683 MycoKeys MycoKeys 121: 357-374 (2025) DOI: 10.3897/mycokeys.121.158142 Research Article Three new species of Clavariadelphus (Clavariadelphaceae, Gomphales) from the Hengduan Mountains region, China Mei-jia Li'©, Peng-tao Deng'™®, Zuo-hong Chen", Ping Zhang'® 1 College of Life Science, Hunan Normal University, Changsha 410081, China Corresponding author: Ping Zhang (zhangping0000@163.net) OPEN Qaceess Academic editor: Bao-Kai Cui Received: 7 May 2025 Accepted: 5 July 2025 Published: 2 September 2025 Citation: Li M-jia, Deng P-tao, Chen Z-hong, Zhang P (2025) Three new species of Clavariadelphus (Clavariadelphaceae, Gomphales) from the Hengduan Mountains region, China. MycoKeys 121: 357-374. https://doi.org/10.3897/ mycokeys.121.158142 Copyright: © Mei-jia Li 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 Specimens of three unnamed club-shaped fungi, collected from the Hengduan Mountains region in China, are formally described as Clavariadelphus acuminatus, C. miniatus, and C. pseudoelongatus. Based on ITS, nrLSU, ATP6, and TEF1 sequence data, a multigene phylogenetic analysis indicated that the three new species each formed a distinct lineage within the Clavariadelphus clade with strong support. Descriptions, illustrations, and the results of molecular phylogenetic analyses for the three newly discovered species are presented. A taxonomic key for the known Clavariadelphus species in China is provided. Key words: Club-shaped fungi, morphological characteristics, new taxa, phylogenetic analysis, taxonomy Introduction Clavariadelphus Donk (Clavariadelphaceae, Gomphales, Basidiomycota) is a genus of symbiotic or saprophytic clavarioid fungi (Deng et al. 2020). Some members of the genus are reported to be mycorrhizal with diverse hosts, such as oak, pine, and spruce species (Hanif et al. 2022). The genus was initially proposed in 1933 to accommodate three species, comprising C. pistillaris (L.) Donk, C. ligula (Schaeff.) Donk, and C. truncatus (Quél.) Donk, with C. pistillaris designated as the type species (Corner 1950). The genus includes unbranched clavarioid fungi with simple, erect, club-shaped basidiomata; a thick hyme- nium composed of (2-)4-spored basidia and clavate leptocystidia; ellipsoid to amygdaliform, smooth, thin-walled, inamyloid basidiospores; and clamped tramal hyphae (Corner 1950, 1970; Methven 1990). The spore length-to-width ratio is an important distinguishing character for Clavariadelphus. The genus was divided into sect. Clavariadelphus and sect. Ligula based on the spore length-to-width ratio (Wells and Kempton 1968). The spores of some Clavaria- delphus species are ellipsoid to broadly ellipsoid (Q_ < 2.5), as in sect. Clavari- adelphus, whereas in other species the spores are narrowly ellipsoid (Q_, 2 3), as in sect. Ligula (Wells and Kempton 1968). Clavariadelphus has a widespread distribution in temperate forests of the Northern Hemisphere. The majority of early-described Clavariadelphus species were reported from Europe and North America. In Index Fungorum (Partnership 357 Mei-jia Li et al.: Three new species of Clavariadelphus from China 2010), 47 fungal records have been assigned to Clavariadelphus, of which 33 species are formally accepted (Deng et al. 2020; Huang et al. 2020; Xia and Fan 2020), and the remaining records are classified in other fungal groups. In 2020, seven Clavariadelphus species had been recorded exclusively from China, namely, C. alpinus J. Zhao & L.P. Tang, C. amplus J. Zhao, L.P. Tang & Z.W. Ge, C. aurantiacus P. Zhang, C. gansuensis J. Zhao & L.P. Tang, C. griseoclavus L. Fan & L. Xia, C. khinganensis J. Zhao, L.P. Tang & P. Zhang, and C. tenuis P. Zhang. The Hengduan Mountains region in China is one of 34 global biodiversity hotspots (Myers et al. 2000; Mittermeier et al. 2005). Previously, the species diversity of macrofungi from the Hengduan Mountains region has been investi- gated, and many new species have been discovered from the region (Cui et al. 2018; Dai et al. 2021; Ji et al. 2022; Liu et al. 2023; Song et al. 2025). In the pres- ent study, using our collections of Clavariadelphus specimens, three new Cla- variadelphus species from the Hengduan Mountains region are identified, which are described herein as C. acuminatus, C. miniatus, and C. pseudoelongatus. Materials and methods Fungal material Three new species of Clavariadelphus were collected from the Hengduan Moun- tains region during 2018-2024. The fresh basidiomata were photographed and recorded relevant characteristics