<8) MycoKeys MycoKeys 111: 229-248 (2024) DOI: 10.3897/mycokeys.111.125933 Research Article Molecular phylogeny and morphology reveal three new plant pathogenic fungi species (Septobasidiales, Basidiomycota) from China Qianquan Jiang'’”®, Zhengli Kang'?®, Xubo Wang'”®, Changlin Zhao’?© 1 Yunnan Provincial Key Laboratory for Conservation and Utilization of In-forest Resource, the Key Laboratory of Forest Resources Conservation and Utilization in the Southwest Mountains of China Ministry of Education, Southwest Forestry University, Kunming 650224, China 2 College of Forestry, Southwest Forestry University, Kunming 650224, China Corresponding author: Changlin Zhao (fungi@swfu.edu.cn) OPEN Qaccess This article is part of: Diversity, taxonomy, and systematics of macrofungi from tropical Asia Edited by Olivier Raspé, Rui-Lin Zhao, Jennifer Luangsa-ard Academic editor: Rui-Lin Zhao Received: 22 April 2024 Accepted: 28 November 2024 Published: 30 December 2024 Citation: Jiang Q, Kang Z, Wang x, Zhao C (2024) Molecular phylogeny and morphology reveal three new plant pathogenic fungi species (Septobasidiales, Basidiomycota) from China. MycoKeys 111: 229-248. https://doi.org/10.3897/ mycokeys.111.125933 Copyright: © Qianquan Jiang 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 Three new fungal species, Septobasidium macrobasidium, S. puerense and S. wuliang- shanense, are proposed based on a combination of the morphological features and mo- lecular evidence. The taxon S. macrobasidium is characterized by the coriaceous basid- iomata with a cream surface, cylindrical basidia, straight, 4-celled, subglobose or ovoid probasidia and thin-walled, narrowly cylindrical basidiospores with septa, measuring as 7-9 x 3.5-4.5 um, the haustoria consisting of irregularly coiled hyphae; in addition, this fungus was found associated with the insect of Diaspididae. The species S. puerense is characterised by resupinate coriaceous basidiomata with a cinnamon brown to chestnut brown surface, cylindrical or slightly irregular basidia, 2-3-celled, slightly curved, subglo- bose to pyriform probasidia, probasidia cell persistent after the formation of the basidia and the haustoria with two types consisting of irregularly coiled hyphae and spindle-shape. The fungus was found associated with the insect species Pseudaulacaspis pentagona. The species S. wuliangshanense is characterised by the coriaceous basidiomata with a slightly brown surface, cylindrical or slightly irregular basidia, 2-3-celled, straight or slight- ly curved, pyriform, subglobose or ovoid and probasidia, haustoria consisting of irregu- larly coiled hyphae, associated with the insect genus Aulacaspis. Sequences of internal transcribed spacer region (ITS) were analysed maximum likelihood, maximum parsimony and Bayesian inference methods. The new species S. macrobasidium was clustered with S. maesae. Furthermore, S. puerense was retrieved as a sister to S. carestianum. The phy- logenetic tree, inferred from the ITS sequences, highlighted that S. wuliangshanense was the sister to S. aguilariae with strong supports. Application of PHI test to the ITS tree-lo- cus sequences revealed no recombination level within phylogenetically related species. Key words: Forest disease, phylogenetic analyses, taxonomy, wood-inhabiting fungi, Yunnan Province Introduction Fungi are a diverse, monophyletic group of eukaryotes and these organisms show immense ecological and economic impacts for playing an important role in the ecosystems as diverse as soil, trees, with hidden layers within their substrate 229 Qianquan Jiang et al.: Three new pathogenic fungi species from China (James et al. 2020). Approximately 150 thousand species of fungi have been described (Cui et al. 2019; Dai et al. 2021; Wang and Zhao 2021; Sun et al. 2022; Liu et al. 2023). But the potential biodiversity of the group is likely to be 2.2-3.8 millions of species (Blackwell 2011; Taylor et al. 2014; Hibbett 2016; Dong et al. 2023). The diversity for flora of seed plants in Yunnan Province is higher than that in other areas of China. The number of discovered new fungal species to- talled 1,345 from this province from 2000 to 2020 years. Endemic woody plants are rich in Yunnan, supplying rich and varied substrates for wood-decaying fungi. The pathogenic fungi with industrial, medicinal and economic value, comprise most basidiomycetes and ascomycetes growing on various kinds of living trees (Dai et al. 2015, 2021; Vinay et al. 2015; Wu et al. 2019; Luo et al. 2022). Mutualistic and parasitic symbioses between fungi and plants are widely acknowledged to have profound influences on the evolution and ecology of terrestrial life, but less well-known are the symbioses between fungi and in- sects (Henk and Vilgalys 2007). The insects are protected from their enemies by the fungus and the fungus draws its nutriment from, and is distributed by, the