universidad Autonoma de Tlaxcala, Centro de Investi- gacion en Ciencias Bioldgicas. Final report SNIB-CONABIO project BKO043. Mexico, D. F. Exeter RL, Norvell L, Cazares E (2006) Ramaria of the Pacific Northwest. United States, Salem, Oregon, 157 pp. MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 169 Xin Tong et al.: New species of Phaeoclavulina from China Franchi P Marchetti M (2022) | Funghi Clavarioidi in Italia (Clavarioid Fungi in Italy). Oviedo, Spain. Gao Y, Tong X, Zhou H, Wang HQ, Li C, Hou CL (2024) Three new species of the genus Clavu- lina (Hydnaceae, Cantharellales) from North China based on morphological and phyloge- netic analysis. MycoKeys 108: 75-94. https://doi.org/10.3897/mycokeys.108.124004 Giachini AJ (2004) Systematics, Phylogeny, and Ecology of Gomphus Sensu Lato. Doc- toral Dissertation. Oregon State University, Corvallis. Giachini AJ, Castellano MA (2011) A new taxonomic classification for species in Gom- phus sensu lato. Mycotaxon 115: 183-201. https://doi.org/10.5248/115.183 Giachini AJ, Hosaka K, Nouhra E, Spatafora J, Trappe JM (2010) Phylogenetic relationships of the Gomphales based on nuc-25S-rDNA, mit-12S-rDNA, and mit-atp6-DNA combined sequences. Fungal Biology 114: 224-234. https://doi.org/10.101 6/j.funbio.2010.01.002 Gonzalez-Avila A, Martinez-Gonzalez R, Espinosa D, Estrada-Torres A (2020) Phaeoclavu- lina liliputiana sp. nov. (Gomphaceae, Gomphales) a new endemic species from Tlax- cala, Mexico. Phytotaxa 470: 155-164. https://doi.org/10.11646/phytotaxa.470.2.4 Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New al- gorithms and methods to estimate maximum likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59(3): 307-321. https://doi. org/10.1093/sysbio/syq010 Herrera MJ, Guzman L, Rodriguez O (2002) Contribucién al conocimiento de la micobio- ta de la region de San Sebastian del oeste, Jalisco, México. Acta Botanica Mexicana 589s 50: Hillis DM, Bull JJ (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology 42: 182-192. https://doi. org/10.1093/sysbio/42.2.182 Index Fungorum (2020) Index Fungorum. https://www.indexfungorum.org/Publica- tions/Index%20Fungorum%20n0.457.pdf [accessed Novermber, 2020] Index Fungorum (2024) Index Fungorum. http://www.indexfungorum.org/names/ names.asp [accessed September, 2024] Kang J, Kang Y, Ji X, Guo Q, Jacques G, Pietras M, Luczaj N, Li D, Ltuczaj t (2016) Wild food plants and fungi used in the mycophilous Tibetan community of Zhagana (Tewo County, Gansu, China). J Ethnobiol Ethhomed 12(1): 1-21. https://doi.org/10.1186/ $13002-016-0094-y Katoh K, Toh H (2010) Parallelization of the MAFFT multiple sequence alignment program. Bioinformatics 26(15): 1899-1900. https://doi.org/10.1093/bioinformatics/btq224 Kim DH, Chung HC, Ohga S, Lee SS (2003) ITS primers with enhanced specificity to de- tect the ectomycorrhizal fungi in the roots of wood plants. Mycobiology 31: 23-31. https://doi.org/10.4489/MYCO.2003.31.1.023 Liu WH, Yan J, Deng PT, Qin WQ, Zhang P (2022) Two new species of Phaeoclavulina (Gomphaceae, Gomphales) from Hunan Province, China. Phytotaxa 561(1): 27-40. https://doi.org/10.11646/phytotaxa.561.1.3 Larsson KH (2007) Rethinking the classification of corticioid fungi. Mycological Re- search 111(9): 1040-1063. https://doi.org/10.1016/j.mycres.2007.08.001 Marr CD, Stuntz DE (1973) Ramaria in western Washington. Bibliotheca Mycologica 38: 1-232: Martin MP, Danils PP, Erickson D, Spouge JL (2020) Figures of merit and statistics for detecting faulty species identification with DNA barcodes: A case study in Ramaria and related fungal genera. PLoS ONE 15(8): e€0237507. https://doi.org/10.1371/jour- nal.pone.0237507 MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 170 Xin Tong et al.: New species of Phaeoclavulina from China Miller KR, Mittermeier RA, Werner TB, Reid WV, Mcneely JA (1991) Conserving the world’s biological diversity. Gland, Switzerland, International Union for Conservatio of Nature and Natural Resources IUCN, 1990, 3(2): 131-133. Norvell L, Exeter R (2004) Ectomycorrhizal Epigeous Basidiomycete Diversity in Oregon Coast Range Pseudotsuga menziesii forests preliminary Observations. Fungi in For- est Ecosystems: Systematics, Diversity, and Ecology. The New York Botanical Gar- den, New York, 159-189. Petersen RH (1981) Ramaria subgenus Echinoramaria. Bibliotheca Mycologica 79: 1-261. Petersen RH (1988) The clavarioid fungi of New Zealand. DSIR Science Information Pub- lishing, Wellington, New Zealand, 170 pp. Petersen