<8) MycoKeys MycoKeys 121: 21-65 (2025) DOI: 10.3897/mycokeys.121.155432 Research Article Species diversity and systematic taxonomy of Sarcosomataceae (Ascomycota, Pezizales), with an emphasis on subtropical regions of China Jin Rong Liu'?®, Deng Li®, Si Ang Chen*®, Yan Cheng Zhang'®, Guang Fu Mou", Yan Liu™®, Guang Rong Zhou®®, Zhou Rong Tan®, Jian Hua Zhang’”® 1 Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrai, Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, Guilin 541006, Guangxi, China ND oO FSF W DY College of Life Science, Guangxi Normal University, Guilin 541006, Guangxi, China College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou, 350002, China College of Agriculture, Fujian Agriculture and Forestry University, Fuzhou 350002, China Managing Bureau of Damingshan National Nature Reserve, Wuming 530114, Guangxi, China Managing Bureau of Huaping National Nature Reserve, Guilin 531199, Guangxi, China Managing Bureau of Mao’ershan National Nature Reserve, Guilin 541316, Guangxi, China Corresponding authors: Guang Fu Mou (guangfuMOU@gxib.cn); Yan Liu (gxibly@163.com) OPEN Qaccess This article is part of: Exploring the Hidden Fungal Diversity: Biodiversity, Taxonomy, and Phylogeny of Saprobic Fungi Edited by Samantha C. Karunarathna, Danushka Sandaruwan Tennakoon, Ajay Kumar Gautam Academic editor: Danushka Sandaruwan Tennakoon Received: 10 April 2025 Accepted: 9 July 2025 Published: 21 August 2025 Citation: Liu JR, Li D, Chen SA, Zhang YC, Mou GF, Liu Y, Zhou GR, Tan ZR, Zhang JH (2025) Species diversity and systematic taxonomy of Sarcosomataceae (Ascomycota, Pezizales), with an emphasis on subtropical regions of China. MycoKkeys 121: 21-65. https://doi. org/10.3897/mycokeys.121.155432 Copyright: © Jin Rong Liu 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 The aim of this study is to update the species diversity of Sarcosomataceae in sub- tropical China. Taxonomic and phylogenetic studies were carried out through ecological observation, morphological examination, and molecular phylogenetic analyses inferred from five molecular loci (ITS + nrLSU + rpb7 + rpb2 + tef1-a). By using Bayesian inference (BI) and maximum likelihood (ML) methods, six lignicolous species were identified, in- cluding four novel taxa: Plectania damingshanensis sp. nov., Plectania huapingensis sp. nov., Pseudoplectania aureonigrescens sp. nov., and Urnula auricularioides sp. nov.; and two new distribution records: Plectania lutea and Urnula ailaoshanensis. Keys to species of Plectania, Pseudoplectania, and Urnula were provided. Key words: New taxa, phylogeny, Plectania, Pseudoplectania, subtropical fungus, Urnula Introduction Sarcosomataceae Kobayasi is regarded as an apothecial family of Pezizales within Ascomycota. This family has been studied for many decades, and its systematics could even date back to the 19" century, spanning more than 170 years. Kobayasi (1937) originally proposed Sarcosomataceae (originally writ- ten as “Sarcosomaceae”) as an independent family based on its dark apothecia and highly gelatinized tissues, typified by the genus Sarcosoma Casp. How- ever, based on the color of ascomata and the presence of subcystidia, Le Gal (1946) proposed a preliminary classification with an invalid family without diag- nosis, Sarcoscyphaceae Le Gal ex Eckblad, further divided into two tribes: the bright-colored tribe Sarcoscypheae Le Gal and the brown-black tribe Urnuleae 21 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Le Gal. Subsequently, this single family was validated by Eckblad. Following the transfer of Sarcosoma into Urnuleae, the family Sarcosomataceae was tem- porarily considered a synonym (Eckblad 1968). Notably, taxa that primarily be- longed to the tribe Urnuleae have now been classified under Sarcosomataceae. In 1970, Korf proposed an ameliorative concept to fit the natural classifi- cation of these two families and divided Sarcosomataceae into two tribes: Galielleae Korf and Sarcosomatae Korf. With the restoration of Sarcosoma- taceae as an independent family, ten genera were successively recognized and classified into this family: Chorioactis Kupfer ex Eckblad, Desmazierel- la Libert, Galiella Nannf. & Korf, Nannfeldtiella Eckblad, Neournula Paden & Tylutki, Plectania Fuckel, Pseudoplectania Fuckel, Sarcosoma, Urnula Fr., and Wolfina Seaver ex Eckblad (Eckblad 1968; Korf 1970, 1972, 1973). Previous studies (Samuelson 1975; Samuelson et al. 1980; Brummelen 1981) have de- tailed the characteristics of the suboperculum and operculum in the asci of various members of Sarcosomataceae, including Galiella, Pseudoplectania, Sarcosoma, and Urnula. These studies indicated that these fungi exhibit the greatest morphological similarity to the non-eccentric type, particularly in their dehiscent structure, which features a thin-walled central cylinder mea- suring between 0.5 um and 0.8 um. Additionally, the apical annulus is typically thinner than the lateral wall, distinguishing these characteristics from those observed of Sarcoscyphaceae species. Furthermore, other research focusing on septal structures and spore ontogeny, utilizing numerical analysis meth- ods and chemotaxonomic markers, has provided additional support for Korf's perspective (Li and Kimbrough 1995, 1996; Képcke et al. 2002). With the rapid development of molecular techniques and their application in the taxonomy of macrofungi (Licking et al. 2021), DNA sequence data have been widely used in systematic studies of Sarcosomataceae. Harrington et al. (1999) used 18S rDNA sequences to conduct an in-depth exploration of the in- tergeneric relationships within Discomycetes, including 10 genera of Sarcoso- mataceae and 11 genera of Sarcoscyphaceae. The results showed that Sarco- somataceae was not monophyletic. Subsequently, Pfister et al. (2008) carried out a phylogenetic study inferred from nrLSU, SSU, and rpb2 sequence data, de- scribing the new family Chorioactidaceae Pfister, with four genera transferred: Chorioactis, Desmazierella, Neournula, and Wolfina, while Sarcosomataceae was restricted to a strongly supported group with six sexual genera: Donadinia, Galiella, Plectania, Pseudoplectania, Sarcosoma, and Urnula. The abbreviations for the generic names referenced in this study are as follows: D. = Donadinia, G. = Galiella, P = Plectania, Ps. = Pseudoplectania, U. = Urnula. Carbone et al. (2013a) clarified the taxonomic framework of Sarcosoma- taceae, which supported the monophyly of these six genera, as shown in the phylogeny based on a broader sequence dataset. Nonetheless, the mono- typic genus Korfiella D.C. Pant & V.P. Tewari, described from India (Pant and Tewari 1970), still lacks related molecular data. In addition, two asexual genera are currently accepted in Sarcosomataceae, viz. Conoplea Pers. and Strumella Fr., whose status in classification remains unclear and doubtful (Wijayawardene et al. 2017; Pfister and LoBuglio 2018). Previous phyloge- netic studies (Harrington et al. 1999; Carbone et al. 2013a; Zeng et al. 2021) seemed to reveal the dispersed phylogenetic position of these asexual taxa within clades of other sexual genera. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 22 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Currently, species of Sarcosomataceae are accepted to accommodate lignicolous or terricolous ascomycetous fungi, mainly macroscopically distin- guished by brown or black, discoid or cupulate to deep funnel-shaped, leath- ery or gelatinous, sessile or substipitate to stipitate apothecia (Carbone et al. 2013a; Zeng et al. 2021). In terms of microscopic morphology, Sarcosomata- ceae species present a definite arrangement of excipulum consisting of a ge- latinous medullary excipulum and an ectal excipulum with one or two types of external hairs; a hymenium consisting of two types of paraphyses; subopercu- late, unitunicate, inamyloid asci; and globose, ellipsoid, or allantoid ascospores with or without ornamentation and gelatinous sheath (Calonge et al. 2003; Carbone et al. 2014a, 2015a; Ekanayaka et al. 2018; Lu et al. 2023; Lin et al. 2024). Particularly, spore shapes and ornamentation, combined with molecular evidence, are recognized as the main characteristics for taxonomic treatment at the genus level (Carbone et al. 2013a), whereas a reliable criterion is lacking for distinguishing species (Korf 1972; Calonge and Mata 2002; Calonge et al. 2003; Carbone et al. 2013a; Wang et al. 2018; Lu et al. 2023). The genus Galiella was proposed by Nannfeldt and Korf (Korf 1957a) with the typification of Galiella rufa (Schwein.) Nannf. & Korf. In Korf’s generic concept, all bulgarioid or Sarcosomaz-like fleshy species with cyanophilous and warty ascospores were assigned to this genus. However, based sole- ly on morphological examination, the concept of Galiella has changed in- termittently. Galiella was regarded as a synonym of Sarcosoma by Boedijn (1959) and Le Gal (1958, 1960), while the independent status of Galiella was widely recognized by most authors (Korf 1972, 1973; Otani 1980; Moravec 1983; Cao et al. 1992; Zhuang and Wang 1998; Zhuang 2004; Carbone et al. 2015b; Popov and Carbone 2021). Furthermore, Samuelson et al. (1980) conducted an in-depth study on the apical characteristics of asci in Galiel- la and Sarcosoma, demonstrating that there were significant differences in ultrastructure and cell chemistry between them. By using the only G. rufa sequence data available for Galiella, Harrington et al. (1999) and Carbone et al. (2013a) initially confirmed the distinct position of this genus in Sar- cosomataceae. However, as the type species Trichaleurina polytricha Rehm was re-examined, two subtropical or tropical Galiella species were subse- quently transferred to the restored genus Trichaleurina, viz. Galiella celebica (Henn.) Nannf. and Galiella javanica (Rehm) Nannf. & Korf (Carbone et al. 2013b). At present, only Galiella amurensis (Lj.N. Vassiljeva) Raitv. and G. rufa have been confirmed through both morphological examination and mo- lecular data (KO6pcke et al. 2002; Carbone et al. 2013a; Popov and Carbone 2021). According to the recent concepts of Carbone et al. (2015b), the main distinguishing characteristics of Galiella species are as follows: gelatinous apothecia, warty or verrucose ascospores, and smooth external hairs. At present, two Galiella species (Galiella amurensis and Galiella sinensis J.Z. Cao) are known from China (Zhuang 2004; Popov and Carbone 2021). The taxonomic status of Galiella sinensis is noteworthy, given that this species was considered a synonym of Galiella celebica (Henn.) Nannf. (as Trichaleurina celebica (Henn.) M. Carbone, Agnello & P. Alvarado) by Zhuang and Wang (1998). However, Carbone et al. (2013b) pointed out that the type specimen (HMAS 29566) of G. sinensis was more closely related to a true Galiella species based on the smooth hairs they observed. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 23 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa The genus Plectania was established by Fuckel (1870) and was typified by Plectania melastoma (Sowerby) Fuckel (Sowerby 1799). Moreover, this genus was divided into five sections based only on spore shapes and ornamentation (Korf 1957a; Paden JW 1983; Carbone et al. 2012): Plectania sect. Plectania Fuckel, Plectania sect. Plicosporae Korf, Plectania sect. Curvatisporae Korf, Plectania sect. Sphaerosporae Paden (= Pseudoplectania), and Plectania sect. Donadinia (Bellem. & Mel.