insects. Under natural conditions, whereas the fungus cannot live without the insects, the insects are able to live without the fungus, but the life of insects unprotected by the fungus is precarious (Couch 1938). Basidiomycete fungi have evolved many symbiotic associations with plants and animals, but fungi in the order Septobasidiales are the only large group of basidiomycetes that are obligately parasitic on insects. Understanding the evolution of insect parasitism and switches from plant parasitism in the Ba- sidiomycota requires a phylogeny to place the order Septobasidiales within the Pucciniomycotina and to determine whether the different forms of insect parasitism in the order Septobasidiales and Septobasidium Pat. have a single origin. Septobasidium is a type of fungi that has a mutualistic relationship with insects. Although Septobasidium sterilize the individuals they parasitize, the fungi may protect uninfected individuals and thereby benefit the population of scale insects (Couch 1938). All fungi in the order Septobasidiales do not dis- play this type of symbiosis. Some may be wholly parasitic because they do not form substantial protective structures. This fungus-insect symbiosis is import- ant because of its unique altruistic and parasitic characteristics and its phylo- genetic position within the Basidiomycota (Henk and Vilgalys 2007). Septobasidium (Septobasidiaceae, Septobasidiales), erected by Patouillard (1892), typed by Septobasidium velutinum Pat., which is a large cosmopolitan genus characterised by perennial colonies on the surfaces of plant structures with colonies of scale insects, the basidiomata are usually white to cream, yel- lowish brown or brown hymenophore, rarely more brightly coloured, normally resupinate and felty in texture; their surfaces may be smooth, warty, or spiny (Patouillard 1892; Couch 1929); a monomitic hyphal system with simple septa, with or without probasidia, 2-4 celled cylindrical, curved, or straight basidia, ba- sidiospores that are hyaline, thin-walled, smooth and cylindrical or fusiform and haustoria consisting of coiled or spindle-shaped hyphae (Ma et al. 2019). Ina previous study, Couch (1938) divided it primarily based upon morphological characters related to the structure of the basidia and probasidia. Basidia may be curved or straight and may have one, two, three, or four cells. The probasidia either remains as an empty cell at the base of the mature basidia (persistent) or becomes a spore-bearing cell in the mature basidia (not persistent). Other MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 230 Qianquan Jiang et al.: Three new pathogenic fungi species from China characters used as indicators of major groups in Septobasidium include the layered nature of the thallus and the presence of pillar structures (Oberwinkler 1989). Septobasidium occurs on living leaves, stems and branches of a great variety of perennial plants, including gymnosperms, monocots and dicots. Inex- perienced collectors may mistake them for corticioid basidiomycetes or even lichens (Couch 1929, 1935, 1938, 1946). As with most basidiomycetes, basidia are produced so that they project toward the ground. Thus, the resupinate basidiomata often are found on the lower sides of branches. They occur on living rather than dead plant parts and away from the extreme tips of branches which distinguishes them from some resupinate species of Aphyllophorales (Couch 1938; Henk and Vilgalys 2007). About 305 species have been accepted into the genus worldwide (Patouillard 1892; Bresadola and Saccardo 1897; Burt 1916; Lloyd 1919; Couch 1929, 1935, 1938, 1946; Yamamoto 1956; Gomez and Henk 2004; Henk 2005; Lu and Guo 2009a, 2009b, 2010a, 2010b, 2011; Chen and Guo 2011a, 2011b, 2011c, 2012; Lu et al. 2010; Li and Guo 2013, 2014; Ma et al. 2019). Molecular systematics has played a powerful role in inferring phylogenies within fungal groups since the early 1990s (White et al. 1990; Binder et al. 2013; Dai et al. 2015; Choi and Kim 2017). Classification of the kingdom of fungi has been updated continuously, based on the frequent inclusion of data from DNA sequences in many phylogenetic studies (Wijayawardene et al. 2020). Howev- er, molecular studies involving Septobasidium are rare (Henk and Vilgalys 2007; Zhao et al. 2017). One phylogenetic study of a single origin of insect symbiosis in the class Pucciniomycetes suggested that there is little or no support for Septobasidium as a monophyletic group (Henk and Vilgalys 2007). The previ- ous study introduced a six-gene phylogenetic overview of Basidiomycota and allied phyla and confirmed that S. carestianum Bres. nested within the order Septobasidiales and grouped with Helicobasidium mompa Nobuj. Tanaka and Thanatophytum crocorum (Pers.) Nees (Zhao et al. 2017). During investigations into the wood-inhabiting fungi in Pu’er, Yunnan of Chi- na, samples representing