RH, Hughes KW, Justice J (2014) Two new species of Ramaria from Arkansas. Mycokeys 8: 17-29. https://doi.org/10.3897/mycokeys.8.7356 Posada D, Crandall KA (1998) Modeltest: Testing the model of DNA substitution. Bioin- formatics 14: 817-818. https://doi.org/10.1093/bioinformatics/14.9.817 Rannala B, Yang Z (1996) Probability distribution of molecular evolutionary trees: A new method of phylogenetic inference. Journal of Molecular Evolution 43: 304-311. https://doi.org/10.1007/BF02338839 Rathnayaka AR, Tennakoon DS, Jones GE, Wanasinghe DN, Bhat DJ, Priyashantha AH, Stephenson SL, Tibpromma S, Karunarathna SC (2024) Significance of precise doc- umentation of hosts and geospatial data of fungal collections, with an emphasis on plant-associated fungi. New Zealand Journal of Botany: 1-28. https://doi.org/10.10 80/0028825X.2024.2381734 Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572-1574. https://doi.org/10.1093/bioinformat- ics/btg180 Schild E (1990) Ramaria-Studien. Zeitschrift fiir Mykologie 56(1): 131-150. Sims K, Watling R, De la Cruz R, Jeffries P (1997) Ectomycorrhizal fungi of the Philippines: Apreliminary survey and notes on the geographic biodiversity of the Sclerodermatales. Biodiversity and Conservation 6: 45-58. https://doi.org/10.1023/A:1018371515051 Stamatakis A (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analy- ses with thousands of taxa and mixed models. Bioinformatics 22(21): 2688-2690. https://doi.org/10.1093/bioinformatics/btl446 Stamatakis A, Hoover P Rougemont J (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57(5): 758-771. https://doi. org/10.1080/10635150802429642 Sui XN, Guo MJ, Zhou H, Hou CL (2023) Four new species of Phyllosticta from China based on morphological and phylogenetic characterization. Mycology 14(3): 190- 203. https://doi.org/10.1080/21501203.2023.2225552 Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: Molecular evolu- tionary genetics analysis version 6.0. Molecular Biology and Evolution 30(12): 2725- 2729. https://doi.org/10.1093/molbev/mst197 van Overeem C (1923) Beitrage zur Pilzflora von Niederlandisch Indien. Bulletin du Jar- din Botanique de Buitenzorg Série Ill 5: 247-293. Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238-4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990 Villegas M, Cifuentes J, Estrada-Torres AE (2005) Sporal characters in Gomphales and their significance for phylogenetics. Fungal Diversity 18: 157-175. https://doi. org/10.1016/j.femsyr.2005.01.001 MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 171 Xin Tong et al.: New species of Phaeoclavulina from China Wang YT, Huang ZH, Wang J, Tong Z, Cui GF (2021) The population structure and dy- namic characteristics of Phellodendron amurense in Yanshan Mountains. Acta Eco- logica Sinica 47(7): 2826-2834. https://doi.org/10.5846/stxb202003300743 Wannathes N, Kaewketsri R, Suwannarach N, Kumla J, Lumyong S (2018) Phaeoclavuli- na pseudozippelii sp. nov. (Gomphales, Basidiomycota) from Northern Thailand. Phy- totaxa 362: 211-219. https://doi.org/10.11646/phytotaxa.362.2.7 White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR Protocols: A Guide to Methods and Applications. Academic Press, San Diego, 315-322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1 Yan J, Wang XY, Wang XH, Chen ZH, Zhang P (2020) Two new species of Clavaria (Agaricales, Basidiomycota) from Central China. Phytotaxa 477: 71-80. https://doi. org/10.11646/phytotaxa.477.1.5 Zheng DG, Lu WH, Han MY, Elgorban AM, Yang JY, Zhou YQ, Suwannarach N, Tibpromma S, Karunarathna SC (2024) Morphology and phylogeny reveal a new ramarioid spe- cies of Phaeoclavulina (Gomphaceae, Basidiomycota) from Yunnan Province, China. Phytotaxa 670: 109-118. https://doi.org/10.11646/phytotaxa.670.2.3 Zhou H, Hou CL (2019) Three new species of Diaporthe from China based on morpho- logical characters and DNA sequence data analyses. Phytotaxa 422(2): 157-174. https://doi.org/10.11646/phytotaxa.422.2.3 Zhou H, Wang QT, Tong X, Hou CL (2021) Phylogenetic analysis of Engleromyces sinen- sis and identification of cytochalasin D from culture. Mycological Progress 20(10): 1343-1352. https://doi.org/10.1007/s11557-021-01739-z Zhou H, Cheng GQ, Sun XM, Cheng RY, Zhang HL, Dong YM, Hou CL (2022) Three new species of Candolleomyces (Agaricomycetes, Agaricales, Psathyrellaceae) from the Yanshan Mountains in China. MycoKeys 88: 109-121. https://doi.org/10.3897/my- cokeys.88.81437 MycoKeys 120: 157-172 (2025), DOI: 10.3897/mycokeys.120.138950 172