-Howell) M. Carbone & Agnello (= Donadinia). As shown by the phylogenetic analysis of Carbone et al. (2013a), the Plectania clade is monophyletic based on limited sequence data sets and molecular loci. However, the work of Zeng et al. (2021) and Hyde et al. (2024), which adopted more sequence data, showed that Plectania species did not cluster together and instead formed a paraphyletic group closely related to Galiella. In the current sense, Plectania is characterized by the following features: the ascocarp is discoid to deeply cupulate, sessile to stipitate, with a dark brown to nearly black external surface covered with dark hairs; the ec- tal excipulum is composed of angular cells, and the medullary excipulum consists of a textura intricata and is often gelatinous; the mature ascospores are elliptical, smooth or ornamented (transversally striated or verrucose), with or without a ge- latinous sheath (Carbone et al. 2013a, 2015a; Ekanayaka et al. 2018). Based on a study of Chinese specimens of Sarcosomataceae, Zhuang (2004) catalogued and described six species of Plectania in China: P campylo- spora (Berk.) Nannf. (a synonym of Urnula campylospora (Berk.) Cooke), P me- lastoma (Sowerby) Fuckel, P. nannfeldtii Korf (a synonym of Donadinia nigrella (Seaver) M. Carbone, Agnello & P. Alvarado), P. platensis (Speg.) Rifai, P. rhyt- idia (Berk.) Nannf. & Korf, and P. yunnanensis W.Y. Zhuang (Zhuang and Wang 1998; Zhuang 2004). In recent years, two novel species have been reported from subtropical China: Plectania lutea T. Bau & G.F. Mou (Mou and Bau 2021) and P. sichuanensis M. Zeng, Q. Zhao & K.D. Hyde (Zeng et al. 2021). The genus Pseudoplectania was initially erected by Fuckel in 1870. The typical representative of this genus, Ps. nigrella (Pers.) Fuckel, had already been docu- mented by Persoon (1801) and was formally designated as the type species of the genus by Korf (1972). For a long time, Pseudoplectania was considered an in- dependent genus by Seaver (1928, 1942), Nannfeldt (1949), Le Gal (1953), Sanwal (1953), Kreisel (1962), and Berthet (1964), standing out from other sarcosomata- ceous fungi by its spherical spores. Nevertheless, Korf (1982) and Paden (1983) argued that this genus should be treated as a synonym of Plectania, amending it due to its Conoplea conidial state and the rounded spores of Plectania when young. In 2013, the work of Carbone et al. (2013a) inferred that the genus Pseudo- plectania is a monophyletic clade based on phylogenetic analysis. To date, spe- cies of Pseudoplectania are distinguished by tiny to medium-large ascomata, sessile, shortly stipitate to distinctly stipitate, with a dark brown to black exter- nal surface often bearing basal tomentum; an ectal excipulum with a textura subglobulosa to prismatica angularis; one or two types of straight, curved, flexu- ous, or coiled external hairs; and, in particular, smooth, thick-walled, globose to subglobose mature spores with or without a gelatinous sheath, distinguishing them from other genera in Sarcosomataceae (Carbone et al. 2013a, 201 4a; Lin et al. 2024). Only four species of Pseudoplectania have been reported in China, namely Pseudoplectania globospora M. Zeng & Q. Zhao, Ps. mystica Jia Y. Lin, Ling Han Guo & Kun L. Yang, Ps. nigrella, and Ps. sinica Qiao Zhang & Jie Zhang (Zhuang 2018; Zhang and Zhang 2020; Lin et al. 2024; Zeng et al. 2024). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 24 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa The genus Urnula was established by Fries, with Urnula craterium (Schwein.) Fr. designated as the type species (Fries 1849, 1851). However, some Urnula species were previously placed in Plectania and Gloeocalyx Massee (a later synonym of Ur- nula) due to the significant morphological differences from the type species U. cra- terium, e.g., U. campylospora (Korf 1957b; Rifai 1968; Carbone and Agnello 2013). Subsequently, Carbone et al. (2013a) conducted an in-depth phylogenetic study, revealing a monophyletic group of Urnula species with strong support. Several spe- cies originally placed in Urnula, such as Urnula megalocrater Malengon & Le Gal, U. lusitanica Torrend & Boud., and U. platensis Speg., were transferred to other gen- era. Currently, this genus is recognized by its dark, bowl- to funnel-shaped ascoma- ta; stipitate or sessile fruitbodies; absence of crystals in the hymenium; one or two types of external hairs; gelatinous medullary excipulum consisting of a textura in- tricata; ectal excipulum with textura subglobulosa to angularis; and especially by its smooth, elliptical to allantoid mature ascospores, distinguishing it from other genera in Sarcosomataceae (Carbone et al. 2013a; Wang et al. 2018; Lu et al. 2023). These exceedingly similar morphological features provide little interspecific heterogene- ity, making Urnula species difficult to distinguish (Wang et al. 2018; Lu et al. 2023). There are relatively few previous studies or reports on the classification and diversity of Urnula species in China. Only five published species have been re- ported: Urnula ailaoshanensis J.R. Lu, Ying Zhang & Q. Zhao, U. campylospora (Berk.) Cooke, U. craterium (Schwein.) Fr., U. subcrateria J.R. Lu, F.M. Yu & Q. Zhao, and U. versiformis Y.Z. Wang & Cheng L. Huang (Nannfeldt 1949; Zhuang and Wang 1998; Zhuang 2004; Wang and Huang 2015; Lu et al. 2023). In this study, twelve specimens collected from subtropical China were newly examined and sequenced. Through field sampling, microscopic studies, ultra- structural observation, and phylogenetic analysis based on five molecular loci (ITS + nrLSU + rpb1 + rpb2 + tef1-a), four new species were confirmed: Plectania damingshanensis, P huapingensis, Pseudoplectania aureonigrescens, and Urnu- la auricularioides. Two new distribution records are also reported: P /utea and U. ailaoshanensis. Keys to the accepted species of Pseudoplectania, Urnula, and Chinese species of Plectania are also provided. Materials and methods Sampling, morphological observations, and descriptions Twelve specimens for this study were collected from Guangxi Zhuang Auton- omous Region (abbreviated as Guangxi below) and Fujian Province in China and deposited in the Herbarium of Guangxi Institute of Botany (IBK, Guangxi, China). Macro-morphological descriptions and measurements were based on fresh specimens. Micro-morphological data were obtained from dried speci- mens and observed under an optical microscope and scanning electron micro- scope. Color codes followed the standards established by Kornerup and Wan- scher (1978). Microscopic features were studied on air-dried samples after hand sectioning and rehydrating in distilled water and, if necessary, mounting in Congo red under suitable light conditions. The iodine reaction of asci and ascospores was tested using Melzer’s reagent. For optimal microscopic ob- servation, this study utilized a Nikon E80i microscope (Japan) to examine the sections at magnifications of x100, x200, x400, x600, and x1000. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 25 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa For electron microscopy, the ultrastructure of ascospores was observed us- ing a Zeiss EVO18 scanning electron microscope (Carl Zeiss IMT (Shanghai) Co., Ltd., Shanghai, China). The ascospore dimensions are described as fol- lows, excluding ornamentation: [a/b/c] length x width, and the factor Q, which is the ratio of spore length to width. The measured value from a ascospores of b ascomata in c specimens is shown as (d) e-f (g), where e-f represents at least 90% of the values, d is the minimum, and g is the maximum value. Thirty or more mature and rehydrated ascospores were measured. In describing the morpho- logical characteristics, terminology follows Carbone et al. (2013a, 2014a), Zeng et al. (2021), Lu et al. (2023), Lin et al. (2024), and Zeng et al. (2024). DNA extraction and sequence amplification DNA was extracted from dried specimens using the NuClean Plant Genomic DNA Kit (CWBIO). The primer pairs used included ITS4 and ITS5 for the internal transcribed spacer region (ITS) (White et al. 1990); LROR and LR5 for the large subunit nuclear ribosomal RNA (nrLSU) (Vilgalys and Hester 1990); RPB1-Af and RPB1-Cf for the first largest subunit of RNA polymerase | gene (rpb7) (Matheny et al. 2002); fRPB2-5F and fRPB2-7.1R for the second largest subunit of RNA polymerase II gene (rpb2) (Matheny 2005); and EF1-983F and EF1-1567R for the translation elongation factor 1-a gene (tef7-a) (Rehner and Buckley 2005). Polymerase chain reaction (PCR) was performed in a total volume of 30 UL, consisting of 15 pL of 2x Es Taq MasterMix, 9 uL of ddH,O, 1.5 uL of each forward and reverse primer, and 3 uL of DNA template. The PCR pro- cedure for ITS and nrLSU was as follows: initial denaturation at 94 °C for 4 min; followed by 35 cycles at 94 °C for 40 s, 52 °C for 40 s, and 72 °C for 1 min; with a final extension at 72 °C for 10 min. The PCR procedure for rpb7, rpb2, and tef1-a was as follows: initial denaturation at 94 °C for 2 min, fol- lowed by denaturation at 94 °C for 40 s, annealing at 60 °C for 40 s with an increment of 1 °C per cycle, and extension at 72 °C for 2 min. This step was repeated 8 times. Then, denaturation at 94 °C for 45 s, annealing at 53 °C for 1.5 min, and extension at 72 °C for 2 min, repeated 36 times. Finally, a final extension was performed at 72 °C for 10 min. Successful PCR products were sent to Sangon Biotech (Shanghai) Co., Ltd., for sequencing. Sequence results were checked using Chromas v2.6.6 (https://technelysium.com.au/ wp/chromas), and sequence data from the specimens were submitted to GenBank (https://www.ncbi.nIm.nih.gov/genbank). Data analysis Sequences of Sarcosomataceae downloaded from GenBank mainly referred to Carbone et al. (2013c, 2014a), Zeng et al. (2021), Lu et al. (2023), and Lin et al. (2024). Additional unpublished sequences were also downloaded from Gen- Bank (https://www.ncbi.nim.nih.gov/genbank). Detailed information on the se- quences included is shown in Table 1. A total of 128 ITS, 111 nrLSU, 11 rpb7, 17 rpb2, and 25 tef1-a sequences from Sarcosomataceae were used for molecular phylogenetic analysis. Sequences were aligned using Clustal W in BioEdit v7.1.3 with default parameters (Hall 1999). This study selected five species from two families, Chorioactidaceae Pfister and Pyronemataceae Corda, as outgroups MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 6 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa of Sarcosomataceae (Perry et al. 2007; Pfister et al. 2008). Alignments were refined in MEGA 11 (Tamura et al. 2021). Individual alignments were concate- nated for ITS + nrLSU + rpb1 + rpb2 + tef1-a using SequenceMatrix (Vaidya et al. 2011). The concatenated dataset was submitted to TreeBASE (ID: 32052). The optimal evolutionary model for the matrix was selected using MrMod- eltest v2.3 based on the Akaike Information Criterion (AIC) (Nylander et al. 2004). Maximum likelihood (ML) and Bayesian inference (Bl) were both used for phylogenetic reconstruction. ML analysis was performed in RAxMLGUI v1.5b1 (Stamatakis et al. 2005) using the bootstrap method with 1,000 repli- cates and the rapid bootstrapping algorithm. BI analysis was conducted using MrBayes v3.2.7 (Ronquist and Huelsenbeck 2003) with Markov chain Monte Carlo (MCMC) simulation set at 5,000,000 generations, sampling trees and pa- rameters every 1,000" generation (two parallel runs; burn-in = 25%). Bayesian posterior probability (BPP) was used to assess branch support. Significance thresholds were defined as follows: MLB = 50% and BPP = 0.90 for well-sup- ported branches; either MLB < 50% or BPP < 0.90 for moderate support; and MLB < 50% and BPP < 0.90 for unsupported branches. FigTree v1.4.4 was used to visualize the phylogram (Rambaut 2018). Results Phylogenetic analyses A total of 3,599 aligned nucleotide sites were used for the phylogenetic recon- struction, including positions 1-501 bp for ITS, 502-1382 bp for nrLSU, 1383- 2169 bp for rpb71, 2170-3055 bp for rpb2, and 3056-3599 bp for tef7-a. The best-fit substitution models selected for reconstructing the phylograms were GTR+I+G for ITS, nrLSU, and tef7-a; GTR+G for rpb1; and SYM+I+G for rpb2. The output topology of the Bayesian inference (BI) tree was nearly identical to that of the maximum likelihood (ML) analysis, so only the ML tree is shown in Fig. 1. In this multi-locus phylogenetic analysis, the family Sarcosomataceae formed a monophyletic group with strong support (MLB = 100%, BPP = 1). The tree also demonstrated that Donadinia, Pseudoplectania, Sarcosoma, and Urnu- la are monophyletic, while Galiella and Plectania are polyphyletic. Based on the overall topology, five single, well-supported clades were clearly inferred with- in Sarcosomataceae. These five clades, comprising 137 taxa representing 45 species, were as follows: the Donadinia clade (MLB = 99%, BPP = 1), Plectania s. lat. clade (MLB = 78%, BPP = 1), Pseudoplectania clade (MLB = 69%, BPP = 0.93), Sarcosoma clade (MLB = 100%, BPP = 1), and Urnula clade (MLB = 95%, BPP = 1). Notably, the Pseudoplectania clade was significantly supported as a sister group to the Sarcosoma clade (MLB = 88%, BPP = 1). The Urnula and Donadinia clades formed a monophyletic group (MLB = 92%, BPP = 1), which was sister to the Plectania s. lat. clade (MLB = 87%, BPP = 1). Within the Plectania s. lat. clade, the two Galiella lineages—Galiella amuren- sis and G. rufa—clustered with Plectania species. Plectania damingshanensis formed a strongly supported lineage (MLB = 100%, BPP = 1) with four speci- mens and was recovered as sister to P /utea. Meanwhile, HP2024071903 and HP2024072014 formed a relatively independent branch with strong support (MLB = 100%, BPP = 1) and are regarded here as P. huapingensis. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 7 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Table 1. Taxa, sequences, and collections analyzed in this study. Collections in bold are newly sequenced in this study. The letters HT, ET, IT, and NT after the sample ID/voucher indicate holotype, epitype, isotype, and neotype, respectively. NA indicates sequences that are not available. GenBank accession Specie Sample ID/Voucher ae aT 7B be ere Reference Chorioactis geaster DHP 02.497 NA KC012672 NA NA KC109211 Hansen et al. (2013) Conoplea fusca CBS: 113475 EU552114 NA NA NA NA Carbone et al. (2013a) C. fusca CHTAR77 NA GU048612 NA NA NA Carbone et al. (2013a) C. globosa CBS:233.91 AF485076 NA NA NA NA K6pcke et al. (2002) C. globosa CBS:438.51 AF485069 NA NA NA NA Képcke et al. (2002) Conoplea sp. JCM12000 JQ972061 NA NA NA NA NCBI Donadinia echinacea HKAS 107659" MW077543 | MW079923 | MW084979 NA MW094269 Zeng et al. (2021) D. echinacea HKAS 107660 MW077544 | MW079924 NA MW085092 NA Zeng et al. (2021) D. echinacea HKAS 107661 MW077545 | MW079925 NA NA NA Zeng et al. (2021) D. echinacea HKAS 107662 MW077546 | MW079926 NA NA MW094270 Zeng et al. (2021) D. helvelloides LY PB 940" JX669834 | JX669872 NA NA NA Carbone et al. (2013a) D. helvelloides SMPM206 KP794618 | KP794617 NA NA NA NCBI D. helvelloides MCVE 28377 KP204907 | KP204914 NA NA NA Carbone et al. (2014b) D. lusitanica MCVE 28378 KP204906 | KP204913 NA NA NA Carbone et al. (2014b) D. lusitanica TUR-A 195791 JX669811 | JX669847 NA NA NA Carbone et al. (2013a) D. lusitanica TUR-A 195792 JX669810 | JX669846 NA NA NA Carbone et al. (2013a) D. nigrella TUR-A 195793 JX669836 | JX669874 NA NA NA Carbone et al. (2013a) D. nigrella WTU-F-017148 KP204912 | KP204919 NA NA NA Carbone et al. (2014b) D. nigrella WTU-F-017150 KP204911 | KP204918 NA NA NA Carbone et al. (2014b) D. nigrella CBS:694.71 MH860304 NA NA NA NA NCBI D. nigrella KH-97-16 (FH) NA AY945853 | JX943641 | DQ017592 | KC109214 Hansen et al. (2013) D. seaveri CUP-Whetz.B.F. 01884" | KC249999 NA NA NA NA Carbone et al. (2014b) D. seaveri FH 00458441 KY794720 | KY794715 NA NA NA Harrington et al. (1999) D. seaveri FH 01142449" KY794717 | KY794712 NA NA NA Carbone et al. (2013a) D. sibirica MCVE 28374'7 KP204910 | KP204917 NA NA NA Carbone et al. (2014b) D. sibirica MCVE 28375 KP204908 | KP204915 NA NA NA Carbone et al. (2014b) Donadinia sp. mh 699 (FH) NA DQ220329 NA DQ017593 NA Perry et al. (2007) Galiella amurensis LE 236216 MW879699 | MW879115 NA NA NA Popov and Carbone (2021) G. amurensis LE 323821 MW879700 | MW879116 NA NA NA Popov and Carbone (2021) G. amurensis KE323822"" MW879701 | MW879117 NA NA NA Popov and Carbone (2021) G. rufa CBS:135.92 AF485070 | FJ176869 NA NA NA K6pcke et al. (2002) G. rufa CBS:762.85 AF485072 | KC012674 NA NA NA Pfister et al. (2018) G. rufa DHP 05.600 (FH) NA KC012674 | JX943642 NA KC109213 Hansen et al. (2013) G. rufa AFTOL-ID 1297 NA FJ176869 NA FJ238352 FJ238401 Schoch et al. (2009) Neournula pouchetii TUR-A 195798 JX669837 | JX669875 NA NA NA Carbone et al. (2013a) N. pouchetii MO-205345 KT968605 | KT968655 NA NA NA NCBI Plectania damingshanensis MES2024040606 PQ691392 | PQ682446 | PV247119 NA PV295999 This study P. damingshanensis DMS2024022001"" PP864720 | PP859016 | PV247118 NA NA This study P. damingshanensis DMS2024022002 PP864721 | PP859017 NA NA PV295998 This study P. damingshanensis DMS2024022003 PP864722 | PP859018 NA NA NA This study P fisherae MST-F22868 PQ066526 | PQ060437 NA PQ067358 NA Yu et al. (2024) P. harnischii TUR-A 195785 NR_174881 | NG_079625 NA NA NA Carbone et al. (2021) P. harnischii WTU-F-68864 MZ713186 | MZ713201 NA NA NA Carbone et al. (2021) P. huapingensis HP2024071903'" PQ691391 | PQ682445 NA PV296007 | PV296000 This study P. huapingensis HP2024072014 PQ691390 | PQ682444 NA NA NA This study P lutea HMJAU 57091" NR_189838 | NG_242497 NA NA NA Mou and Bau (2021) P lutea IBKMO1 MW794211 | MW794213 NA NA NA Mou and Bau (2021) MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 28 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa GenBank accession Specie Sample ID/Voucher P. lutea MES2024033005 | PP859394 | PP981371 | PV247120 | PV296006 | PV296001 P. megalocrater TUR-A 195803 JX669809 | JX669845 Zz > Zz > NA N Ps. sinica CGMCC 3.19892 NA Ps. sinica CGMCC 3.19892 NA Pseudoplectania sp. 3-1-7-2-4-1 NA Pseudoplectania sp. DO87 NA Pseudoplectania sp. ER1858 MZ091916 | MZ019023 Pseudoplectania sp. QU0896 MZ091877 | MZ019019 a > a A Zz > Zz > Zz > N NA NA > Zz A Zz > NA N > Zz > P. cf. melastoma TUR-A 195785 Jx669804 | JX669840 P. melastoma TUR-A 195783 JX669805 | JX669841 P melastoma TUR-A 195784 JX669814 | JX669850 P. melastoma MH679 MN103434 | MN103422 P. milleri OSC 104436 FU652354 | EU652389 P milleri OSC 73165 FU652355 | EU652390 P milleri TUR-A 190823 JX669812 | JX669848 P. rhytidia PDD 90028 JX669832 | JX669871 P. rhytidia TUR-A 195786 JX669813 | JX669849 P. rhytidia TUR-A 195787 JX669815 | JX669851 P. rhytidia TUR-A 195788 JX669816 | JX669852 NA P. sichuanensis HKAS 107664"" | MW077547 | MW079927 Mwo08s5093 | Mwo094271 P. submilleri HKAS 129665" | OR500919 | OR500933 P. submilleri HKAS 129666 0R500920 | OR500934 Plectania sp. Mumblo 2007090-01 | JX310423 | JX287505 Plectania sp. PA39 MG543948 NA P. zugazae AVM1467#" JX669817 | JX669854 P zugazae AVM2086 JX669818 | JX669855 P. zugazae TUR-A 199785 KM610322 | KM610324 Pseudoplectania affinis PDD 818424" JX669826 | JX669865 Ps. affinis RBG 7287 ON989641 OP066979 | OP066929 Ps. africana TUR-A 216892 PP590544 | PP587623 Ps. africana ZE59/18"" MT496892 | MT496884 OR961521 | OR961523 Ps. africana TUR-A 216593 OR940185 | 0R939326 OR961522 | OR961524 Ps. aureonigrescens CSA-797 PQ863322 PQ863491, NA | NA | PV296004 Ps. aureonigrescens CSA-798"" PQ895821 | PQ863683 | PV247121| NA | PV296005 Ps. episphagnum TUR 064173 KF305711 | KF305724 Ps. ericae MCVE 27581 KF305721 | KF305731 Ps. ericae TUR-A 195789 JX669822 | JX669862 Ps. ericae MPU: JCD 82-775 | MT273641 MT274711 Ps. globospora HKAS 127988" | PP357153 | OR879981 Ps. globospora HKAS 127989 PP357154 | OR879982 Ps. lignicola HR89756 MT496886 | MT496882 Ps. lignicola SAV 105/17 MT496881 | MT496883 Ps. melaena MCVE 27433 JX669806 | JX669842 Ps. melaena MCVE 27579 KF305717 | KF305728 Ps. melaena NY 54130 KF305718 | KF305730 Ps. mystica HKAS133073"" | PP422101 | PP422115 Ps. mystica HKAS133074 PP422102 | PP422114 Ps. nigrella KL BK-4914"" JX669807 | JX669843 Ps. nigrella MCVE 27396 KF305715 | KF305725 Ps. nigrella TUR 169888 JX669821 | JX669859 Ps. nigrella FH 00822741 MN103433 | MN103421 —s17241 | wa0es200 | NA | nA | NA na 057 wrososiz wa) SNA | SONA ON MycoKeys 121: 21-65 (2025 a , DOI: 10.3897/mycokeys.121.155432 Reference This study Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2013a) Pfister et al. (2019) NCBI NCBI Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2013a) Zeng et al. (2021) Hyde et al. (2024) Hyde et al. (2024) NCBI NCBI Pfister et al. (2019) Pfister et al. (2019) Carbone et al. (2015) Carbone et al. (2014a) NCBI Pfister et al. (2019) Pfister et al. (2019) Pfister et al. (2019) This study This study Carbone et al. (2013a) Carbone et al. (2013a) Carbone et al. (2014a) Carbone et al. (2013a) Pfister et al. (2019) Zeng et al. (2024) Zeng et al. (2024) Pfister et al. (2019) Pfister et al. (2019) Carbone et al. (2015) Carbone et al. (2015) Carbone et al. (2015) Lin et al. (2024) Lin et al. (2024) Carbone et al. (2013a) Carbone et al. (2015) Carbone et al. (2013a) Pfister et al. (2019) NCBI Zhang and Zhang (2020) Li et al. (2016) NCBI Healy et al. (2022) Healy et al. (2022) 29 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa GenBank accession Specie Sample ID/Voucher Reference ITS nrLSU rpb1 rpb2 tef1-a Ps. tasmanica MCVE 27584 KF305723 | KF305733 NA NA NA Carbone et al. (201 4a) Ps. tasmanica MCVE 275831" KF305722 | KF305732 NA NA NA Carbone et al. (201 4a) Pseudosarcosoma TUR-A 195801 JX669819 | JX669856 NA NA NA Carbone et al. (2013a) latahensis Sarcosoma globosum KH.07.04 (S) FJ499393 NA JX943640 | JX943753 | KC109215 NCBI S. globosum LE-BIN 3794 KY344789 NA NA NA NA NCBI S. globosum AJ1324 OR529395 NA NA NA OR543971 NCBI Trichaleurina tenuispora TUR-A 195800 JX669839 | JX669876 NA NA NA Carbone et al. (2013a) T. javanica TUR-A 195799 JX669838 | JX669861 NA NA NA Carbone et al. (2013a) T. javanica HKAS 88981 MG871291 | MG871326 NA MG980716 | MG980693 NCBI Urnula ailaoshanensis HKAS129212"" 0Q941816 | 0Q941895 NA NA NA Lu et al. (2023) U. ailaoshanensis HKAS129213 0Q941817 | 0Q941896 NA NA NA Lu et al. (2023) U. ailaoshanensis MES2023081104 PQ187434 | PQ187435 | PV247122 NA NA This study U. auricularioides CSA-455" PQ489315 | PQ187431 | PV247123 | PV296008 | PV296003 This study U. auricularioides CSA-454 PQ489472 | PQ187433 NA NA PV296002 This study U. campylospora PDD 83522 JX669830 | JX669869 NA NA NA Carbone and Agnello (2013) U. campylospora PDD 88805 JX669831 | JX669870 NA NA NA Carbone and Agnello (2013) U. craterium 30.15.291.11 KF311230 | KF305734 NA NA NA Carbone et al. (2014a) U. craterium TUR-A 195794 JX669820 | JX669857 NA NA NA Carbone et al. (2013a) U. craterium DHP 04-511 (FH) NA AY945851 | JX943639 | DQ0O17595 | KC109216 Hansen et al. (2013) U. hiemalis TUR 136909 JX669827 | JX669867 NA NA NA Carbone et al. (2013a) U. hiemalis TUR 196076 JX669828 | JX669868 NA NA NA Carbone et al. (2013a) U. hiemalis TUR-A 195795 JX669835 | JX669873 NA NA NA Carbone et al. (2013a) U. himalayana CAL 16734" NR_159073 | NG_064532 NA NA NA Wang et al. (2018) U. himalayana CAL 1674 MH179123 | MH179124 NA NA NA Wang et al. (2018) U. mediterranea MCVE 28636 KU933925 | KU933926 NA NA NA Marziani et al. (2016) U. mediterranea TUR-A 195796 JX669808 | JX669844 NA NA NA Carbone et al. (2013a) U. mediterranea TUR-A 195797 JX669824 | JX669864 NA NA NA Carbone et al. (2013a) U. padeniana WTU-F-3305147 JX669825 | JX669866 NA NA NA Carbone et al. (2013a) U. subcrateria HKAS129214"7 0Q944116 | 0Q941897 NA NA NA Lu et al. (2023) U. subcrateria HKAS129215 0Q944117 | 0Q941898 NA NA NA Lu et al. (2023) Urnula sp. YC2023090928 PQ550998 | PQ187037 NA NA NA This study Urnula sp. EN-22 KT358892 NA NA NA NA Healy et al. (2022) Urnula sp. GL1-D-D1 KX100405 NA NA NA NA Strobel et al. (2017) Urnula sp. PDD 81259 JX669829 NA NA NA NA Carbone and Agnello (2013) Urnula sp. TNM F15544 KJ577537 NA NA NA NA Wang and Huang (2015) U. versiformis TNM F11312 KJ577534 NA NA NA NA Wang and Huang (2015) U. versiformis TNM F11317"7 KJ577535 NA NA NA NA Wang and Huang (2015) U. versiformis TNM F13875 KJ577536 NA NA NA NA Wang and Huang (2015) In Pseudoplectania, based on distinguishing morphological characters (see details in “Discussion’), high statistical support, and clear genetic relationships, a further division is proposed here: subclade | (MLB = 79%, BPP = 0.78), sub- clade II (MLB = 100%, BPP = 1), subclade III (MLB = 100%, BPP = 1), and sub- clade IV (MLB = 84%, BPP = 0.7). A lineage labeled Pseudoplectania aureoni- grescens was strongly supported (MLB = 100%, BPP = 1) with two specimens and was affiliated with Ps. affinis and Ps. globospora within subclade III. In Urnula, subclade A was significantly supported (MLB = 100%, BPP = 1). Within subclade A, the lineage representing U. auricularioides (MLB = 100%, BPP = 1) grouped together with U. himalayana. The topology also showed that MES2024033005 belongs to P /utea, while MES2023081104 belongs to U. ailaoshanensis, and YC2023090928 was recovered in the genus Urnula. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 30 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa ioonf Ponadinia echinacea HKAS 107661 China @ VE Donadinia echinacea HKAS 107662 China Donadinia echinacea HKAS 107660 China > VES Donadinia echinacea HKAS 107659 HT China Donadinia seaveri CUP-Whetz.B.F.0188 HT UK © SA ory _100/1 Donadinia seaveri FH 01142449 IT UK ‘@) Donadinia sp. mh 699 (FH) Unknown (\GE 87/0.58' Donadinia seaveri FH 00458441 USA O S 100/1] Ponadinia /usitanica TUR-A 195792 Greece © co Gas 51/- Donadinia lusitanica MCVE 28378 Greece © ee : ww se 2070.99 1900/1”! Donadinia lusitanica TUR-A 195791 Italy © Donadiniai Donadinia helvelloides SMPM206 France oO : SES i007) Donadinia helvelloides MCVE 28377 France © \ 100/1) 90/0.93L Donadinia helvelloides LY PB 940 IT France © ©) se 10017 Donadinia sibirica MCVE 28375 Russia lan 100/1 Donadinia sibirica MCVE 28374 HT Russia © ) s6c oo Donadinia nigrella TUR-A 195793 HT Russia 4 if Donadinia nigrella CBS:694.71 Canada - SGE on 82/0.93| , Donadinia nigrella KH-97-16 (FH) Unknown 95/1| Donadinia nigrella WTU-F-017148 USA © 100/-"! Donadinia nigrella WTU-F-017150 USA © Conoplea globosa CBS:438.51 Unknown 7/0.87) sao.e7) Urnula hiemalis TUR 136909 Finland © 100/1* Urnula sp. GL1-D-D1 Greenland Urnula hiemalis TUR-A 195795 Finland 66/0782 Urnula hiemalis TUR 196076 Finland © Conoplea globosa CBS:233.91 Unknown 10/1 Urnula craterium 30.15.291.11 France i Urnula craterium TUR-A 195794 Italy © 58/0.68 Urnula craterium DHP 04-511 (FH) Unknown 54/0.59|) Urnula subcrateria HKAS 129214 HT China © sri] 1°9/9-95" Urnula subcrateria HKAS 129215 China @ Urnula padeniana WTU-F-33051 HT USA © Urnula sp. YC2023090928 China toon? Urnula mediterranea TUR-A 195796 Italy © ee tien rosa Hoops aie 195797 France © 86/0.56 ‘onoplea sp. rance 96/0.99! Urnula mediterranea MCVE 28636 Italy (@) Ur. nu, I. a 87/1 toon Uraula sp. EN22 Australia 1 Urnula campylospora PDD 88805 New Zealand © nti 100/18 Urnula campylospora PDD 83522 New Zealand © at Urnula sp. PDD 81259 New Zealand Urnulasp. TNM F15544 China Urnula ailaoshanensis MES2023081104 China © oo7ih Urnula ailaoshanensis HKAS 129212 HT China © 52/0.93 98/1 100/0.83-7] Urnula ailaoshanensis HKAS 129213 China () A 56/0.86] 199/47 Urnula versiformis TNM F11312 China @ qoore Urnula versiformis TNMF11317 HT China © N| | Urnula versiformis TNM F13875 China re) 100/iPay Urnula auricularioides CSA-455 HT China © Urnula auricularioides CSA-454 China (e) 69/0.561 ) Urnula himalayana CAL 1673 HT India ie) 100/1 Urnula himalayana CAL 1674 India ie) 59/--— Plectania fisherae MST-F22868 HT Australia 1001] ———_ Conoplea fusca CBS:113475 South Africa Conoplea fusca CHTAR77 Mexico avoes|( Plectania rhytidia PDD 90028 New Zealand © 69/0.86] [ Plectania rhytidia TUR-A 195788 Italy (2) 34/0.94| Plectania rhytidia TUR-A 195786 Italy 3] 70/-¥L Plectania rhytidia TUR-A 195787 Italy () 78/1 - Plectania melastoma MH679 Unknown yo0/1) Plectania huapingensis HP2024072014 China oO 59/0.55 [ Plectania er te dpa a ee 903 HTChina © . Galiella amurensis LE323822 ET Russia ; Galiella amurensis LE323821 Russia r4 Galiella amurensis 51/0.99 Galiella amurensis LE236216 Russia ie) 100/0.96 Plectania sichuanensis HKAS 107664 HT China © 98/4) Galiella rufa DHP 05.600 (FH) Unknown 100/1| ' Galiella rufa CBS:762.85 Unknown © : 001 Galiella rufa CBS:135.92 Unknown © Galiella rufa 99/1! Galiella rufa AFTOL-ID 1297 Unknown 76/0.91 100/15 Plectania submilleri HKAS 129666 China N Plectania submilleri HKAS 129665 HT China e Ma Plectania milleri OSC 104436 Unknown Plectanias. \at. Plectania milleri OSC 73165 Unknown Plectania milleri TUR-A 190823 USA © 80/1 Plectania sp. Mumblo2007090-01 Unknown Plectania megalocrater TUR-A 195803 Greece © roof Plectania lutea MES2024033005 China T} Plectania lutea HMJAU 57091 HT China © P67"! Plectania lutea IBKM01 China @ 100/1 Plectania damingshanensis DMS2024022003 China 10071) Plectania damingshanensis DMS2024022002 China 1-4" Plectania damingshanensis DMS2024022001 HT China 100/0,96 «4 2°!" | Plectania damingshanensis MES2024040606 China : Plectania harnischii WTU-F-68864 USA oO 100/0.97 Plectania harnischii TUR-A 195785 HTUSA =O 93/0974 |" Plectania cf. melastoma TUR-A 195785 USA sooty Plectania zugazae TUR-A 199785 Cyprus O 67/- in! Plectania zugazae AVM2086 Spain 81/0. ju Plectania zugazae AVM1467 HT Spain (3 Plectania sp. PA39 France 100/0.94) Plectania melastoma TUR-A 195784 Italy © 100/- Plectania melastoma TUR-A 195783 Italy © @000 Figure 1. Phylogenetic tree of Sarcosomataceae inferred from the dataset with the aligned ITS + nrLSU + rpb7 + rpb2 + tefl-a contig. Nodes are separated by ‘-/-’ (MLB/BPP) and indicated if bootstrap support values for ML = 50% and/or posterior probability for BI = 0.90. Newly collected specimens of the current study are highlighted in red. The letters HT, ET, IT, and NT, and the country highlighted in blue after the specimens stand for holotype, epitype, isotype, neotype, and the original country, respectively. Icons depicting the shape and ornamentation of spores and the presence and position of a gelatinous sheath are indicated after the specimens and correspond to abbreviations. VE indicates verrucose or vermicular ellipsoid spores without a gelatinous sheath. VES indicates verrucose or vermicular ellipsoid spores with a gelatinous sheath. SA indicates a smooth allantoid spore without a gelatinous sheath. GE indicates grooved ellipsoid spores without a gelatinous sheath. SE indicates smooth ellipsoid spores without a gelatinous sheath. SES indicates smooth ellipsoid spores with a gelatinous sheath. SG indicates smooth, globose to subglobose spores without a gelat- inous sheath. SGC indicates smooth globose to subglobose spores with a centrally arranged gelatinous sheath. SGE indicates smooth globose to subglobose spores with an eccentrically arranged gelatinous sheath. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 3] Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa 0.1 100; Sarcosoma globosum AJ1324 Canada e) Sarcosoma globosum KH.07.04 (S) Sweden © [ Sarcosoma 97/0.841 Sarcosoma globosum LE-BIN 3794 Russia ©) 100/1) Pseudoplectania sinica CGMCC 3.19892 China Pseudoplectania sinica CGMCC 3.19892 China Pseudoplectania melaena NY 54130 USA CO I Pseudoplectania melaena MCVE 27579 France © 98/" Pseudoplectania melaena MCVE 27433 Italy e 1001 PSeudoplectania sp. QU0896 Canada Pseudoplectania episphagnum TUR 064173 Finland © 1) 28/9.984 Pseudoplectania episphagnum TUR 064171 Finland © 100/4 Pseudoplectania lignicola HR89756 Czech @) Pseudoplectania lignicola SAV 105/17 Slovakia fe) II Pseudoplectania nigrella TUR 169888 Finland 2) Oo Pseudoplectania nigrella MCVE 27396 Italy © Pseudoplectania affinis RBG7287 Australia o) Pseudoplectania affinis PDD 81842 HT New Zealand © 100/15 Pseudoplectania aureonigrescens CSA-797 China (e) Pseudoplectania aureonigrescens CSA-798 HT China @ Ul Pseudoplectania globospora HKAS 127989 China oO 100/" Pseudoplectania globospora HKAS 127988 HT China © Pseudoplectania nigrella FH00822741 Unknown 100/14" Pseudoplectania tasmanica MCVE 27584 Australia 2) Pseudoplectania tasmanica MCVE 27583 HT Australia © 100/1] 100/1] Pseudoplectania ericae TUR-A 195789 Spain 2) Pseudoplectania ericae MPU:JCD 82-775 France © Pseudoplectania ericae MCVE 27581 Italy oO Pseudoplectania africana TUR-A 216892 Portugal © IV Pseudoplectania africana TUR-A 216593 Portugal @ Pseudoplectania africana ZE59/18 HT South Africa @ Pseudoplectania sp. 3-1-7-2-4-1 China 1007p Pseudoplectania mystica HKAS 133074 China o Pseudoplectania mystica HKAS 133073 HT China @ 100/" Pseudoplectania sp. DO87 China Chorioactis geaster DHP 02.497 (FH) USA Trichaleurina tenuispora TUR-A 195800 Japan 100/1-— Trichaleurina javanica HKAS 88981 Unknown Trichaleurina javanica TUR-A 195799 Seychelles Pseudosarcosoma latahense TUR-A 195801 Canada O utg rou ps 100/1 Neournula pouchetii MO-205345 Unknown Neournula pouchetii TUR-A 195798 Italy Pseudoplectania O 84/0.7] Figure 1. Continued. Taxonomy Plectania damingshanensis G. F. Mou & J. R. Liu, sp. nov. Fungal Names: FN 572018 Figs 2-4 Etymology. The specific epithet refers to the type locality, Damingshan National Nature Reserve, in China. Diagnosis. It is characterized by the cupulate ascomata, the surface of hyme- nium with brownish orange to light brown color, external surface and stipe (when present) densely covered with grey tomentum and tree-like ridges, the elliptical as- cospore is warty under SEM, (21.7) 22.5-32.0 (34.6) x (11.0) 12.1-14.7 (16.5) um. Holotype. CHINA * Guangxi Zhuang Autonomous Region, Nanning City, Damingshan National Nature Reserve, 23°49'N, 108°43'E, ca 1230 m alt., on rotten fallen sticks or wood surrounded by moss in the evergreen broad-leaved forest, 19 Feb. 2024, Guang R. Zhou (DMS2024022001, IBK, holotype!) (ITS: PP864720; nrLSU: PP859016; rpb1: PV247118). Description. Apothecium cupulate, up to 19-27 mm in diameter, 20-30 mm high, subsessile to stipitate (Fig. 2a, b). Hymenium surface glabrous, without ornamentation or wrinkled, brownish orange (6C6) to light brown (6D7), brown- dark, brown (6E8-6F8) after drying, margin with light brown (6D5) tomentum. External surface reddish brown-dark brown (7D8-—7F8), brownish grey (8F2) after drying. Stipe (when present) up to 2-20 mm long, 7.5-9 mm in width, densely covered with grey (8D1—8F1) tomentum and tree-like ridges extending all the way to the margin of the cup (Fig. 2b, d). Flesh solid, wet, gelatinous at a high level, semitransparent, slight grey (8B1). Odor and taste not special. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 32 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Figure 2. Ascocarps of Plectania damingshanensis. Photos by Guang R. Zhou and G. F. Mou. Scale bars: 10 mm (a-d). External hairs curved and flexuous, septate, glabrous, brown (6E7) with an olive tint (1E8), 5-8.5 um in diameter, with obtuse end (Fig. 3g, h). Ectal excipulum of textura angularis made up of thick-walled cells, brownish to dark brown, 13-91 um thick (Fig. 3g), near the margin arranged in separable moniliform hair-like cells, 6.2-10.6 um diam (Figs 3f, 4g). Medullary excipulum of loose textura intricata immersed in a highly gelatinous matrix, approximately 380-1000 um, subhya- line to yellowish white (2A2), composed of 2—4.2 um broad hyphae (Figs 3c, 4f). Subhymenium of a dense textura intricata of closely septate hyphae, olive brown (4D7-4D6), 50-80 um thick. Asci cylindrical, suboperculate, 8-spored, rarely with 7 spores, thick-walled, inamyloid, 322-401 (426) x 11.0-15.6 (Fig. 3j-m, 4a). Para- physes 1.5-3 um diam, filiform, septate, branched, slightly enlarged or narrowed near apex (Figs 3d, 4d). Ascospores uniseriate, ellipsoid, equilateral, hyaline, very finely warty under light microscope (x1000, Fig. 3i), verrucose to vermicular under scanning electron microscope (Fig. 16b), with 1-2 guttules, [40/5/2] (21.7) 22.5— 32.0 (34.6) x (11.0) 12.1-14.7 (16.5) um, Q = (1.7) 1.8-2.3 (2.6), Q., = 2.0. Hymenial hairs cylindrical, comparable in length to the paraphyses, non-septate, 2.1—4.4 um in diameter, concolor with paraphyses due to the homogeneous pigments, aggluti- nating or intermingle with paraphyses to bundles, tips rounded to slightly subcapi- tate, straight or curved, unbranched (Figs 3e, 4c). Anamorph unknown. Habitat. Growing in scattered to gregarious groups on moss-covered decaying wood in broadleaf forests. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 33 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa i Figure 3. Features of Plectania damingshanensis. (photos by J. R. Liu, from holotype: DMS2024022001, IBK!). a. Section of apothecium; b. Hymenium; c. Medullary excipulum; d. Paraphyses and a hymenial hair; e. Hymenial hair; f. Moniliform hair-like cells; g, h. External hairs; i. Ascospores; j—m, 0. Asci; n. Apex of asci. Scale bars: 100 um (a); 20 um (b, j-m); 10 um (c-i, n-o). c, e, min 1% Congo Red solution. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 34 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa a Figure 4. Microscopic structures of Plectania damingshanensis (drawings by J. R. Liu, from holotype DMS2024022001, IBK!). a. b. Asci; c. Hymenial hair; d. Paraphyses; e. Ascospores; f. Medullary excipulum; g. External hairs and ectal excip- ulum; h. Moniliform hair-like cells. Scale bars: 20 um (a, b); 10 um (c, d, f-h); 5 pm (e). Geographic distribution. So far, only known from Guangxi Zhuang Autono- mous Region, China. Other specimens examined. CHINA * Damingshan National Nature Reserve, 23°49'N, 108°43'E, alt. 1225 m, on wood surrounded by moss in the evergreen broad-leaved forest, 19 Feb. 2024, Guang R. Zhou, DMS2024022002 (IBK!) (ITS: PP864721; nrLSU: PP859017; tef1-a: PV295998); DMS2024022003 (IBK!) (ITS: PP864722; nrLSU: PP859018). Mao’ershan National Nature Reserve, 25°88'N, 110°37'E, ca 720 m alt., on rotten fallen sticks, 6 April 2024, Yan C. Zhang, MES024040606 (IBK!) (ITS: PQ691392; nrLSU: PQ682446; rpbb7: PV247119; tef1-a: PV295999). Notes. In the phylogenetic tree (Fig. 1), PR damingshanensis and P lutea were grouped together with strong support (MLB = 100%, BPP = 1). Both species share ellipsoid and verrucose ascospores with similar dimensions: P. daming- shanensis measures (21.7) 22.5-32.0 (34.6) x (11.0) 12.1-14.7 (16.5) pm, while P. lutea measures (24.5) 27.0-30.0 (32.0) x (12.2) 13.0-15.0 (16.0) um (Mou and Bau 2021). However, the external surface of P damingshanensis is densely covered with grey tomentum and tree-like ridges that extend to the margin, and its apothecium is larger with a longer and thicker stipe. While the external surface of P /utea has fewer brown hairs and irregular, vascular ridge- like protrusions, with ascomata measuring only 11.0-18.0 mm in diameter. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 35 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa P. sichuanensis is distinguished from P damingshanensis by its hymenium, which is ochre in color and lacks a noticeable covering of black hairs, and its smaller, smooth ascospores (22-26 x 11-13 um) (Zeng et al. 2021). From a microscopic perspective, the ascospores of P damingshanensis, as well as those of P zugazae Calonge & Alb. Garcia, RP melastoma described in older mycological literature, and P. harnischii M. Carbone, Agnello, A.D. Parker & P. Alvarado, are all elliptical and verrucose (Carbone et al. 2015a, 2021). Howev- er, there are significant differences in ascospore size among the four species, P. damingshanensis having larger ascospores [(21.7) 22.5-32.0 (34.6) x (11.0) 12.1-14.7 (16.5) um], while ascospores of P. zugazae [18-22 x 12-14 um] and P melastoma [21.8-25 x 10-12.4 um] are notably smaller. As for P. harnischii, it exhibits the narrower ascospores with a size of 20.2-24.8 x 8.1-11.2 um. Plectania damingshanensis exhibits numerous similarities to species within the Galiella genus, featuring brightly colored ascocarps, highly ge- latinized fleshy tissue, and ascospore surfaces adorned with wart-like or- namentation. However, on the phylogenetic tree, P damingshanensis does not cluster with G. amurensis and G. rufa. Additionally, it is noteworthy that the ascospores of G. amurensis lack oil droplets, and its asci (measuring 400-450 x 12-17 um) and ascospores [measuring (26) 28-37.5 (40) x 11.5-17.5 (18) pm] are larger compared to those of P damingshanensis. Conversely, the asci (measuring 270 x 12 um) and ascospores [measuring (17) 18-21 (23) x 8-10 um] of G. rufa are smaller (Carbone et al. 2015b). Plectania huapingensis G. F. Mou & J. R. Liu, sp. nov. Fungal Names: FN 572466 Figs 5-7 Etymology. The specific epithet refers to the type locality, Huaping National Nature Reserve, in China. Diagnosis. It is characterized by having ascomata, are clustered, disc-shaped, or margin irregular disc-shaped, and the outer surface of the apothecium is adorned with brown-orange to brownish-red particles; under scanning electron microscope (SEM) observation, the elliptical ascospores have a smooth sur- face, with a size of 17.3-21.4 (23.1) x 8.1-10.9 (11.7) um, and 7- or 8-spored asci with a size of 363-388 (425.8) x 9.2-14.1 um. Holotype. CHINA * Guangxi Zhuang Autonomous Region, Guilin City, Huaping National Nature Reserve, 25°63'N, 109°92’E, ca 950 m alt., on buried wood on the ground mixed with evergreen broad-leaved trees and bamboo, 19 July 2024, J. R. Liu (HP2024071903, IBK, holotype!) (ITS: PQ691391; nrLSU: PQ682445; rpb2: PV296007; tef1-a: PV296000). Description. Apothecium clustered, sometimes single, disc-shaped or mar- gin irregular disc-shaped, reaching a diameter of 18-26 mm, with a thickness of 1 mm, subsessile (Fig. 5a). Hymenium surface has wart-like protrusions, some of which are adorned with tiny tomentum, dark brown (Fig. 5b, e). External sur- face shares a similar color to the hymenium, being relatively rough and adorned with brownish-orange to brownish-red particles, which are typically confined to the cup margin but occasionally cover the entire outer surface, relatively tough in texture (Fig. 6c, d). Odor and taste not special. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 36 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa ae « ("hh [ a ee r i tok iw “ —— « | Tee : ba aby cS Figure 5. Ascocarps of Plectania huapingensis (photos by J. R. Liu). a, d. Habitat of specimen HP2024071903 (IBK!), found on senescing or dead bamboo rhizomes; b, c. Habitat of specimen HP2024072014 (IBK!), growing on dead woody branches on the ground or dead bamboo rhizomes; e. Vertical section showing dark violet thin flesh; f. Details of the External hairs are nearly cylindrical, septate, with a diameter ranging from (5.1) 6.0-7.2 (7.5) um, slightly curved, and smooth. They originate from the outer cortical layer and appear brown due to epidermal pigmentation, with walls thick- ened up to 1 um (Figs 6f—-h, 7h). Ectal excipulum consists of a textura angularis, thick-walled cells that are dark brown and measuring 115-135 um, with slight- ly thick walls. Medullary excipulum composed of a textura intricata, measuring 480-550 um in thickness, subhyaline to yellowish white, and consisting of hy- phae 4.1—7.0 um in diameter (Figs. 6c, 7g). Subhymenium is a dense textura in- tricata of closely septate hyphae, olive-colored (2F5), with a thickness of (46.6) 62.2-110 um. Hymenium (329.5) 363.5-422.8 um thick. Asci cylindrical, sub- operculate, 8-spored (occasionally with 7 spores), thick-walled, inamyloid, mea- suring 363-388 (425.8) x 9.2-14.1 um (Figs 6i, j, 7a). Paraphyses 1.9-3.4 um in diameter, filiform, septate, branched, and slightly enlarged or narrowed near the apex (Figs 6l-m, 7f). Ascospores uniseriate, ellipsoid, equilateral, hyaline, smooth, usually contain 1-2 guttules and measuring [37/6/2] 17.3-21.4 (23.1) x 8.1-10.9 (11.7) um (Figs 6d, e, 7b), with Q |... = (1.5) 1.7-2.2 (2.4) and an average Q of 2. Crystals few to abundant, present among the hairs and in the outer sur- face of the ectal excipulum. Hymenial hairs cylindrical, comparable in length with paraphyses, non-septate but with a single septum in the basal part, 2.2-3.3 um wide, concolor with paraphyses due to the homogeneous pigments, tips rounded, straight to slightly curved, unbranched (Figs 6k, 7e). Anamorph unknown. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 37 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa sama hie = ikl OW Figure 6. Features of Plectania huapingensis (photos by J. R. Liu, from HP2024071903, IBK!). a. Section of apothecium; b. External hairs; c. Medullary excipulum; d, e. Ascospores; f—h. External hairs; i, j. Asci; k. Hymenial hair; |, m. Paraphyses. Scale bars: 50 ym (a); 10 um (b, ¢, i-m); 5 um (d-h). Habitat. Half-buried in the litter layer in moist forest mixed with evergreen broad-leaved trees and bamboo. Geographic distribution. By now, only known from Guangxi Zhuang Autono- mous Region, China. Other material examined. CHINA * Guangxi Zhuang Autonomous Region, Gui- lin City, Huaping National Nature Reserve, 25°63'N, 119°92'E, ca 850 m alt., 20 July 2024, J. R. Liu, HP2024072014 (ITS: PQ691390, nrLSU: PQ682444). Notes. P huapingensis and P melastoma share macroscopic characteristics of apothecial external surfaces and margins covered with orange granules. However, upon comparing their microscopic features, it becomes evident that the asci of P. huapingensis [asci: 363-388 (425.8) x 9.2-14.1 um] and its ascospores [17.3- 21.4 (23.1) x 8.1-10.9 (11.7) um] are smaller than those of P melastoma (asci: 400-450 x 12-15 um; ascospores: 21.8-25 x 10-12.4 um) (Agnello and Carbone 2012; Zeng et al. 2021). This difference serves as a distinguishing factor between the two species. Furthermore, with the distinctive feature of its apothecial external surface and margin being covered with orange granules, P huapingensis can be clearly differentiated from other species within the genus Plectania. P huapingensis shares morphological similarities with Korfiella karnika D.C. Pant & V.P. Tewari in both macroscopic and microscopic characteristics. How- ever, molecular sequence data from K. karnika and its allied species remain un- available. Given that P huapingensis aligns with the genus Plectania in molecular phylogeny and micromorphological features, P huapingensis exhibits morpho- logical distinctions from K. karnika, specifically the presence of hymenial hairs (absent in K. karnika) and mature asci containing 7-8 ascospores (versus 2-4 ascospores per ascus in K. karnika), and it should be classified as a member of Plectania s. lat clearly from phylogenetic evidence (Pant and Tewari 1970). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 38 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa A ieee Figure 7. Microscopic structures of Plectania huapingensis (drawings by J. R. Liu). a, d. Asci; b. Ascospores; c. Apex of asci; e. Hymenial hair; f. Paraphyses; g. Medullary excipulum; h. External hairs. Scale bars: 10 um (a, c, d); 5 um (b, e-h). Plectania lutea T. Bau & G. F. Mou, Phytotaxa, 509(3), 288. 2021. Fig. 8 Material examined. CHINA * Guangxi, Guilin city, Mao’ershan National Nature Reserve, 25°87'N, 110°47'E, ca 540 m alt., on rotten fallen sticks, 30 Mar 2024, J. R. Liu (MES2024033005, IBK!) (ITS: PP859394; nrLSU: PP981371; rpb7: PV247120; rpb2: PV296006; tef1-a: PV296001). Geographic distribution. Currently known in Guangxi Zhuang Autonomous Region in China. Notes. The morphological characteristics of this specimen are identical to the original description of P /utea provided by Mou and Bau (2021). Taxonomic Keys for Species Classification of Plectania in China i et ASCOSHOLSS-CIND SONG. 5c oUt mca terre gmt, cents laemm, 9 Un coon rele iB a res oats Beas! 2 - Ascospores inequilateral ellipsoid with a flattened side .................. ee. 3 25 ole ASCOSBOLES SUITACE SIMOOUN:...h. cassyrvadhaecececransss shade ord uatorusheiee Sbucnanenaes 4 - Ascospores surface covered with verrucose ornamentation..................... 5 3. Ascospores one side bearing 13 to 17 transverse grooves, 19.6-27.4 (290 A exe(G OD) 7 Ome 2 SNM, wrath shots Aaah testes th vaca teed allt P. rhytidia - Ascospores one side bearing 8 to 12 rib-like horizontal furrows, 18-22 x A AZADI EY ss Pere creek Ege ert a Snes et ee ach ete en ee i P. platensis 4 Apothecium sessile, ascospores relatively large, 35-42 x 13-18 um........ Babu gareaden Tagen ecetea ee ecetaninn gcc ehote peas md epebs asta cessnidtea set eerade ta Aue Acre P. yunnanensis - Apothecium substipitate, ascospores Smallef...............ccccceeessceeesteeeeees 6 5 Apothecium11.0-18.0 mm, with short stipe, external surface sparsely COVELEC*WATATEOMONM OMIA 2 8.8. cppesateanestcdaeserecsavansntigtpestpeuitbeamenttideess4 P. lutea - Apothecium19-27 mm, with longer and thicker stipe, external surface densely covered with grey tomentum................0::cceee P. damingshanensis MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 39 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa 6 Apothecia external surface and margin covered by orange granules ....... 