three additional species belonging to genus Septo- basidium were collected. Three new Septobasidium taxa were found that could not be assigned to any described species. To clarify the placement and rela- tionships of these specimens, we carried out a phylogenetic and taxonomic study based on the ITS sequences. A description, illustrations, and phylogenet- ic analysis results of the new species are provided. Materials and methods Sample collection and herbarium specimen preparation Fresh fruiting bodies of fungi growing on angiosperm branches were collected from Pu’er of Yunnan Province, P.R. China. The samples were photographed in situ and fresh macroscopic details were recorded. Photographs were recorded using a Jianeng 80D camera (Tokyo, Japan). All of the photos were stacked and merged using Helicon Focus Pro 7.7.5 software. Specimens were dried in an electric food dehydrator at 40 °C and then sealed and stored in an enve- lope bag and deposited in the herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, PR. China. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 231 Qianquan Jiang et al.: Three new pathogenic fungi species from China Morphology Macromorphological descriptions are based on field notes and photos cap- tured in the field and lab. Color terminology follows Petersen (1996). Micro- morphological data were obtained from the dried specimens when observed under a light microscope following the previous study (Ma et al. 2019). The following abbreviations are used: 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). DNA extraction and sequencing Genomic DNA was obtained from dried specimens using the EZNA HP Fungal DNA Kit, according to the manufacturer's instructions with some modifications. A small piece of dried fungal specimen (about 30 mg) was ground to powder with liquid nitrogen. The powder was transferred to a 1.5 ml centrifuge tube, suspended in 0.4 ml of lysis buffer and incubated in a 65 °C water bath for 60 min. After that, 0.4 ml phenol-chloroform (24: 1) was added to each tube and the suspension was shaken vigorously. After centrifugation at 13,000 rpm for 5 min, 0.3 ml supernatant was transferred to a new tube and mixed with 0.45 ml binding buffer. The mixture was then transferred to an adsorbing column (AC) for centrifugation at 13,000 rpm for 0.5 min. Then, 0.5 ml inhibitor removal fluid was added in AC for a centrifugation at 12,000 rpm for 0.5 min. After washing twice with 0.5 ml washing buffer, the AC was transferred to a clean centrifuge tube and 100 ml elution buffer was added to the middle of adsorbed film to elute the genomic DNA. ITS region was amplified with primer pairs ITS5 and ITS4 (White et al. 1990). The PCR procedure was as follows: initial denaturation at 95 °C for 3 min; followed by 35 cycles of 94 °C for 40 s, 58 °C for 45 s and 72 °C for 1 min; and a final extension of 72 °C for 10 min. The PCR products were purified and directly sequenced at barium Tsingke Biological Technology Limited Company. All newly generated sequences were deposited at GenBank (https://www.ncbi.nim.nih.gov/genbank/) (Table 1). Phylogenetic analyses The sequences were aligned in MAFFT version 7 using the G-INS-i strategy (Ka- toh et al. 2019). The alignment was manually adjusted using AliView version 1.27 (Larsson 2014). Sequences of Helicobasidium mompa and Pachnocybe ferruginea Berk. obtained from GenBank were used as an outgroup to root trees following Henk and Vilgalys (2007) in the ITS analysis (Fig. 1). Maximum parsimony (MP), maximum likelihood (ML) and Bayesian Infer- ence (BI) analyses were applied to the three combined datasets. The phylo- genetic analysis method was adopted by previous study (Zhao and Wu 2017; Yang et al. 2023; Dong et al. 2024). MP analysis was performed in PAUP* ver- sion 4.0610 (Swofford 2002). All of the characteristics 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 addi- tions. Max-trees were set to 5,000, branches of zero length were collapsed and MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 232 Qianquan Jiang et al.: Three new pathogenic fungi species from China Table 1. List of species, specimens and GenBank accession numbers of sequences used in this study. [* Indicates type materials]. Species Helicobasidium mompa Pachnocybe ferruginea Septobasidium alni S. aquilariae S. aquilariae S. aquilariae S. aquilariae S. aquilariae S. arachnoideum S. bogoriense S. broussonetiae S. burtii S. canescens S. carestianum S. castaneum S. cavarae S. fumigatum S. gomezii S. grandisporum S. griseum S. hainanense S. kameii S. macrobasidium S. macrobasidium S. maesae S. marianiae S. marianiae S. michelianum S. pallidum S. pilosum S. pinicola S. pseudopedicellatum S. puerense S. puerense S. ramorum S. septobasidioides S. sinuosum S. taxodii S. velutinum S. westonii S. wilsonianum S. wuliangshanense S. wuliangshanense MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 