in the field. All collected specimens were dried at 50 °C-60 °C in a drying oven or desiccated in silica gel. The dried spec- imens are housed in the Mycological Herbarium of Hunan Normal University (MHHNU), Changsha, China. Morphological descriptions Macromorphological characters were described from detailed field notes and habitat photographs. A section of dried hymenial tissue was taken from the materials and placed in 5% KOH solution containing 1% Congo Red or Melzer’s reagent. The microscopic structure of each species was measured as at least 25 sets of parallel data, including the basidia, basidiospores and hyphae. Color descriptions and codes follow Kornerup and Wanscher (1978) and Ridgway (1912). In the basidiospore description, the size of basidiospores is expressed in the form (a—) b-c (-d), where the range b-c is the majority of measured values, and a and d are extreme values of the spore dimension. The notation [n/m/p] refers to the number of measurements recorded for n basidiospores from m basidiomata of p specimens. The Q value represents the length-to-width ratio of basidiospores, and the Q_ value is the average Q + standard deviation. Scanning electron microscopy A small portion of tissue containing the hymenium was sampled from the dried specimens, mounted onto aluminum stubs, and coated with gold palladium. Scanning electron microscopy was conducted with a TESCAN CLARA Xplore 30 operating at 20 keV. The scanning electron micrographs of basidiospores of three new Clavariadelphus species were shown in Fig. 2. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 358 Mei-jia Li et al.: Three new species of Clavariadelphus from China DNA extraction, PCR amplification, and sequencing Total genomic DNA was extracted from the collected specimens using the Ezup Column Fungi Genomic DNA Purification Kit (Sangon Biotech, Shanghai, China). The universal primers ITS4/ITS5 and LR5/LROR were used to amplify the internal transcribed spacer (ITS) region of ribosomal DNA and the nuclear large subunit (nrLSU) region in the 28S ribosomal RNA gene (Vilgalys and Hester 1990; White et al. 1990; Gardes and Bruns 1993). The ATP6 and TEF71 genes were respectively amplified using the primer pairs ATP6-1R/ATP6-1F and EF-CR/EF-CF or EF-d2/ EF-d1 (Deng 2021). Each PCR amplification reaction was performed on an Ep- pendorf Mastercycler thermal cycler (Eppendorf, Inc., Hamburg, Germany). The 25 pL reaction mixture contained 1 x PCR buffer, 1.5 mM MgCl, 0.2 mM dNTP,, 0.4 uM of each primer, 1.25 U Taq polymerase (Sangon Biotech), and 1 uL DNA template. The PCR protocol for each amplification was as follows: pre-denatur- ation at 94 °C for 5 min, then 34 cycles of denaturation at 94 °C for 30 s, annealing at an appropriate temperature (55 °C for 30s for ITS and nrLSU, 52-54 °C for 40 s for TEF7 and ATP6; He et al. 2023), and extension at 72 °C for 1 min, followed by a final extension at 72 °C for 7 min, and then held at 8 °C. The amplified PCR prod- ucts were separated by 1% agarose gel electrophoresis and the same primers were used for sequencing by Sangon Biotech. All sequences newly generated in this study were deposited in GenBank (accession numbers are listed in Table 1). Alignment and phylogenetic analysis In addition to the newly generated sequences from 25 specimens, the remain- der of the sequences used in this study were publicly available from GenBank. A multiple sequence alignment for each of the four gene fragments, namely, ITS, nrLSU, TEF1, and ATP6, was constructed with MAFFT v7.149 (Katoh and Standley 2016). The aligned sequences were manually adjusted and trimmed as necessary with BioEdit v7.2.5 (Hall 1999). A combined matrix, comprising 146 sequences, was generated with Sequence Matrix v7.2.5 (Hall 1999). Two species of Gomphus (Pers.) Gray, G. clavatus (Pers.) Gray and G. /udovicianus R.H. Petersen, Justice & D.P. Lewis, were selected as the outgroup. Maximum likelihood analysis was conducted with RAxML v7.2.6 (Stamatakis et al. 2005; Stamatakis 2006), using a GTR+Gamma evolutionary model (Stamatakis et al. 2008) with 1000 bootstrap replicates. Bayesian inference (BI) was performed using MrBayes v3.2.7 (Ronquist et al. 2012) with the GTR+I+G optimal evolutionary model, and analyses were run for 2,000,000 generations using four Metropolis-coupled Monte Carlo Markov chains to calculate posterior probabilities (PP). The tree files were visualized with FigTree v1.4.2 (Rambaut 2012) and slightly adjusted for presentation purposes with Photoshop