7 - Apothecia external surface covered or not by orange or brownish granules often restricted at the MAIN .............c cee ceseceeseeceseeeeeseeees P. sichuanensis 7 Asci and ascospores relatively large, asci measuring 400-450 x 12-15 Um, ascospores measuring 21.8-25 x 10-12.4 um............... P. melastoma - Asci and ascospores relatively small, asci measuring 363-388 (425.8) x 9.2-14.1 um, ascospores measuring 17.3-21.4 (23.1) x 8.1-10.9 (11.7) Se ies ek En Rh oes Po NER nT ME A a | P. huapingensis Pseudoplectania aureonigrescens S. A. Chen, D. Li & G. F. Mou, sp. nov. Fungal Names: FN 572467 Figs 9-11 Etymology. The specific epithet “aureonigrescens” refers to the yellowish to orangish color of hymenium surface when young, which changes to grayish black at maturity. Diagnosis. Differs from other known Pseudoplectania species by combina- tion of the following features: sessile ascomata, bowl-shaped, petal-shaped to irregular apothecium up to 25 mm in diameter, hymenium surface yellow-col- ored when young, changing to grey or black when mature; one type of exter- nal hairs straight to distinctly curved or flexuous (not coiled), paraphyses with a mostly straight and rarely diverticulated tip, crystals in the hymenium up to 19.5 um wide, and smooth ascospores without a gelatinous sheath. Holotype. CHINA * Fujian Province, Fuzhou City, Cangshan District, Fuji- an Agriculture and Forestry University, Baizhu Garden, 26°08'N, 119°24'E, ca 50 m alt., on senescing to dead rhizomes of bamboo (probably Oligostachyum lubricum) exposed outside mossy soil walls in a planted bamboo forest, 29 Oct. 2024, D. Li & S. A. Chen (CSA-798, IBK, holotype!) (ITS: PQ895821; nrLSU: PQ863683; rob1: PV247121; tef1-a: PV296005). Description. Ascomata tiny to small, sessile. Apothecium bowl-shaped, pet- al-shaped to irregular, 5-24 mm in diameter, up to 10 mm high; margin entire and involute, cracked after drying. Hymenium surface yellow-tan (2A6), dirty or- ange (4B8) to golden brown (5C7) when young, glabrous, dull, plum grey (8E5), wine grey (9E3) to off-black (4F8) at maturity, color unchanging when bruised, cracked after drying. External surface velvety to strigose, off-black (4F8), densely covered with tomentum or short bristles, without ridges or crests. Stipe absent (Fig. 9f). Subiculum off-black (4F8), cotton-like, dense, soft. Flesh thin, conch grey (22B1), cloud grey (15C1) to pewter grey (10E1). Odor and taste not special. Anamorph not observed. External hairs of one type, hyphoid, septate, cylindrical, straight to distinctly curved or flexuous (not coiled), brass brown (1C7), 4-9.5 um in diameter, with slightly thick to thick walls, up to 2 um thick, tips rounded, sometimes split to two protuberances, unbranched, surface smooth to slightly rough (Figs 10b, 11e). Ectal excipulum of a texture angularis, made up of properly to distinctly thick- walled (up to 3 um) cells, measuring 5.5-23 x 3.5-13 ym, sub-hyaline, tawny (4C5), masala chai brown (4D7) to dark brown (5E8), and dull black (18F8), not or very slightly encrusted, thick-walled, up to 180 um thick (Figs 10a, f, 11h). Medullary excipulum of loose textura intricata immersed in a gelatinous matrix, MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 40 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa ioe te Figure 8. Ascocarps of Plectania lutea, collected from the Mao’ershan National Nature Reserve (photos by J. R. Liu). a-c. Habitat of specimen MES2024033005 (IBK!), growing on dead woody branches on the ground; d. External surface with a detail of the short hirsute hairs. Scale bars: 10 mm (a-d). up to 335 ym thick, with hyphae septate, subhyaline or pale yellow (1A5), thin- walled, flexuous, branched, 2-4 (6) um wide, tawny (4C5) to light brown (4C7), dark coffee brown (4E8) at low magnification (x100, Fig. 10a, e). Subhymenium of a dense textura intricata of septate hyphae, 2-4 um wide, thin-walled, concolor with medullary excipulum, up to 80 um thick, dark coffee brown (4E8) to dark brown (5E8) at low magnification (Fig. 10a). Asci cylindrical, operculate, with a curved or flexuous, tapered base, 8-spored, 195-335 x 10.5-14 um, constricts at the junction of two ascospores, non-amyloid, apex mostly rounded, sometimes with a conical, eccentric, or non-eccentric protrusion (Figs 10g-i, n, 0, 11a, b). Ascospores uniseriate, smooth, with thick walls, globose to subglobose, sub-hy- aline to pale yellow (1A5), [60/4/2] (8.5) 9.5-11.5 (14) x (8.5) 9-11 (14) um, Q vaue = (0-95) 1-1.1 (1.13) including the spore wall, non-amyloid, with or without one to multiple, globose to subglobose contents (Figs 10p—s, 11g). Paraphyses abundant, filiform, not or slightly extending the length of asci, septate, subcylin- drical, simple to bifurcate, straight to slightly curved, sometimes anastomosed, branched both from the lower part and tips, dark khaki brown (5D6) to dark coffee brown (4E8) due to amorphous pigments, (1) 1.5—-3 um in diameter, tips round- ed to finger-shaped, rarely with few protuberances or notches, mostly straight, sometimes slightly enlarged or curved, unbranched or branched 1-2 times MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 4 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Figure 9. Ecological and macroscopic characteristics of Pseudoplectania aureonigrescens, photos by S. A. Chen & D. Li; c—e. from holotype CSA-798 (IBK!); f-j. from CSA-797 (IBK!). a. Habitat where the specimen CSA-798 (IBK!) grew (the white arrow indicates the growth site of ascomata), senescing or dead rhizomes of bamboo exposed outside mossy soil walls; b. Habitat where the specimen CSA-797 (IBK!) grew (white arrows indicate growth sites of ascomata), a planted bamboo forest; c-e. Mature ascomata with a smooth, greyish-black hymenium; f. Off-black external surface covered with abundant tomenta; g. Details of the hairy external surface; h. Vertical section including greyish thin flesh; i, j. Young ascomata with discoid to petal apothecium, dirty orange hymenium, and velvety to strigose external surface. Scale bars: 10 mm (c, e); 5 mm (d, f, h-j); 2 mm (g). (Figs 10j, k, 11d). Hymenial hairs cylindrical, long as the paraphyses, non-septate but with a single septum in the basal part, (1.5) 2—3.5 um in diameter, concolor with the paraphyses due to the homogeneous pigments, tips rounded, straight to slightly curved, unbranched (Figs 10I, m, 11c). Subiculum dark coffee brown MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 42 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Figure 10. Microscopic structures of Pseudoplectania aureonigrescens, photos by S. A. Chen, from holotype CSA-798 (IBK!). a. Cross section through apothecia; b. External hair; c. Hypha of subiculum; d. Hymenium with crystals (white arrows indicate the position of crystals); e. Medullary excipulum in Congo red; f. Ectal excipulum in Congo red; g. Ascus with mature ascospores in Meltzer’s reagent; h. Empty ascus with opened operculum in Congo red; i. Immature ascus without visible ascospores in Congo red; j, k. Paraphyses in Congo red; I, m. Hymenial hair in Congo red (the yellow arrow in m indicates the basal septa of the hymenium hair); n. Rounded apex of asci in Congo red; o. Apex of asci with a conical eccentric protrusion; p, q. Ascospores in distilled water; r, s. Ascospores in Meltzer’s reagent. Scale bars: 100 um (a); 50 um (b-i, k, m); 20 um (j); 10 um (I, n-s). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 43 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa d Figure 11. Microscopic structures of Pseudoplectania aureonigrescens, drawing by S. A. Chen, from holotype CSA-798 (IBK!). a. An ascus with 8 ascospores; b. Empty ascus with its operculum; c. Hymenium hairs; d. Paraphyses; e. External hairs; f. Hypha of subiculum; g. Ascospores; h. A part of ectal excipulum. Scale bars: 50 um (a-f sharing same bars); 10 um (g, h). (4E8) to dark brown (5E8), septate, thick-walled, straight to curved or flexuous, mostly unbranched, rarely branched once, not encrusted, 4—6.5 um in diameter, with a rounded tip (Figs 10c, 11f). Crystals in the hymenium rectangle to angular, sometimes cracked, roughly parallel to other elements in hymenium, concolor with the paraphyses, mainly of two types, although intermediates forms may MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 44 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa exist: 1) thin, 1.9-4.5 um in diameter, up to 24.5 um long; 2) thick, 7-19.5 um in diameter, up to 27.5 um long (Fig. 10d). Habitat. Growing in scattered to gregarious groups on senescing to dead rhi- zomes of bamboo (probably Oligostachyum lIubricum) exposed outside mossy soil walls in bamboo forests. Geographic distribution. So far, only known from Fujian Province, China. Other material examined. CHINA * Fujian Province, Fuzhou City, Cangshan District, Fujian Agriculture and Forestry University, Baizhu Garden, 26°08'N, 119°24'E, ca 50 m alt., on senescing to dead rhizomes of bamboo exposed outside mossy soil walls in a planted bamboo forest (probably Oligostachyum lubricum), 12 Nov. 2023, S. A. Chen, CSA-797 (IBK!) (ITS: PQ863322; LSU: PQ863491; tef7-a: PV296004). Notes. In the phylogenetic analyses, Pseudoplectania aureonigrescens clus- tered with Ps. affinis and Ps. globospora in clade III. The sessile ascomata with entire margin, velvety, rough, blackish external surface, dark hymenium sur- face at maturity, and globose to subglobose ascospores without gelatinous sheath make them easily confused (Carbone et al. 2014a; Zeng et al. 2024). But it can still be recognized by some distinguishing characters. With respect to Ps. affinis, Ps. aureonigrescens has variously shaped apothecium, dull hyme- nium surface at maturity, hymenium with crystals, existing obviously curved or flexuous external hairs, and paraphyses rarely with protuberances or notches. Compared with Ps. globospora, Ps. aureonigrescens can be distinguished by its variously shaped apothecium, one-typed external hairs, smaller cells in ectal excipulum (5.5—23 x 3.5-13 um vs. 17-22 x 14-17 um), existing crystals in hymenium, slightly smaller ascospores (9.5-11.5 x 9-11 um vs. 10.5-12.5 x 9.8-13.5 pm), and also by its lignicolous habitat. Also, the lineage belongs to the newly established species, obtains strong support, and is enough to sepa- rate itself from its relatives by genetic distance. Ecologically, Pseudoplectania aureonigrescens and Ps. mystica, teleomorphs of both, share a similar habitat and are geographically close in distribution, grow- ing groups on senescing to dead rhizomes of bamboo (Lin et al. 2024). It is also notable that Ps. aureonigrescens shares similarities with Ps. mystica in its sessile ascomata, velvety, dark external surface, crystals in hymenium, existing wavy or flexuous external hairs, paraphyses rarely with protuberances or notches. How- ever, besides its distant of genetic, this new species is clearly differentiated by ascomata of various shapes, dull hymenium surface when mature, existing branched subiculum, a lack of crystals in ectal excipulum, and slightly smaller ascospores without sheath (9.5-11.5 x 9-11 um vs. 11-13 x 10.5-12 um). Key to accepted species of Pseudoplectania Pseudoplectania species lacking molecular data or clear original diagnoses are not included (viz., Pseudoplectania carranzae, Ps. kumaonensis, Ps. spongiosa, and Ps. stygia). Ps. sinica is also excluded due to the lack of teleomorph. 1 Mature ascospores with slight ornamentation......Pseudoplectania ryvardenii - Mature ascospores without ornamentation ...............ccccccccccessseceeeeesteeeeees 2 2 Mature ascospores with a gelatinous Sheath ..............cccccccccesesseceeeeeeeees 3 - Mature ascospores without a gelatinous sheath.............. ccc cccceeeteeeeeee 9 MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 45 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa 3 Mature ascospores with centrally arranged gelatinous sheath; ectal excip- ulum composed of subglobose to elongated cells.................. Ps. lignicola - Mature ascospores with eccentrically arranged gelatinous sheath; ectal excipulumwithout-elongated Cells... 24 c-ccccncssteceossenacce yreetedytretrarenchncgenes 4 4 Ascomata distinctly stipitate; paraphyses tips often hooked ..................... ca IR ein re ie Cee at 7 RE Mee for we MRO re PN. AEE We Ps. melaena complex - Ascomata sessile to shortly stipitate; paraphyses tips straight to curved, DUE MOBO OKC Ce rhe ae Bastar eaecsmt etcetera oes het totes ste Potts caer tas ees sab 5 Be “SEXterial lair StlexuoUS:OrOONeUs isk ccc yk bch cs ade OS ehed ale eietaaees 6 - External hairs straight to curved or flexuous but not coiled....................... 7 Gy mika GesCryStals. AUSCMUr aceccucsonpracetteseussdsancresemeeciseseoe Ps. nigrella complex - Large crystals present; known from South Africa and Madira island.......... Pe FE ert SIRE FTE ISSN Ph ot MPS SATU PORE URNS Sa 5 ODE E SONU Er Ps. africana 7 Paraphyses tips mostly bifurcate to trifurcate ........... ccc ecceceeseceeneeens 8 - Paraphyses tips simple to bifurcate.........0. ccc ec ceeseeeeseeees Ps. mystica 8 Apothecium up to 10 mm; crystals absent in hymenium; known from Eu- [ca] Of yee. Recs eer ARK hott tn ey reve. (ured Rennie eae ew Marien. t Smregt Rent erm. Ps. ericae - Apothecium up to 30 mm; crystals present in hymenium; known from Oce- | Fe ar ea ee he Reh Ale aimee es._ W shal ae rhe. A aR Ps. tasmanica 9 Apothecium mostly up to 10 mm; external hairs flexuous or coiled ........... Er CIN et Peer any RUT eat ECOL S TORT OemiS rae Ps. episphagnum - Apothecium available more than 10 mm; external hairs straight to curved Omtiexious#DULMOLCONE aye et. Fe Rnd Se ita pain Ek isalee 5 esse 10 NOs = EXTEN Mai SAWwOrky DSO! 2.4.8 20.4. 092s. den Aude es Nets ded Nevada 835: Ps. globospora =" CSEXTenMall Meas OMe shy POG mas cease tas ete taten Se cea Pot tars cy eudeanes drassPue teen wii 11 11. Hymenium surface shiny or polished when mature; external hairs straight to slightly curved; crystal absent in the hymenium...................... Ps. affinis - Hymenium surface dull when mature; external hairs straight to distinctly curved or flexuous; crystals present in the hymenium..... Ps. aureonigrescens Urnula ailaoshanensis J. R. Lu, Y. Zhang & Q. Zhao, Phytotaxa 619 (1): 91. 