Specimen No. DAH h1 DAH pf1 DAH FP3 CLZhao 6610 CLZhao 6611 CLZhao 6612 CLZhao 6613 CLZhao 6614 DAH 025 998434 998436 DAH 062 DAH 323 DJM 644 DAH 052 DJM FP1 DAH 005 DAH 031 DAH 065 DAH 016 998437 998432 CLZhao 9624* CLZhao 9658 998433 LJF 7006 DAH 283b DAH FP5 998435 DAH 020 DAH 013 DAH 044 CLZhao 9430* CLZhao 4298 DAH 045a DAH 032 DAH 036 DAH 194C DAH 024 DAH FP2001 DAH 037 CLZhao 5809* CLZhao 3666 GenBank accession No. ITS DQ241472 DQ241473 DQ241441 MK804520 MK804521 MK804522 MK804523 MK804524 DQ241443 HM209414 HM209416 DQ241444 DQ241446 DQ241448 DQ241447 DQ241445 DQ241451 DQ241462 DQ241453 DQ241454 HM209417 HM209412 PP532758 PP532759 HM209413 MK809161 DQ241456 DQ241457 HM209415 DQ241458 DQ241459 DQ241460 PP532760 PP532761 DQ241450 DQ241461 DQ241464 DQ241466 DQ241467 DQ241468 DQ241469 PP532756 PP532757 Preference Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Ma et al. 2019 Ma et al. 2019 Ma et al. 2019 Ma et al. 2019 Ma et al. 2019 Henk and Vilgalys 2007 Lee et al. 2023 Unpublished Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Lee et al. 2023 Lee et al. 2023 Present study Present study Unpublished Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Unpublished Henk and Vilgalys 2007 Henk and Vilgalys 2007 Lee et al. 2023 Present study Present study Lee et al. 2023 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Lee et al. 2023 Henk and Vilgalys 2007 Henk and Vilgalys 2007 Present study Present study 233 Qianquan Jiang et al.: Three new pathogenic fungi species from China A Type species @ Persistent @ Not persistent = & Coiled ® Spindle ™ Absent @ Present - /100/0.95 99/100/1.00| Probasidia Haustoria Pillars 100/100/1.00 r= Septobasidium sinuosum DAH 036 ) 7 a 87/91/1.00 Septobasidium griseum DAH 016 & ¢ = Septobasidium fumigatum DAH 005 C) ¢ | ~168/0.99 Septobasidium grandisporum DAH 065 C C2 ie] 95/99/1.00) Septobasidium westonii DAH FP2001 t) ¢ | a Septobasidium septobasidioides DAH 032 @ 2 i_ ; Septobasidium gomezii DAH 031 ® ¢e L_ Septobasidium castaneum DAH 052 C ) ¢ L_ Septobasidium canescens DAH 323 © ¢ = =/>0ee Septobasidium burtii DAH 062 @ + = 92/98/1,007— Septobasidium pilosum DAH 020 c) e [-] Septobasidium arachnoideum DAH 025 @ e iat 5/9 10:38 L - /58/1.00) Septobasidium wilsonianum DAH 037 C) e | Lavinia Septobasidium bogoriense 998434 & & a Loo/100/ roo Septobasidium broussonetiae 998436 c * fe) Septobasidium taxodii DAH 194C c ) ¢ a Septobasidium pinicola DAH 013 @ e = Septobasidium michelianum DAH FP5 C ) C2 w Septobasidium ramorum DAH 045a C ) e TT] 100/100/1.00 | Septobasidium alni DAH FP3 © 4 =) Septobasidium kameii 998432 C) ¢ _] py Septobasidium marianiae LIF 7006 ; 94/99/1.00 Septobasidium marianiae DAH 283b e ¢ _ N Septobasidium velutinum DAH 024 A C ) 2 ia ‘Septobasidium puerense CLZhao 9430 uM C) x 2 | Septobasidium puerense CLZhao 4298 Septobasidium carestianum DJM 644 so] + L_| Septobasidium pallidum 998435 @ e aa Septobasidium cavarae DJM FP1 @ G Septobasidium aquilariae CLZhao 6610 Septobasidium aquilariae CLZhao 6614 Septobasidium aquilariae CLZhao 6611 e e = Septobasidium aquilariae CLZhao 6613 Septobasidium aquilariae CLZhao 6612 Septobasidium wuliangshanense CLZhao 5809 C) e "Lb Septobasidium wuliangshanense CLZhao 3666 ; = Septobasidium pseudopedicellatum DAH 044 ie e fe 100/100/Loq Septobasidium macrobasidium CLZhao 9658 Septobasidium macrobasidium CLZhao 9624 e ¢ = Septobasidium maesae 998433 ( } CD es Septobasidium hainanense 998437 i 10 Figure 1. Maximum parsimony strict consensus tree illustrating the phylogeny of the three new species and related species in Septobasidium. Branches are labelled with maximum likelihood bootstrap values = 70%, parsimony bootstrap values = 50% and Bayesian posterior probabilities = 0.95, respectively. all most-parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1,000 replicates (Felsenstein 1985). Descriptive tree statistics tree length (TL), the consistency index (Cl), the retention index (RI), the rescaled consistency index (RC) and the homoplasy index (HI) were calculated for each most-parsimonious tree generated. ML was inferred using RAxML-HPC2 through the CIPRES Science Gateway (Miller et al. 2012). Branch support (BS) for ML analysis was determined using 1,000 bootstrap replicates and evaluated under the gamma model. MrModeltest 2.3 (Posada and Crandall 1998; Nylander 2004) was used to determine the best-fit evolution model for each data set for Bayesian inference (BI). Bayesian inference was calculated with MrBayes3.1.2 with a general time reversible (GTR+G+l) model of DNA substitution and a gamma distribution rate variation across sites (Ronquist et al. 2012). Four Markov chains were run for 2 runs from random starting trees for 2.9 million generations (Fig. 1) and trees were sampled every 100 generations. The first one-fourth generations were dis- carded as burn-in. A majority rule consensus tree of all remaining trees was cal- culated. Branches were considered as significantly supported if they received maximum likelihood bootstrap (ML) = 70%, maximum parsimony bootstrap (MP) = 50%, or Bayesian posterior probabilities (PP) = 0.95. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 234 Qianquan Jiang et al.: Three new pathogenic fungi species from China Pairwise homoplasy test The Genealogical concordance phylogenetic species recognition analysis (GCPSR) is a tool used to check significant recombinant events. The data were analysed us- ing SplitsTree 4 with the pairwise homoplasy ®w, PHI test to determine the recom- bination level within closely related species (Bruen et al. 2006; Huson and Bryan 2006; Quaedvlieg et al. 2014). One-locus dataset (ITS) with closely related species were used for the analyses. PHI results lower than 0.05 (®w < 0.05) indicates a sig- nificant recombination is present in the dataset. The relationships between closely related taxa were visualised by constructing split graphs from the concatenated datasets, using the LogDet transformation and splits decomposition options. Result Molecular phylogeny The dataset based on ITS (Fig. 1) comprises sequences from 43 fungal spec- imens representing 35 species. The alignment length of this dataset is 474 characters, of which 267 characters are constant, 45 characters are variable with no information and 162 characters have no information. Maximum parsi- mony analysis yielded three equally parsimonious trees (TL = 967, Cl = 0.3516, HI = 0.6484, RI = 0.4543, RC = 0.1597). Bayesian analysis and ML analysis re- sulted in a similar topology as MP analysis with an average standard deviation of split frequencies of 0.007729 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 861. The phylogenetic tree (Fig. 1), inferred from the ITS sequences, highlighted that Septobasidium macrobasidium was clustered with S. maesae C.X. Lu & L. Guo. Fur- thermore, S. puerense was retrieved as a sister to S. carestianum. The new species S. wuliangshanense was the sister to S. aquilariae C.L. Zhao with strong supports. Application of PHI test to the ITS tree-locus sequences revealed no recombi- nation level within phylogenetically related species. The test results of ITS se- quence dataset show ®w = 0.5697 (®w > 0.05) no recombination is present in the new species Septobasidium wuliangshanense with S. aquilariae, S. cavarae Bres. and S. pseudopedicellatum Burt (Fig. 2). Septobasidium wuliangshanense CLZhao 3666 Septobasidium wuliangshanense CLZhao 5809 Septobasidium aquilariae CLZhao 6610 Septobasidium aquilariae CLZhao 6614 Septobasidium pseudopedicellatum DAH 044 Septobasidium cavarae DJM FP1 -——+0.01 Figure 2. Split graphs showing the results of PHI test for the ITS data of Septobasidium wuliangshanense and closely relat- ed taxa using LogDet transformation and splits decomposition. PHI test results ®w < 0.05 indicate that there is significant recombination within the dataset. New taxa are in red while closely related species to new species are in other colours. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 235 Qianquan Jiang et al.: Three new pathogenic fungi species from China Taxonomy Septobasidium macrobasidium Q.Q. Jiang & C.L. Zhao, sp. nov. MycoBank No: 853680 Figs 3, 4 Holotype. CHINA * Yunnan Province, Pu’er, Jingdong County, Wuliangshan Na- tional Nature Reserve, 24°29'17"N, 100°40'27'E, altitude: 1800 m a.s.l., on the living tree of angiosperm, leg. C.L. Zhao, 6 January 2019, CLZhao 9624 (SWFC). Diagnosis. Differs from other Septobasidium species by the coriaceous ba- sidiomata with a cream surface, cylindrical basidia (48.5-83 x 6.5-13 um), and thin-walled, narrowly cylindrical basidiospores with septa, measuring as 7-9 x 3.5-4.5 um. Etymology. Macrobasidium (Lat.): refers to the larger basidia of the type specimen. Description. Basidiomata perennial, resupinate, easy to separate from sub- strate, coriaceous upon drying, up to 9 cm long, 5 cm wide, 2 mm thick. Hy- menial surface smooth, slightly cream when fresh, cream upon drying. Sterile margin narrow, white to cream, up to 0.5 mm. Hyphal system monomitic, generative hyphae with simple septa, pale brown, thick-walled, frequently branched, interwoven. In section 850-1850 um thick; subiculum pale brown, 30-80 um thick; pillars brown, 350-550 um high, 85- 200 um wide, hyphae of pillars 2-5 um thick, hyaline or brown, forming 2-3 horizontal layers. Basidia arising directly from the generative hyphae, basidia cylindrical, straight, 4-celled, 48.5—83 x 6.5-13 um, colourless. Probasidia subglobose or ovoid, 13-23 x 7—20.5 um, colourless, persistent. Haustoria consisting of irreg- ularly coiled hyphae. Basidiospores narrowly cylindrical, colourless, thin-walled, with septa, (6.5-)7— 9(-9.5) x (3-)3.5-4.5(—5) um, L = 8.09 um, W = 3.94 um, Q = 1.55-2.83 (n = 60/2). Habitat and distribution. Growing on the plant Betulaceae Gray, associated