CS6 and Illustrator CS6 (Adobe Systems, Inc., San Jose, CA, USA). Results Phylogenetic analyses The final multigene dataset, which constituted 146 sequences (66 ITS, 22 ATP6, 21 TEF1, and 37 nrLSU), was used for the ML and BI analyses. The matrix of con- catenated sequences comprised 2776 bp. The phylogenetic tree included 22 MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 359 Mei-jia Li et al.: Three new species of Clavariadelphus from China Table 1. Details of sequences used and produced in phylogenetic analysis. The sequences newly generated in this study are highlighted in bold. The type specimens and type species in Clavariadelphus are indicated with asterisks (*); — represents data unavailability. Taxa voucher GenBank No. | GenBank No. | GenBank No. | GenBank No. Leestion Rataraneas ITS nrLSU TEF1 ATP6 Clavariadelphus acuminatus MHHNU 12163* PV463549 PV490621 - PV523761 China This study C. acuminatus MHHNU 12164 PV463550 PV490622 *. PV523762 China This study C.americanus MycoMap # 1288 MK575228 a a = USA unpublished C. amplus HKAS 76577 MK705851 MK704443 MK736675 = China Huang et al. (2020) C. amplus HKAS 54876* MK705857 MK704444 MK736676 = China Huang et al. (2020) C. amplus HKAS 49229 MK705854 MK704447 MK736677 = China Huang et al. (2020) C. amplus HMAS 250466 MK705858 MK704448 = = China Huang et al. (2020) C. aurantiacus MHHNU 9256* MT580787 PV490611 PV523772 PV523753 China Deng et al. (2020), this study C. aurantiacus MHHNU 10085 MT580792 PV490612 PV523773 PV523754 China Deng et al. (2020), this study C. aurantiacus HKAS 53889 MT580788 = = = China Deng et al. (2020) C. elongates SWATO000559 MG768848 = a = Pakistan | Sher et al. (2018) C. elongates LAH31397 MG768847 7 = = Pakistan Sher et al. (2018) C. elongates HMAS 260746 MK705845 MK704441 MK736672 = China Huang et al. (2020) C. elongates HKAS 50742 MK705843 MK704440 = = China Huang et al. (2020) C. elongates MHHNU 9250 PV463551 PV490604 PV523767 PV523746 China This study C. elongates MHHNU 9933 PV463552 PV490605 PV523768 PV523747 China This study C. griseoclavus BJTC FM964 MT302370 7 = = China Xia and Fan (2020) C. griseoclavus BJTC FM965 MT302371 < “ = China Xia and Fan (2020) C. gansuensis HKAS 76487 MK705847 MK704442 MK736673 = China Huang et al. (2020) C. himalayensis HKAS 50684 MK705863 = = = China Huang et al. (2020) C. himalayensis HKAS 58811 MK705864 = 2 = China Huang et al. (2020) C. himalayensis MHHNU 9167 PV463553 PV490610 PV523771 PV523750 China This study C. khinganensis MHHNU 7789* MK705865 MK704451 MK736680 PV523758 China Huang et al. (2020), this study C. khinganensis MHKMU H.Y. Huang 368 | M1T447468 _ 5 2 China Huang et al. (2020) C. khinganensis MHHNU 7822 PV463554 PV490616 PV523776 PV523759 China This study C. ligula OMDL iNat # 180557549 PP959633 = = > USA Unpublished C. ligula NAMPA2215-22 OP225575 = = = USA Unpublished C. ligula AMB 18570 MT055950 = = = Italy Unpublished C. ligula 3833 KM248918 - = Canada Unpublished C. ligula MHHNU 10452 PV463555 PV490608 a PV523751 China This study C. ligula MHHNU 10453 PV463556 PV490609 = PV523752 China This study C. mucronatus OSC 1064138 EU526000 = 4 - USA Smith et al. (2002) C. miniatus MHHNU 9952* PV463557 PV490619 PV523777 PV523760 China This study C. miniatus MHHNU 12089 PV463558 PV490620 = = China This study C. occidentalis OSC 104664 EU669308 - a - USA Unpublished C. occidentalis OSC 114281 EU846242 - = - USA Unpublished C. pakistanicus mh129901* HQ379937 = = = Pakistan Hanif et al. (2014) C. pakistanicus MHHNU 9282 MT580789 PV490603 PV523766 PV523745 China Deng et al. (2020), this study C. pistillaris* NAMA 2017-123 MH979250 - - = USA Unpublished C. pistillaris* FLAS-F-60521 MH281842 - = - USA Unpublished C. pseudoelongatus MHHNU 32323* PV463559 PV490617 4 - China This study C. pseudoelongatus MHHNU 12123 PV463560 PV490618 = - China This study C. sachalinensis OSC 96213 EU834196 F 7 = USA Unpublished C. sachalinensis p059i EU624410 " = 7 USA Unpublished C. sachalinensis MHHNU 7816 MT580791 MK704452 = PV523744 China | Huang etal. (2020), this study C. sachalinensis p058i EU624411 = = = USA Unpublished C. subfastigiatus OSC 119587 EU669207 EU669259 = ~ USA Unpublished MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 360 Mei-jia Li et al.: Three new species of Clavariadelphus from China Taxon C. subfastigiatus C. truncates C. truncates C. truncates C.tenuis C.tenuis C.tenuis C. unicolor C. yunnanensis C. yunnanensis C. yunnanensis C. yunnanensis C. yunnanensis L. byssiseda L. byssiseda L. patouillardii