2023. Fig. 12 Material examined. CHINA * Guangxi Zhuang Autonomous Region, Guilin City, Mao’ershan National Nature Reserve, on rotten fallen sticks surrounded by mosses, ca 1990 m alt., 11 Aug 2023, J. R. Liu (MES2023081104, IBK!) (ITS: PQ187434; nrLSU: PQ187435; rpb1: PV247122). Habitat. On the fallen, rotten wood surrounded by mosses. Geographic distribution. Currently known in Yunnan Province and Guangxi Zhuang Autonomous Region in China. Notes. According to Lu et al. (2023), Urnula ailaoshanensis was originally reported in Yunnan Province, China, and it is recognized by saucer- or bowl- shaped cup apothecia, reddish-brown to ocher hymenium with glabrous sur- face, one type of external hair, cylindrical asci with a tapered or attenuated base, and ellipsoid, sometimes reniform-like ascospores. In this study, these features are mostly consistent with the observation of our specimens. Howev- er, compared to the type specimens, our specimens have darker hymenium sur- face, smaller asci (340-450 x 21-26 um vs. 465-600 x 21-30 um), and slightly MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 46 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa ; f. os 7 ins a “—s photos by S. A. Chen, from IBKMES2023081104 (IBK!). a. Fresh ascomata in a field, growing on rotten wood surrounded by mosses; b. Fresh ascomata with a dark hymenium surface and a reticulate or corrugated external surface; c. Cross Section through apothecia; d. Subhymenium and medullary excipulum; e. Paraphyses; f. External hairs; g. Hymenial hairs; h. Ascospores in water; i. apical apparatus of ascus with developed ascospores; j. An ascus with immature ascospores. Scale bars: 100 um (c, j); 50 um (d); 20 ym (f-i); 10 um (e). longer ascospores (29-38.5 x 13.5-17 um vs. 29-34.5 x 11.5-17 um). Such variable color of hymenium surface may be associated with the maturity of ascomata and/or environmental factors, which makes U. ailaoshanensis easily confused with other Urnula species, viz., U. versiformis Y. Z. Wang & Cheng L. Huang, U. himalayana K. Das & D. Chakr. But U. ailaoshanensis can be iden- tified through these microscopic features: one type of external hair, ellipsoid to reniform-like ascospores, and molecular evidence (Wang and Huang 2015; Wang et al. 2018). In addition, we found a few hymenial hairs in the hymenium, cylindrical, non-septate, up to 3.5 um in diameter, with rounded tips (Fig. 12g). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 47 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Urnula auricularioides S. A. Chen, D. Li & G. F. Mou, sp. nov. Fungal Names: FN 572194 RIgSs Mosul Etymology. The specific epithet “auricularioides” refers to the appearance and habitat of this species similar to Auricularia species. Diagnosis. Differs from other known Urnula species by the combination of the following features: external surface velvety to warty and with ridges, hymenium surface with verrucose bulges to intestinal folds when mature, two types of ex- ternal hairs composed of strongly encrusted true hairs and smooth, slender short hairs, ellipsoid to bean-shaped, smaller ascospores (25—30.5 x 9-11 um). Holotype. CHINA * Fujian Province, Fuzhou City, Minhou County, Sandiejing Na- tional Forest Park, 26°26'N, 119°17'E, ca 550 m alt., on rotten fallen sticks or wood surrounded by mosses on riverside in moist forest mixed with evergreen broad- leaved trees and bamboo, 9 Nov. 2023, S. A. Chen (CSA-455, IBK, holotype!) (ITS: PQ489315; LSU: PQ187431; rob1: PV247123; rpb2: PV296008; tef1-a: PV296003). Description. Ascomata small to medium-sized, subsessile to distinctly stipi- tate. Apothecium cupulate, bowl-shaped to long funnel-shaped, up to 15-60 mm in diameter, 20-30 mm in high; margin entire and involute. Hymenium surface glabrous, initially without ornamentation or wrinkled, with verrucose bulges to intestinal folds at maturity, burlywood (3E8), yellowish-brown (4D8) to rosy brown (6E8), lighter in margin, color unchanging when bruised, darkening and cracked after drying. External surface velvety, densely dotted with droppable and fine grey (9D1) warts, with radial and forked ridges or crests not reaching margin, tapering towards the base or stipe, slate grey (12D1), dim grey (21D1) to off-black (4F8), covered by subiculum. Stipe (when present) up to 5 mm in di- ameter, 19 mm in high, uneven or foveolate, concolor with the external surface. Subiculum off-black (4F8), mussel base byssal-like, dense, soft, and fibrous, mixed or flexuous. Flesh thin, gelatinous, elastic, coco-nata white (1A1). Odor and taste not special. Anamorph not observed. External hairs mainly of two types, although intermediates forms may exist: 1) short, hyphoid, septate, cylindrical, slightly to distinctly curved or flexuous, subhyaline with a very pale yellow tint but occasionally dark brown (5E8) to dull black (18F8) for a granular intracellular pigment in part, thin-walled, 2-4 um in diameter, subcylindrical, tips rounded to slightly subcapitate, surface smooth (Figs 14b, 15e); 2) long, true hairs, undulated, almost non-septate, some branched, tips rounded to truncated, surface unevenly covered by a granular to agglomerate encrustation, 4-7 um in diameter, with moderately thick walls, up to 1 um thick, light brown (4C7) or grayish olive (29D5) to dark brown (5E8) and off-black (4F8) (Figs 14c, 15f). Ectal excipulum of a textura subglobulosa to tex- tura angularis, textura irregularis made up of properly thick-walled (up to 4 um) cells up to 5-35 um in length and/or width, tawny (4C5) to dark brown (5E8) and dull black (18F8), not or slightly encrusted, 95-240 um thick (Fig. 14f). Medullary excipulum of a loose textura intricata immersed ina highly gelatinous matrix, approximately 260-510 um thick (Figs 14e, 15h), mainly with two types of hyphae: I) short, septate, subhyaline or pale yellow (1A5), thin-walled, arbori- form and flexuous, 2—3.5 um in wide; II) long (relative to type |), septate, concol- or with type | hyphae, thin-walled, cylindrical, almost no branched, roughly par- allel to subhymenium, 1.5—3 (5) um in wide. Subhymenium of a dense textura MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 48 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Figure 13. Ecological and macroscopic characteristics of Urnula auricularioides, photos by S. A. Chen; b, d, e, f, g. from holotype CSA-455 (IBK!); c, h. from CSA-454 (IBK!). a. Habitat where the specimen grew, a moist forest mixed with ever- green broad-leaved trees and bamboo on riverside; b, c. Fresh ascomata with a hymenium surface that is unfolded when immature and variously wrinkled when mature, growing on rotten branch surrounded by mosses; d, f. Dried materials; g. Hymenium surface with dense verrucose bulges; h. Velvety external surface of fresh specimens with greyish warts. Scale bars: 10 mm (b-h). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 49 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa te -—_ —= ee te ae. Ap Figure 14. Microscopic structures of Urnula auricularioides, photos by S. A. Chen, from holotype CSA-455 (IBK!). a. Cross section through apothecia; b. Type 1 of external hairs; c. Type 2 of external hairs; d. Hymenium and subhymenium in Congo red; e. Medullary excipulum with two types of hyphae (type | indicated by black arrows; type II indicated by white arrows) in Congo red; f. Ectal excipulum and external hairs in Congo red; g. Asci with mature and immature ascospores; h. An ascus with 8 developed ascospores in Meltzer's reagent; i. Apical apparatus of ascus in Congo red; j. Ascospore in distilled water; k. Ascospore in Meltzer’s reagent; |. Ascospore in Congo red; m. Straight to hooked hymenial hair and paraphyses; n. Tips of paraphyses in Congo red; 0. Hymenial hair in Congo red; p. Hypha of subiculum. Scale bars: 100 um (a); 50 um (g, 0, p); 20 um (b, d—-f); 10 pm (i-n). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 50 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa (IBK!). a. An ascus with 8 developed ascospores; b. Hymenium hairs; c. Ascospores; d. Paraphyses; e. Type 1 of external short hairs; f. Type 2 of external long hairs; g. Hypha of subiculum; h. A part of ectal excipulum. Scale bars: 50 um (a); 20 um (b, d, f, g, h; b, d sharing the same bars); 10 um (c, e). intricata of closely septate hyphae, surface smooth, up to 3 um in diameter, thick-walled, tawny (4C5) to light brown (4C7), dark brown (5E8) at low magni- fication, 50-220 um thick (Fig. 14d). Asci cylindrical, operculate, with a tapered or attenuated base, rounded at apex, 8-spored, 240-410 x 11-18 um, non-am- yloid (Figs 14g-i, 15a). Ascospores uniseriate, smooth, sometimes curved, thick-walled, rounded at both ends, heterogeneous in shapes depending on the spacial position and growth stage of the spores, pale yellow (1A5) or char- treuse (30A7) to grayish olive (29D5), incipiently ellipsoid to allantoid with walls up to 3 um thick, becoming oblong to bean-shaped with walls 1-2.5 um thick, [40/4/2] (23.5) 25-30.5 (32) x (8.5) 9-11 (11.5) um, Q = (2.31) 2.46-3.1 values MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 51 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa EHT = 10.00 kV Signal A= SE1 Date :5 Nov 2024 EHT = 10.00 kV Signal A = SE1 Date :13 May 2024 WD = 8.5mm Mag= 2.50KX Time :16:48:04 WD = 8.5mm Mag= 2.50KX Time :11:24:05 EMT = 10.00 kV Signal A = SE1 Date :11 Dec 2024 ! EHT = 10.00 kV Signal A = SE1 Date :30 May 2024 WD = 8.5mm Mag= 2.50KX Time :18:55:24 WD= 85mm Mag= 2.50KX Time :15:36:03 Figure 16. Scanning electron micrograph of basidiospores of the four new species of Sarcosomataceae from China (photos by J. R. Liu). a. Plectania huapingensis; b. Plectania damingshanensis; c. Pseudoplectania aureonigrescens; d. Urnula auricularioides. Scale bars: 2 um (a-d). (3.3), non-amyloid, with 0-5 lipid bodies (Figs 14i, k, |, 15c). Paraphyses fili- form, not or slightly extending the length of asci, septate, cylindrical, straight to slightly curved, dark coffee brown (4E8) due to the extracellular amorphous pigments, darkening toward the upper part but deposited by dense off-black (18F8) pigments near the base, (1) 1.5-2.5 um in diameter, tips rounded to fin- ger-shaped, sometimes branched 1-3 times (Figs 14m, n, 15d). Hymenial hairs abundant, cylindrical, long as the paraphyses, non-septate, 2.5-3.5 (4.5) um in diameter, concolor with the paraphyses due to the homogeneous pigments, agglutinating or intermingle with the paraphyses to bundles, tips rounded to slightly subcapitate, straight, curved, or flexuous to almost hooked (Figs 140, 15b). Subiculum grayish olive (29D5), tawny (4C5), and rosy brown (6E8) to dark brown (5E8), septate, with a slightly thick wall, up to 1 um thick, slightly curved, unbranched, not or slightly encrusted by an extracellular dark brown (5E8) to dull black (18F8) pigment, (2) 4.5-6 pm in diameter (Figs 14p, 15g). Habitat. Growing in gregarious groups on rotten fallen branches or twigs surrounded by mosses on riverside, half-buried in the litter layer in moist forest mixed with evergreen broad-leaved trees and bamboo. Geographic distribution. So far, only known from Fujian Province, China. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 52 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Other material examined. CHINA * Fujian Province, Fuzhou City, Minhou County, Sandiejing Forest Park, 26°26'N, 119°17'E, ca 550 m alt., 9 Nov. 2023, S. A. Chen, CSA-454 (IBK!) (ITS: PQ489472, LSU: PQ187433; tef1-a: PV296002). Notes. Urnula auricularioides is morphologically similar to Urnula versiformis, sharing most characteristics but distinctly differing from the latter by its slender external short hairs (2-4 um in wide vs. 5-7 pm in wide) and small ascospores (25-30.5 x 9-11 um vs. 30-35 x 10-12 um) (Wang and Huang 2015). U. auricu- larioides is also similar to U. campylospora and U. ailaoshanensis in morphology. However, U. campylospora produces reticulate external surface and two-typed external hairs composed of heavily encrusted short hairs (5-6 um in wide) and mainly smooth true hairs (Carbone and Agnello 2013), while U. auricularioides is evidently distinguished by its ridged external surface and smooth, slender exter- nal short hairs (2-4 um in wide) and strongly encrusted true hairs; U. ailaoshan- ensis characteristically produces reticulate external surface and one-typed ex- ternal hairs (Lu et al. 2023). In the phylogenetic analysis, Urnula auricularioides is relatively close to Urnula himalayana, but it can be easily separated on mul- tiple morphological characteristics. U. auricularioides produces entire margin, smooth and slender external short hairs (2-4 um in wide vs. 6-7 um in wide), and smaller ascospores (25-30.5 x 9-11 um vs. 24-33.6 x 10-13.8 um). Addi- tionally, the external surface appearance of both species is also clearly different. U. himalayana has a reticulate pattern without warts formed by vertically joined folds or veins (Wang et al. 2018), whereas U. auricularioides possesses a ridged surface covered with vertical folds or crests with visible warts. Key to accepted species of Urnula Urnula species lacking molecular data or a clear original diagnosis are not in- cluded (viz., Urnula brachysperma, U. groenlandica, U. mexicana, U. microcrater, and U. viridirubescens). 