with the insect of Diaspididae Ferris. Additional specimen examined (paratype). CHINA * Yunnan Province, Pu'er, Jingdong County, Wuliangshan National Nature Reserve, 24°29'17'N, 100°40'27'E, altitude: 1800 m a.s.I., on the living tree of angiosperm, leg. C.L. Zhao, 6 January 2019, CLZhao 9658 (SWFC). Septobasidium puerense Q.Q. Jiang & C.L. Zhao, sp. nov. MycoBank No: 853681 Figs 5, 6 Holotype. CHINA * Yunnan Province, Pu’er, Jingdong County, The Forest of Pine- apple, 24°30'58"N, 100°52'31"E, altitude: 2000 m a.s.I., on the living tree of an- giosperm, leg. C.L. Zhao, 4 January 2019, CLZhao 9430 (SWFC). Diagnosis. Differs from other Septobasidium species by a cinnamon brown to chestnut brown surface, subglobose to pyriform probasidia (10.5- 19.5 x 5.5-9 um), and two types of haustoria consisting of irregularly coiled hyphae and spindle-shape. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 236 Qianquan Jiang et al.: Three new pathogenic fungi species from China D probasidia E basidia F basidiospore G haustoria H hyphae | scale insect on branches. Scale bars: 1 cm (A); 1 mm (B); 100 um (C); 10 um (D-H); 1 mm (I). MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 237 Qianquan Jiang et al.: Three new pathogenic fungi species from China fn a KOO an > eld C1 GID GRIGOA \N I ES © Figure 4. Microscopic structures of Septobasidium macrobasidium (holotype, SWFC 9624) A basidiospores B generative hyphae from hyphal layer C probasidia and basidia. Scale bars: 5 um (A); 10 um (B, C). Etymology. Puerense (Lat.): refers to the locality (Pu’er) of the type specimen. Description. Basidiomata perennial, resupinate, hard to separate from sub- strate, coriaceous upon drying, up to 15 cm long, 1 cm wide,1 mm thick. Hy- menial surface smooth, pale brown when fresh, cinnamon brown to chestnut brown upon drying. Sterile margin slightly brown, up to 1 mm. Hyphal system monomitic, generative hyphae with simple septa, pale brown, thick-walled. In section 380-650 um thick; subiculum pale brown, 10-30 um thick; pillars brown, 170-380 um high, 40-85 um wide, hyphae of pillars 1.5- 3.5 um thick, colorless, with closely packed parallel upright threads, forming 2-3 horizontal layers. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 238 Qianquan Jiang et al.: Three new pathogenic fungi species from China 3 1 — .s ’ eS e Figure 5. Septobasidium puerense (holotype, CLZhao 9430) A, B basidiomata on branch C sections of basidiomata D probasidia E basidia (arrow) F hyphae G haustoria consisting of irregularly coiled hyphae H the spindle-shaped hausto- ria I, J scale insect on branches. Scale bars: 1 cm (A); 1 mm (B); 100 um (C); 10 um (D-H); 1 um (I). MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 239 Qianquan Jiang et al.: Three new pathogenic fungi species from China Figure 6. Microscopic structures of Septobasidium puerense (holotype, CLZhao 9430) A probasidia and basidia B gener- ative hyphae from hyphal layer. Scale bars: 10 um (A, B). Basidia arising directly from the generative hyphae, cylindrical or slightly ir- regular, slightly curved, 2-3-celled, 17-30.5 x 3-6.5 um, colourless. Probasidia subglobose to pyriform, 10.5-19.5 x 5.5—9 um, colorless, probasidia cell per- sistent after the formation of the basidia. Basidiospores not seen. Haustoria with two types: 1) consisting of irregularly coiled hyphae; 2) spindle-shape. Habitat and distribution. Growing on the plant Berberidaceae Juss, associ- ated with the insect species Pseudaulacaspis pentagona (Targioni Tozzetti). Additional specimen examined (paratype). CHINA * Yunnan Province, Puer, Jingdong Country, Wuliangshan National Nature Reserve, 24°29'17'N, 100°40'27"E, altitude: 1800 m a.s.I., on the living tree of angiosperm, leg. C.L. Zhao, 5 October 2017, CLZhao 4298 (SWFC). Septobasidium wuliangshanense Q.Q. Jiang & C.L. Zhao, sp. nov. MycoBank No: 853679 Figs 7,8 Holotype. CHINA * Yunnan Province, Pu’er, Zhenyuan County, Heping Town, Li- angzi Village, Wuliangshan National Nature Reserve, 24°29'17"N, 100°40'27'E, altitude: 1860 m a.s.I., on the living tree of angiosperm, leg. C.L. Zhao, 15 Janu- ary 2018, CLZhao 5809 (SWFC). Diagnosis. Differs from other Septobasidium species by the coriaceous ba- sidiomata with a slightly brown surface, cylindrical or slightly irregular basidia, pyriform to subglobose or ovoid probasidia (7.5-13 x 4.5—9 um), and the haus- toria consisting of irregularly coiled hyphae. Etymology. Wuliangshanense (Lat.): refers to the locality (Wuliangshan) of the type specimen. Description. Basidiomata perennial, resupinate, easy to separate from sub- strate, coriaceous upon drying, up to 10 cm long, 2 cm wide, 1 mm thick. Hy- menial surface smooth, cream to pale brown when fresh, slightly brown upon drying. Sterile margin cream to slightly brown, up to 2 mm. MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 240 Qianquan Jiang et al.: Three new pathogenic