L. patouillardii Gomphus clavatus* G. ludovicianus GenBank No. GenBank No. GenBank No. GenBank No. Voucher ITS nrL SU TEF1 ATP6 Location References MICH 73554 JX275756 = — = USA Unpublished SIM278 HQ650728 = = = Canada Kranabetter et al. (2009) QHU20413 OM974113 OM942741 = = China Unpublished UBC: F30993 MZ836052 - = = Canada Unpublished MHHNU 9897 MT580786 PV490613 PV523774 PV523755 China Deng et al. (2020), this study MHHNU 9900 MT580793 PV490614 — PV523756 China Deng et al. (2020), this study MHHNU 9934* MT580790 PV490615 PV523775 PV523757 China Deng et al. (2020), this study Mushroom Observer # MN906166 - = = USA Unpublished 112193 HMAS 250510 MK705874 MK704458 MK736685 = China Huang et al. (2020) HKAS 54849 MK705869 MK704453 MK736681 = China Huang et al. (2020) HKAS 63558 MK705870 MK704454 MK736682 = China Huang et al. (2020) MHHNU 9244 PV463561 PV490606 PV523769 PV523748 China This study MHHNU 9977 PV463562 PV490607 PV523770 PV523749 China This study MHHNU 10765 PV463563 PV490623 = PV523763 China This study MHHNU 9087 PQ248145 PQ242649 = PV523764 China Li et al. (2025), this study MHHNU 10443 PQ248147 PQ242651 = PV523765 China Li et al. (2025), this study HMJAU 26892 KU870449 = = = China Liu et al. (2017) EL 64/03 EU118628 - = = Sweden Larsson (2007) TFB14476 clone c8* KJ655570 7 = = USA Petersen et al. (2014) Clavariadelphus taxa, 2 Lentaria Corner taxa, and two Gomphus species (G. clava- tus and G. ludovicianus) as the outgroup. The ML tree (Fig. 1) was highly similar in topology to the BI tree (not shown). The bootstrap support (BS) values and posterior probabilities (PP) of relevant nodes, which were greater than or equal to 50% and 0.95, respectively, are shown in Fig. 1. In this study, six samples of three newly gathered Clavariadelphus species were placed in different Clavariadelphus subclades with strong support (all PP 1, BS 100%) and each formed a distinct lin- eage. Clavariadelphus acuminatus and C. pseudoelongatus were sister taxa within a well-supported subclade (PP 1, BS 100%). The novel saprophytic species C. min- iatus formed a distinct lineage sister to another saprophytic species, C. sachlin- ensis (S. Imai) Corner, with strong statistical support (PP 1, BS 100%). The results of the molecular phylogenetic analyses supported the distinctness of the three putative new Clavariadelphus species, which are formally described herein. Taxonomy Clavariadelphus acuminatus X.L. Gao & P. Zhang, sp. nov. MycoBank No: 858803 Figs 3,4 Diagnosis. Differs from other Clavariadelphus species in that the apex of the basidiomata is acuminate and pale to brownish yellow or golden-brown in color. Type. CHINA * Yunnan Province, Lufeng County, 25°24'02"N, 101°76'10"E, 2300 m asl., 4 September 2023, leg. X.L. Gao (holotype MHHNU 12163). Etymology. Acuminatus (Latin) refers to the acuminate apex of the basidiomata. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 361 Mei-jia Li et al.: Three new species of Clavariadelphus from China 1/100 0.05 Figure 1. Maximum likelihood (ML) phylogenetic tree of Clavariadelphus species and Lentaria species inferred from the concatenated multigene sequence dataset (ITS, nrLSU, ATP6, and TEF7). The Bayesian inference posterior probabilities > 0.95 and ML bootstrap values = 50% are shown at the nodes; “— olds. Accessions of the three new Clavariadelphus species are in bold. Clavariadelphus khinganensis MHHNU 7822 Clavariadelphus khinganensis MHHNU 7789 1/100! Clavariadelphus khinganensis MHIKMU H.Y.Huang 368 Clavariadelphus occidentalis OSC 114281 W100LClavariadelphus occidentalis OSC104664 {rascanadelel us pseudoelongatus MHHNU32323 /fiavariadelphus pseudoelongatus MHHNU12123 Clavariadelphus acuminatus MHHNU 12163 1 ea acuminatus MHHNU 12164 Clavariadelphus americanus MycoMap # 1288 Clavariadelphus himalayensis HKAS 50684 1/100! Clavariadelphus himalayensis HKAS 58811 iG laverratiel airs himalayensis MHHNU 9167 "Clavariadelphus pistillaris FLAS-F-60521 1 giavariadelphus pistillaris NAMA 2017-123 Ka lavar iadelphus subfastigiatus OSC 119587 Clavariadelphus subfastigiatus MICH 73554 Clavariadelphus mucronatus OSC 1064138 Clavariadelphus unicolor $.D, Russell Mushroom Observer # 112193 Clavariadelphus tenuis MHHNU 9934 1/100! Clavariadelphus tenuis MHHNU 9897 Clavariadelphus tenuis MHHNU 9900 Clavariadelphus elongatus HMAS 260746 Clavariadelphus elongatus LAH31397 Clavariadelphus elongatus MHHNU 9933 eavariogelalin elongatus MHHNU 9250 Clavariadelphus elongatus SWAT000559 Clavariadelphus elongatus HKAS 50742 Clavariadelphus yunnanensis MHHNU 9244 Clavariadelphus yunnanensis MHHNU 9977 Clavariadelphus yunnanensis HKAS 54849 Clavariadelphus yunnanensis HKAS 63558 Clavariadelphus yunnanensis HMAS 250510 Nal eae a gansuensis HKAS 76487 vioo) Clavariadelphus truncatus QHU20413 Clavariadelphus truncatus SIM278 1/99 Glavertade Talia truncatus UBC F30993 Clavariadelphus aurantiacus HKAS 33889 Clavariadelphus aurantiacus MHHNU 10085 Clavariadelphus aurantiacus MHHNU 9256 Clavariadelphus pakistanicus mh129901 Clavariadelphus pakistanicus MHHNU 9282 9.98/98! Clavariadelphus amplus HKAS 76577 Clavariadelphus amplus HKAS 54876 Clavariadelphus amplus HKAS 49229 Clavariadelphus amplus HMAS 250466 W109 Clavariadelphus griseoclavus BJTC FM964 Clavariadelphus griseoclavus BITC FM965 0.96/57 0.99/67 1/98 1/100 1/100 -/56 Clavariadelphus sachalinensis OSC 96213 Clavariadelphus sachalinensis p058i 98} Clavariadelphus sachalinensis p059i 1/100] Clavariadelphus sachalinensis MHHNU7816 Claviriageiphus miniatus MHHNU 12089 5/1 Clavariadelphus miniatus MHHNU 9952 Lentaria patouillardii MHHNU 12089 ie Lentaria patouillardii HMJAU 26892 wio9|Lentaria byssiseda MHHNU 10765 Lentaria byssiseda MHHNU 9087 Clavariadelphus ligula 3833 -/86| Clavariadelphus ligula NAMPA22 15-22 Clavariadelphus ligula OMDL iNat # 180557549 Clavariadelphus ligula MHHNU 10452 Clavariadelphus ligula AMB 18570 Clavariadelphus ligula MHHNU 10453 Gomphus ludovicianus TFB14476 clone c8 Gomphus clavatus EL 64/03 (GB) MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 V 9pelD dq 9Pe[D D 9petD ” indicates that the values were less than these thresh- Mei-jia Li et al.: Three new species of Clavariadelphus from China Figure 2. Scanning electron micrographs of basidiospores of the three new Clavariadelphus species. A, B. C. acuminatus (MHHNU 12163, holotype); C, D. C. miniatus (MHHNU 9952, holotype); E, F. C. pseudoelongatus (MHHNU 32323, holo- type). Scale bars: 1 um. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 Mei-jia Li et al.: Three new species of Clavariadelphus from China oe Bik? vit _ — Se —_ Figure 3. Basidiomata of Clavariadelphus acuminatus. A. MHHNU 12163; B. MHHNU 12164. Scale bars: 1 cm. Description. Basidiomata up to 15 cm high, 0.4-0.8 cm in diameter at the base, 0.8-1.2 cm in diameter in the middle, slightly tapering at both ends, simple, initially narrowly cylindrical, narrowly fusiform after maturity, mostly flexuous; hymenium initially smooth, then longitudinally rugose with age, pale (2A2) to pale orange (5A3) to apricot (5B6); apex fertile, obtuse to acute, smooth to rugose, darker than the hymenium with age, pale to brownish yellow (5C8) or golden brown (5D7); base terete, almost smooth, white to cream (4A3); context solid when young, gradually becoming soft and spongy with age. Odor and taste not recorded. Hymenium extending over the apex of the basidioma, composed of basidia and leptocystidia. Basidia 75-99 x 8.5-13 um, narrowly clavate, pale yellow in KOH, smooth, thin-walled, clamped, 4 sterigmata (rarely 2), 6-12 um tall, with numerous granular contents and guttules. Basidiospores [60/3/2] 8—9.6(- 10.0) x 6-7.8(-8.0) um [Q = (1.12-)1.14-1.50(-1.60), Q. = 1.36 + 0.10], el- lipsoid to broadly ellipsoid, with a prominent apiculus, and several oleiferous guttules within the spores, pale yellow in KOH, inamyloid, thin-walled, smooth. Leptocystiada 45-69 x 2-4 um, mostly clavate, hyaline, smooth, clamped, with branches. Contextual hyphae 4-6 um in diameter, thin-walled, clamped. Ecological information and distribution. Solitary, scattered or gregarious on the ground in mixed coniferous and broadleaved forest at elevations of 2300- 2400 m. Southwestern China. Additional material examined. CHINA * Yunnan Province, Gucheng Dis- trict, Kainan Street, 26°77'41"N, 100°26'77"E, 2400 m asl., 9 September 2023 (MHHNU 12164). Comments. Clavariadelphus acuminatus is distinguishable from other Cla- variadelphus species by the distinctive acuminate apex of the basidiomata. Morphologically, the taxon is similar to C. khinganensis in the color of the ba- sidiomata, but the apex in C. khinganensis is obtuse or broadly rounded. In the phylogenetic analysis, C. acuminatus was the closest relative to C. pseudoe- longatus with strong support (PP 1, BS 100%). Although the two species oc- cupy a similar habitat, and have a subacute apex and flexuous basidiomata, in C. acuminatus the basidiomata have a brighter color and more strongly acute apex. Clavariadelphus acuminatus has pale to pale orange to apricot