fungi species from China Figure 7. Septobasidium wuliangshanense (holotype, CLZhao 5809) A, B basidiomata on branch C sections of basidi- omata D probasidia (arrow) E basidium (arrow) F hyphae G haustoria H, I scale insect on branches. Scale bars: 1 cm (A); 1 mm (B); 100 um (C); 10 um (D-G); 1 mm (H); 10 mm (I). MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 241 Qianquan Jiang et al.: Three new pathogenic fungi species from China Figure 8. Microscopic structures of Septobasidium wuliangshanense (holotype, CLZhao 5809) A probasidium and basid- ia B generative hyphae from hyphal layer. Scale bars: 10 um (A, B). Hyphal system monomitic, generative hyphae with simple septa, pale brown, thick-walled. In section 660-1200 um thick; subiculum pale brown, 20-50 um thick; pillars brown, 150-300 um high, 30-150 um wide, hyphae of pillars 3-4 um thick, brown, forming 2-3 horizontal layers. Basidia arising directly from the generative hyphae, cylindrical or slightly ir- regular, colourless, straight or slightly curved, 2-3-celled, 21.5-29 x 5.5-9 um. Probasidia pyriform, subglobose or ovoid, 7.5-13 x 4.5—9 um, colorless, proba- sidia cell persistent after the formation of the basidia. Haustoria consisting of irregularly coiled hyphae. Basidiospores not seen. Habitat and distribution. Growing on the plant Fagaceae Dumort, associated with the insect genus Aulacaspis Cockerell. Additional specimen examined (paratype). CHINA * Yunnan Province, Pu’er, Jing- dong County, Wuliangshan National Nature Reserve, in association with the genus Aulacaspis on Rosaceae, 24°29'17'N, 100°40'27’E, altitude: 1800 m a.s.l., on the living tree of angiosperm, leg. C.L. Zhao, 2 October 2017, CLZhao 3666 (SWFC). Discussion Many recently described wood-inhabiting fungal taxa have been reported world- wide, including in the genus Septobasidium (Patouillard 1892; Bresadola and Sac- cardo 1897; Burt 1916; Lloyd 1919; Couch 1929, 1935, 1938, 1946; Yamamoto 1956; Gomez and Henk 2004; Henk 2005; Chen and Guo 2009a, 2009b, 2010, 2011a, 2011b, 2011c, 2012; Lu and Guo 2009a, 2009b, 2010a, 2010b 2011, 2013, 2014; Lu et al. 2010). The diversity of Septobasidium is rich in China. Prior to this study, the following 58 Septobasidium species were reported from China, espe- cially in subtropics and tropics (Lu and Guo 2009a, 2009b, 2010a, 2010b, 2011; Lu et al. 2010; Chen and Guo 2011a, 2011b, 2011c, 2012; Li and Guo 2013, 2014; Ma etal. 2019). Several Septobasidium species have been described from Yunnan Province (Lu and Guo 2010b, 2011; Ma et al. 2019). In the present study, three new species, S. macrobasidium, S. puerense and S. wuliangshanense are described based on phylogenetic analyses and morphological characteristics. In addition, MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 242 Qianquan Jiang et al.: Three new pathogenic fungi species from China the PHI test (Fig. 2) was carried out to confirm that there is no recombination pres- ent in the new species S. wuliangshanense compared with closely related taxa. Based on ITS topology (Fig. 1), the phylogenetic tree includes the type spe- cies Septobasidium velutinum, which is collected from Costa Rica and most species of this genus have persistent probasidia, except for S. aquilariae, S. gomezii Henk, S. hainanense C.X. Lu & L. Guo, S. pallidum Couch, S. septo- basidioides (Henn.) Hohn. & Litsch. and S. westonii Couch. Most Septobasidi- um species have irregularly coiled haustoria and the spindle-shaped haustoria of S. fumigatum Burt, S. griseum Couch, S. grandisporum Couch, S. pilosum Boedijn & B.A. Steinm and S. sinuosum Couch. However, S. puerense has two types of haustoria: 1) consisting of irregularly coiled hyphae; 2) spindle-shape. Most Septobasidium species have pillars, except for S. arachnoideum (Berk. & Broome) Bres., S. burtii Lloyd, S. canescens Burt, S. cavarae, S. fumigatum, S. grandisporum, S. michelianum (Caldesi) Pat., S. pilosum, S. pinicola Snell, S. sinuosum, S. taxodii Couch and S. wilsonianum L.D. Gomez & Kisim.-Hor. Phylogenetically, based on the ITS topology (Fig. 1), Septobasidium macroba- sidium is clustered with S. maesae. The new species S. puerense is closely relat- ed to S. carestianum and S. wuliangshanense is sister to S. aquilariae with strong supports. But morphologically S. maesae differs from S. macrobasidium by its greyish-brown hymenial surface and smaller basidia (28-55 x 7.5-11.5 um), brown, fusiform basidiospore without septa (18-19.5 x 4—5 um; Lu and Guo 2009a). The species S. carestianum differs from S. puerense by its iuventute avellaneas deinde cinnamomeas, superficiei sub lente pruinosula hymenial surface and the larger sphaerica probasidia (15.1-11.3 um) and 4-celled, api- cem acuta, longer cylindrica basidia in senectute brunnea (62-71 x 5-6.7 um), ellipticae and flexae sporae (21-23 x 4.2-5 um; Gdmez and Henk 2004). The species S. aguilariae differs from