basidioma- ta, whereas C. pseudoelongatus has ecru-drab to light purple-drab basidioma- ta. In addition, microscopic features distinguish these species: C. acuminatus MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 364 Mei-jia Li et al.: Three new species of Clavariadelphus from China has smaller basidiospores (8-9.6 x 6—7.8 um vs. 9.8-11.0 x 6.4—9.5 um) and shorter basidia (75-99 x 8.5-13 um vs. 81-113 x 8-13 um). Thus, C. acumina- tus and C. pseudoelongatus can be distinguished by the color of the basidioma- ta, and dimensions of the basidiospores and basidia. QOOQ Or QOO0C Figure 4. Microscopic features of Clavariadelphus acuminatus (MHHNU 12163). A. Basidiospores; B. Basidia. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 365 Mei-jia Li et al.: Three new species of Clavariadelphus from China Clavariadelphus miniatus P. Zhang & M.J. Li, sp. nov. MycoBank No: 858804 Figs 5, 6 Diagnosis. Differs from other Clavariadelphus species in the small and pale yellow to salmon to purple-drab basidiomata with age. Type. CHINA * Yunnan Province, Deqin Prefecture, 27°24'15"N, 98°57'55'E, 3750 m asl., 17 August 2018, leg. P. Zhang (holotype MHHNU 9952). Etymology. Miniatus (Latin) refers to the small basidiomata of this species. Description. Basidiomata 4-10 cm tall, 0.2-0.5 cm in diameter, simple, nar- rowly cylindrical to clavate with age, slightly flexuous when young, occasionally branched; hymenium initially smooth, longitudinally rugose at maturity, pale yel- low (2A3) to salmon (6A4) to purple-drab (11E5); apex fertile, smooth to rugose, obtuse to subacute, occasionally truncated with age, monochromatic with hyme- nium; base terete, almost smooth, pale orange (5A3); context solid when young, gradually becoming soft and spongy with age. Odor and taste not recorded. Hymenium extending over the apex of basidiomata, composed of basidia and leptocystidia. Basidia 72-120 x 10-13 um, narrowly clavate, pale yellow in the KOH, smooth, thin-walled, clamped, 4 sterigmata (rarely 2), 6-12 um tall, with nu- merous granular contents and guttules. Basidiospores [66/3/2] (16.5—) 18.0-26.6 (—27.0) x 4-6 (-7.2) pm [Q = (2.75-) 3.00-5.67 (6.50), Q_ = 4.35 + 0.79], ellipsoid to oblong, with a prominent apiculus, hyaline in the KOH, thin-walled, smooth, inam- yloid. Leptocystida 30-82 x 3-4 um, narrowly clavate, hyaline, smooth, clamped, with branches. Contextual hyphae 3-7 um in diameter, thin-walled, clamped. Ecological information and distribution. Scattered or gregarious on the ground or on twigs in forest dominated by Picea in an elevational range of 3750-3800 m. Southwestern China. Additional material examined. CHINA * Yunnan Province, Degin Prefecture, Baima Snow Mountain, 28°15'28'"N, 99°15'20"E, 3800 m asl., 19 August 2024 (MHHNU 12089). Comments. Unlike the other two new species identified in this study, C. minia- tus is saprophytic and can be distinguished by the small size and pale yellow to salmon to purple-drab basidiomata. This taxon is confusable with C. sachalin- ensis. With regard to morphological characters and molecular sequence data, C. miniatus is similar to C. sachalinensis. Both species were resolved in a clade Figure 5. Basidiomata of Clavariadelphus miniatus. A. MHHNU 9952; B. MHHNU 12089. Scale bars: 1 cm. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 366 Mei-jia Li et al.: Three new species of Clavariadelphus from China 10um Figure 6. Microscopic features of Clavariadelphus miniatus (MHHNU 9952). A. Basidiospores; B. Basidia. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 367 Mei-jia Li et al.: Three new species of Clavariadelphus from China phylogenetically close to the two species of Lentaria, but their basidiospores are oblong, thin-walled, and hyaline in the KOH. Clavariadelphus sachalinensis has tawny or light walnut-brown to light brown basidiomata; thus, it is confusing to distinguish it from C. miniatus based onthe color of the basidiomata alone. How- ever, microscopic characteristics differ between C. miniatus and C. sachalinen- sis. Specifically, C. miniatus has longer basidiospores (18.0-26.6 x 4—6 um vs. 21-24 x 4-6 um) and basidia (72-120 x 10-13 um vs. 65-105 x 8-12.5 um). Clavariadelphus pseudoelongatus P. Zhang & Z. H. Chen, sp. nov. MycoBank No: 858805 Figs 7,8 Diagnosis. Differs from other Clavariadelphus species in having ecru-drab to light purple drab basidiomata with age, becoming cinnamon after cutting or bruising. Type. CHINA: Yunnan Province, Shangri-La Prefecture, 27°13'13"N, 100°02'20"E, 3828 m asl., 26 August 2020, leg. P. Zhang (holotype MHHNU 32323). Etymology. Pseudoelongatus (Latin) refers to having longer basidiomata than other Clavariadelphus species, but differing from C. elongatus. Description. Basidiomata 6-14 cm tall, 0.7-0.9 cm in diameter, simple, narrow- ly cylindrical to clavate in age, erect or slightly flexuous, occasionally branched; hymenium longitudinally apparent rugose, ecru-drab (5A2-4) to light purple drab (12D3-12D4), slowly stained cinnamon (6D6) after cutting or bruising; apex fertile, smooth to rugose, obtuse to subacute with age, monochromatic with hymenium, rarely dark violet (16F8); base terete, almost smooth, white; context solid when young, gradually becoming soft and spongy in age. Odor and taste not recorded. Hymenium extending over the apex of the basidiomata, composed of basidia and leptocystidia. Basidia 81-113 x 8-13 um, narrowly clavate, pale yellow in KOH, smooth, thin-walled, clamped, 4 sterigmata (rarely 2), 8-15 um tall, with nu- merous granular contents and guttules. Basidiospores [60/3/2] (9.2—) 9.8-11.0 (-13.0) x (6.0-) 6.4-9.5 (-11.0) um [Q = 1.16-1.55(-1.67), Q. = 1.31 + 0.10], ellipsoid to broadly ellipsoid, with a prominent apiculus, and several oleiferous guttules within the spores, pale yellow in KOH, thin-walled, smooth, inamyloid. Leptocystida 31-71 x 2—6 um, narrowly clavate, hyaline, smooth, clamped, with branches. Contextual hyphae 5-9 um in diameter, thin-walled, clamped. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 368 Mei-jia Li et al.: Three new species of Clavariadelphus from China WOOF OOOC Figure 8. Microscopic features of Clavariadelphus pseudoelongatus (MHHNU 32323). A. Basidiospores; B. basidia. : 10um ) y y MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 369 Mei-jia Li et al.: Three new species of Clavariadelphus from China Ecological information and distribution. Scattered or gregarious on the ground in the coniferous and broadleaved mixed forest in an elevational range of 3800-3828 m. Southwestern China. Additional material examined. CHINA « Yunnan Province, Shangri-La Prefec- ture, Qianhu mountain, 27°25'45"N, 99°40'55"E, 3800 m asl., 24 August 2024 (MHHNU 12123). Comments. Clavariadelphus pseudoelongatus can be distinguished by hav- ing longer and ecru-drab to light purple-drab basidiomata. It was initially mis- identified as C. elongatus J. Khan, Sher & Khalid owing to the similar color of the basidiomata, which are light purple at maturity. In the phylogenetic analy- sis, C. pseudoelongatus and C. elongatus were respectively placed in different subclades based on molecular evidence. In addition, C. pseudoelongatus has broader basidiospores (9.8-11.0 x 6.4-9.5 um vs. 9-11.0 x 5.7-7.4 um) and larger basidia (81-113 x 8-13 um vs. 75-95 x 6-10 um) than C. elongatus. Discussion Among Clavariadelphus species, certain species are saprophytic, while others are symbiotic (Hanif and Khalid 2013). In the present phylogenetic tree, Clade A+B and Clade C correspond respectively to symbiotic species and saprophytic species. Other than four Clavariadelphus species (C. miniatus, C. sachalinensis, C. ligula, and C. griseoclavus), 13 Clavariadelphus species that have an obtuse and fertile apex to the basidiomata were grouped in Clade A with strong support (PP 1, BS 98%). The remaining five Clavariadelphus species, which have a trun- cated and sterile apex, were grouped in Clade B with strong support (PP 1, BS 100%). Clavariadelphus ligula is unique in occupying saprophytic and symbiotic habitats. In the phylogenetic analysis, C. ligula was placed in the Clavariadelphus clade, but it formed an independent lineage separated from other Clavariadel- phus species with strong support (PP 1, BS 100%). These results roughly indicate that this taxon may be transitional from saprophytic to symbiotic environments. Although there are clear morphological differences between Clavariadelphus species and other members of Gomphales (Methven 1990), Clavariadelphus has a close relationship with other genera in Gomphales based on molecular phylo- genetic analysis, such as Ramaria Holmskj6ld, Gomphus, and Lentaria (Pine et al. 1999; Giachini et al. 2010). In particular, some species of Clavariadelphus have similar microscopic features and a saprophytic habitat comparable to Lentaria (Deng et al. 2020; Huang et al. 2020; Li et al. 2025), but few studies have closely investigated the relationship between Clavariadelphus and Lentaria. Therefore, a systematic analysis of the genetic