S. wuliangshanense by its smaller basidia (15-26.5 x 4-6 um) and without a probasidia cell and reniform basidiospores (11-19 x 4-7.5 um), habitat and distribution growing in association with Pseu- daulacaspis sp. on Aquilaria sinensis (Lour.) Spreng. (Ma et al. 2019). Further, application of PHI test to the ITS tree-locus sequences revealed no recombina- tion level within phylogenetically in these two species (Fig. 2). Morphologically, Septobasidium cokeri Couch. differs from S. macrobasidi- um by its pure white hymenial surface and restricted growth on Quercus rubra L. (GOmez and Henk 2004). The species S. hainanense differs in its purple hy- menial surface and smaller basidia (25-36 x 7-13 um; Lu and Guo 2010a). The taxon S. maesae differs by its perennial basidiomata peeled off after maturity and smaller basidia (28-55 x 7.5-11.5 um; Lu and Guo 2009a). Septobasidium guangxiense Wei Li bis & L. Guo differs from S. puerense in its yellowish brown hymenial surface with numerous fissures at maturity and larger basidia (27-38 x 5-10 um; Li and Guo 2014). The species S. hoveniae Wei Li bis, S.Z. Chen, L. Guo & Yao Q. Ye differs in its cinnamon-brown hymenial surface and growth on Hovenia acerba (Li and Guo 2013; S. polygoni C.X. Lu & L. Guo differs in its white to cinnamon-brown hymenial surface and growth on Polygonum campan- ulatum Hook. f. (Lu and Guo 2010b); S. reevesiae S.Z. Chen & L. Guo differs in its thicker section (1.65-2.20 mm) and larger basidia (37-55 x 8-13 um) and growth on Reevesia longipetiolata Merr. & Chun (Chen and Guo 2012) and S. transversum Wei Li bis & L. Guo differs in its cinnamon-brown basidiomata, its transverse layer at the pillar bases and larger basidia (42-60 x 9-16 um; Li and Guo 2014). MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 243 Qianquan Jiang et al.: Three new pathogenic fungi species from China Morphologically, several species of Septobasidium broussonetiae C.X. Lu, L. Guo & J.G. Wei, S. brunneum Wei Li bis & L. Guo, S. capparis S.Z. Chen & L. Guo, S. eu- ryae-groffii C.X. Lu & L. Guo, S. fissuratum Wei Li bis & L. Guo and S. gaoligongense C.X. Lu & L. Guo are similar to S. wuliangshanense were found in China. However, S. broussonetiae is distinguished by its cracking basidiomata and growth on Brous- sonetia papyrifera (L.) UHér. ex Vent. (Lu et al. 2010); S. brunneum differs in its pur- ple-brown hymenial surface with many cracks and growth on Eurya sp. (Li and Guo 2014); S. capparis differs by its thicker section (less than 2 mm thick) and larger basidia (45-56 x 8-12 um; Chen and Guo 2012); S. euryae-groffii is distinguished by its cinnamon to chestnut brown hymenial surface and growth on Eurya groffii (Lu and Guo 2010b); S. fissuratum differs in its larger basidia (32-45 x 6-9 um) and growth on Castanea sp. (Li and Guo 2013); S. gaoligongense differs in its dark brown hymenial surface and thinner section (260-580 um; Lu and Guo 2010b). Based on our phylogenetic and morphological research results, 61 species have been reported from China, including newly described in the present study and other recently published papers in this country (G6mez and Henk 2004; Lu and Guo 2009a, 2009b, 2010a, 2010b, 2011; Lu et al. 2010; Chen and Guo 2011a, 2011b, 2011c, 2012; Li and Guo 2013, 2014; Ma et al. 2019). It seems that the species diversity of Septobasidium is rich in China. Although the taxa of Septobasidium are well studied in the present paper and the species diversity, taxonomy and phylogeny of Septobasidium and related genera are still unre- solved. A comprehensive study on this issue is urgently needed. 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 (Project Nos: 32170004, U2102220), Forestry Innovation Programs of Southwest Forestry Univer- sity (Project No: LXXK-2023Z07) and the Yunnan Province College Students Innovation and Entrepreneurship Training Program (Project no. $202310677041; s202310677028). Author contributions Conceptualisation, CZ and QJ; methodology, CZ and QJ; software, CZ, QJ and ZK; valida- tion, CZ and QU; formal analysis, CZ and QJ; investigation, CZ and QJ; resources CZ; writ- ing — original draft preparation, CZ, QU, ZK and XW; writing — review and editing, CZ and QU; visualisation, CZ and QJ; supervision, CZ and QJ; project administration, CZ; funding ac- quisition, CZ. All authors have read and agreed to the published version of the manuscript. Author ORCIDs Qianquan Jiang © https://orcid.org/0009-0003-0644-3638 Zhengli Kang © https://orcid.org/0009-0003-1878-6822 Xubo Wang ® https://orcid.org/0000-0001-5996-6027 Changlin Zhao ® https://orcid.org/0000-0002-8668-1075 MycoKeys 111: 229-248 (2024), DOI: 10.3897/mycokeys.111.125933 244 Qianquan Jiang et al.: Three new pathogenic fungi species from China Data availability All of the data that support the findings of this study are available in the main text. 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