relationship between these two genera is necessary. In the present research, all Clavariadelphus species were grouped in the same subclade except C. ligula. Meanwhile, two species of Lentaria (L. bys- siseda and L. patouillardii) were embedded into Clavariadelphus species, but the statistical support was insufficient to corroborate a close relationship between Clavariadelphus and Lentaria owing to the limited data available. Thus, the rela- tionship between the two genera requires further investigation. The Hengduan Mountains region, a global hotspot of biodiversity, is an ideal habitat for fungi to inhabit, reproduce, and diversify (Mittermeier et al. 2005; Myers et al. 2000). Four newly identified species, C. alpinus, C. amplus, C. aurantiacus, and C. tenuis, were discovered in the Hengduan Mountains MycoKkeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 370 Mei-jia Li et al.: Three new species of Clavariadelphus from China region in 2020, in addition to the three new species described in the present study. This region is rich in Clavariadelphus species diversity. In the future, we predict that a growing number of unnamed Clavariadelphus species will be found in the Hengduan Mountains region. Key to known Clavariadelphus species in China 1 Basidiospores narrowly €llipSOidy-Q.!> 23 ak svsscpacss i ctaeenastovasastulausrn enter von. 2 - Basidiospores broadly ellipsoid to ellipSOid, Q_ < 2.0... ee ceseeeeeteeeeeee 3 a ~Basidiospores:<:1'6:5 pnilong). Qc Ae. Sepa cdeceuarertalieelslccrens C. ligula = s(Basidiospores 16.5. Minin ONG)? Qo Accs esteccrecignceravsviverertpivesiesniverens 4 3. Basidiomata orange; apex sterile, trUNCATE o.oo. cece ceeeestseeeeeeeeaeees 5 - Basidiomata without orange tinge; apex fertile, not truncate ................... 6 4 Basidiomata pale yellow to salmon to purple-drab................... C. miniatus - Basidiomata tawny or light walnut-brown to light brown...C. sachalinensis i =“ Basidiomata-apexs—75-Cl IM- Cl AMete he sscas ders doneseerepaceanderdduaaantocseashereddanes 7 - Basidiomata apex < 2 cm in diamete?l................ccccceeeseeeeeeees C.gansuensis G6 6 “BasidiomatewSually ZOSSOACHWMIGIN dere pan enist pies nnteasshdoinhondibbducs toowoukbeeeebaees 8 =, wBasidlomata.wsyallye << ZOSSHICM si. coe sue rern wibaea tear auras 9 7 Basidiomata yellow or light orange yellow ................:::::ee C. aurantiacus - Basidiomata pinkish orange or grayish orangeé..............::cccsseeee C. amplus 8 Basidiospores narrowly ellipsoid, basidiomata gray-purple....C. elongatus - Basidiospores broadly ellipsoid, basidiomata cinnamon... C. yunnanensis Or BaSidia-ClalMples Si... some sibs, arya. ae anete 2d, er seanaee Baud a, Magnan Ell vert 8 C. tenuis Se AE SICAL AND CCl ay Fizen oy PEI s UNO ST Rea en oa JO, ON ore, a cea 10 10 Basidiomata grayish red to pastel-red ............. ee eeeeeeees C. himalayensis - Basidiomata gray or yellow, without red coloration ..............ccceeeeeseeeees 11 11 Basidiomata gray; basidiospores elliopsoid 10-11 x 5-6.5 um, Q. = 1.89 Sheree eesodut ans elas Ace Moin Neca ae NAcaE 5 TMI Ae Ze lates ANG See ONE C. griseoclavus Py -DaSICIGh iat asy COW... -s.jic-actanin ee greg et eatin eens RR ee neil haa 12 12 Basidiomata yellow; basidiospores broadly ellipsoid, 7.8-9.6 x 5.5- TAM, QOS ASB tiated Schedule oa ill Broth teed is C. alpinus - Basidiomata pale yellow-brown; basidiospores narrowly ellipsoid 9.2- 1230-456 [6 MIM, Oe 1S OFS, tersidan 6 ates tenensstarnus eek pike C. khinganensis 13 Basidiomata pale orange to apricot.......... ee eceeececesteeeeees C. acuminatus - Basidiomata ecru-drab to light purple-drab.................. C. pseudoelongatus Acknowledgements We thank Robert McKenzie, PhD, from Liwen Bianji (Edanz) (www.liwenbianji. cn) for editing the English text of a draft of this manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. MycoKeys 121: 357-374 (2025), DOI: 10.3897/mycokeys.121.158142 37] Mei-jia Li et al.: Three new species of Clavariadelphus from China Use of Al No use of Al was reported. Funding This study was supported by the National Natural Science Foundation of China (No. 31750001) and the National Natural Science Foundation of China (No. 31670015). Author contributions Conceptualization: PZ. Methodology: MJL, PTD. Investigation: MJL, PTD. Resources: PZ, PTD, ZHC, MJL. Writing — original draft: MJL. Writing — review and editing: PZ. Supervi- sion: PZ. Project administration: PZ. Funding acquisition: PZ. 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