1 Mature ascospores ellipsoid to allantoid, bean-shaped or reniform-like...2 - Mature ascospores ellipsoid to short clavate ........... ccc cccceesseceeeeentseeeeee 6 2 Exterrial Surbace velvety: tO War ysncsaccsncosstrers dead snowsuitetrosat decad dusnunetraead Adsad gust 3 - External surface velvety without Warts ............:ccccccssscsssecsseceseecsseenseeesseeenees 5 3. External short hairs mostly smooth, slender............ Urnula auricularioides - External short hairs encrusted to heavily encrusted, thick...................06. 4 4 Ectal excipulum 40-60 um thick; base vertically corrugated; mature asco- SNOlESs 227-3 Line Sul = PaO LIAN eae Aten ee ateronsieee U. campylospora - Ectal excipulum 50-125 um thick; base ridged; mature ascospores 26-32 BAD Se 1G: (MIT Veo Scrca Py cei locke estas aa tai en Meee Alon Ne. erase ses he U. versiformis 5 PIWO-TYpedtexterniglaiall Sit... .ccctetictaebe cial svnabiel Maton Meese. tts U. himalayana — ~@Ohetyped-external Nairs:... isch at ieatasteeestes U. ailaoshanensis 6 External surface with warts or tubercles .............cccccccscecsseeesseceesseeesseeeaes 7 - External surface with patches of issue or toMeNtUM............ cece eee eeeeee 8 7 Ascomata with a well-developed stipe; mature ascospores thick-walled; ectal excipulum with crystals ................ceceeessecesreeeeteeeesneees U. mediterranea - Ascomata with a short to well-developed stipe; mature ascospores thin- walled; ectal excipulum without crystals ............. eee eeeeeeeeteeeeee U. hiemalis MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 53 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa 8 Ascomata sessile; external surface with patches of grey tomentum within TES} ARGS AC MTA Ss ered esscct ce Pied sete econ oe eesone te aren sde tease U. padeniana - Ascomata subsessile to stipitate; external surface with yellow to brown DAtCNES-OTASSUC iW cctssa te: testes anny trator tte. aioe? ee AMR cctgn eae eagiitei eet 9 9 Asci short; paraphyses slender .................cceeeeeeeees U. craterium complex - Asci obviously long; paraphyses thick ...............cccceceeseeeeeees U. subcrateria Discussion As a cosmopolitan family, Sarcosomataceae has been explored for many decades, representing an interesting group of fungal taxa capable of grow- ing in tropical, subtropical, temperate, and even cold climates (Carbone et al. 2013a, 2014a; Blanchette et al. 2016; Van Vooren and Mauruc 2020). Previous studies of sarcosomataceous species were mainly confined to temperate regions (Dissing 1981; Calonge et al. 2003; Carbone et al. 2013a, 2014a, 2015a; Glejdura et al. 2015; Pfister and LoBuglio 2018; Saitta 2020; Popov and Carbone 2021; Carbone and Rocha 2024). The current study lays emphasis on updating the diversity, taxonomy, and delimitation of sarcoso- mataceous fungi in subtropical China, based on ecological observations, morphological characteristics, and multi-gene phylogenetic analysis. Four new species were scientifically verified, namely Plectania damingshanen- sis, P huapingensis, Pseudoplectania aureonigrescens, and Urnula auricu- larioides. In addition, two taxa with new distribution records were revealed: Plectania lutea and Urnula ailaoshanensis. An updated list of Sarcosoma- taceae species in China is shown in Table 2, in which a total of 25 species were recorded in the subtropical region, except for Urnula craterium, indicat- ing the extremely rich diversity of these ascomycetes in subtropical China. Previous studies related to the classification of Sarcosomataceae only per- formed one- or two-locus phylogenetic analyses. In the phylogenetic analysis using ITS and nrLSU sequences by Carbone et al. (2013a), the results revealed that six monophyletic clades corresponded well to the six accepted genera: Donadinia, Galiella, Plectania, Pseudoplectania, Sarcosoma and Urnula. How- ever, judging from our phylogenetic result, Donadinia, Pseudoplectania, Sar- cosoma, and Urnula appeared to be monophyletic with significant support, while Galiella and Plectania did not form two distinct monophyletic groups but instead composed a single complex: the Plectania s. lat. clade. Moreover, with wider taxon sampling, Zeng et al. (2021) also proposed a similar division based on an ITS + nrLSU dataset including 126 Sarcosomataceae taxa. How- ever, upon further comparison with our topology, there are some differences in the division of Galiella and Plectania. In their study, Plectania was divided into three divergent clades (Plectania I-III), and Galiella was represented only by G. rufa, grouped as sister to Plectania clade |. In our study, the Plectania s. lat. clade is composed of several lineages, including Galiella, which is rep- resented by two species (G. amurensis and G. rufa). With the application of more extensive shallow sequencing data, our study revealed inconsistency in the divisions between our multi-locus phylogenetic analysis and the two com- parative cases. This suggests that ITS-based or ITS + nrLSU-based analyses, when based on limited specimen sampling, are insufficient to investigate the relationship between Galiella and Plectania. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 54 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Table 2. List of Sarcosomataceae taxa reported from China. A tick after taxa indicates that its distribution in China is supported by molecular data. Location of taxa is based on relevant reports with or without molecular data. The letters STR, TE, and TR in climate are indicated as subtropical, temperate, and tropical, respectively. Genus Taxa Location Climate References Donadinia (3) D. echinacea V Yunnan STR Zeng et al. (2021) D. cf. helvelloides Tibet (Nyingchi) STR Zhuang (2004) D. nigrella Tibet (Nyingchi) STR Xu (2000); Zhuang (2004) Galiella (2) G. amurensis V Heilongjiang, Jilin, Sichuan, Tibet (Nyingchi) | STR, TE Zhuang (2004); Popov and Carbone (2021) G. sinensis Fujian, Yunnan STR Cao et al. (1992); Zhuang (2004) Plectania (10) P. damingshanensis V Guangxi STR This study P huapingensis V Guangxi STR This study P lutea V Guangxi STR Mou and Bau (2021) P melastoma Tibet (Nyingchi) STR Xu (2000) P nigrella Tibet (Nyingchi) STR Xu (2000) P platensis Hubei STR Xu (2000); Zhuang (2004) P. rhytidia Tibet (Nyingchi) STR Xu (2000); Zhuang (2004) P. sichuanensis V Sichuan STR Zeng et al. (2021) P submilleri V Yunnan STR Hyde et al. (2024) P. yunnanensis Yunnan STR Zhuang and Wang (1998); Zhuang (2004) Pseudoplectania (5) | Ps. aureonigrescens V Fujian STR This study Ps. globospora V Yunan STR Zeng et al. (2024) Ps. mystica V Zhejiang STR Lin et al. (2024) Ps. nigrella V Heilongjiang, Hubei, Sichuan, Tibet (Nyingchi) | STR, TE Xu (2000); Zhuang (2004); Lin et al. (2024) Ps. sinica V Fujian STR Zhang and Zhang (2020) Urnula (6) U. auricularioides V Fujian STR This study U. ailaoshanensis V Guangxi, Yunnan STR Lu et al. (2023); This study U. campylospora Guangxi, Hainan, Yunnan STR, TR Zhuang (2004) U. craterium Heilongjiang TE Zhuang and Wang (1998); Zhuang (2004) U. subcrateria V Yunnan STR Lu et al. (2023) U. versiformis V Taiwan STR Wang and Huang (2015) As mentioned, the Plectania s. lat. clade encompasses both Galiella and Plectania. Species within this complex group exhibit a wide range of mor- phological variation, offering several interspecific distinguishing features. The main diagnostic characteristics for differentiating Plectania species in- clude apothecium size, stipe length, color of the hymenium surface, density of tomentum on the external surface, presence of one or two types of exter- nal hairs, and ascospore size and ornamentation. Two new members were collected from Guangxi: Plectania damingshanen- sis and P. huapingensis. The former is distinguished by cupulate ascomata, a brownish-orange to light-brown hymenium surface, a velvety and ridged exter- nal surface, and elliptical ascospores. Plectania huapingensis is recognized by disc-shaped or irregularly margined disc-shaped ascomata, an external surface adorned with brown-orange to brownish-red particles, and smooth elliptical as- cospores. The macroscopic and microscopic structures of the two species are similar to those of Galiella species in possessing gelatinous flesh and a bright hymenium surface. Further clarification requires more refined morphological analysis and accurate molecular systematic studies. Additionally, on intergeneric aspects, the delimitation between Galiella and Plectania lacks clarity. In existing research, species with Plectania-like and MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 55 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa dark-colored apothecia, thin medullary excipulum, and ellipsoid ascospores were placed in Plectania, while Sarcosoma-like and bright-colored apothecia, thick medullary excipulum, and verrucose ellipsoid ascospores were placed in Galiella (Korf 1957a). Recently, gelatinous apothecia, verrucose ascospores, and smooth external hairs were recognized as the main distinguishing features of the genus Galiella (Carbone et al. 2015b). Exceptionally, RP damingshanensis and P lutea blur these criteria, as they possess gelatinous apothecia; yellow, light brown, or orange-brown hymenium surfaces; and verrucose or vermicular ellipsoid ascospores with smooth external hairs. Notably, P damingshanensis also has a thick and watery medullary excipulum. From a phylogenetic point of view, Galiella and Plectania species cluster into a monophyletic group with strong support (MLB = 78%, BPP = 1). Moreover, the two Galiella lineages—G. amurensis and G. rufa—did not even cluster together with strong support. Based on both morphological characters and topological evidence, this seems to indicate that there is no obvious heterogeneity at the genus level between Galiella and Plectania. However, we adopt a cautious posi- tion regarding this intricate group until more specimens are available and more convincing morphological criteria are proposed. In Pseudoplectania, species within this monophyletic genus share highly similar macroscopic features and are recognized by tiny to medium-sized as- comata and globose to subglobose ascospores. The diagnostic characters, combining macro- and micro-morphological features, chiefly concern apothe- cium size, whether sessile or stipitate; paraphyses with or without hooked tips; one- or two-typed and coiled or straight external hairs; and ascospores with or without ornamentation and the position of the gelatinous sheath. Two other potentially useful characteristics are the presence or absence of the gelatinous sheath and the presence of large or small crystals in the hymenium, though these should be considered in conjunction with the freshness of collections. Subclade | is represented only by the Ps. melaena complex, showing distinctly stipitate fruitbodies, paraphyses with often hooked tips, straight to slightly curved external hairs (but not coiled), and ascospores with an eccentrically positioned gelatinous sheath (Van Vooren et al. 2013). With- in subclade II, Ps. episphagnum and the Ps. nigrella complex (including Ps. lignicola) form a phylogenetically affinitive group. Their teleomorphs commonly share the characteristics of sessile to shortly stipitate fruitbod- ies, paraphyses tips not hooked, the presence of coiled external hairs, and ascospores with or without eccentrically to centrally arranged gelatinous sheaths, except for Ps. sinica (Carbone and Agnello 2012; Carbone et al. 2013a, 2014a). Subclade IIl is composed of three identified species—Ps. af- finis, Ps. aureonigrescens, and Ps. globospora—which possess sessile fruit- bodies, paraphyses tips not hooked, straight to curved or flexuous external hairs (but not coiled), and smooth ascospores without any surrounding ge- latinous sheath (Carbone et al. 2014a; Zeng et al. 2024). Among these, Ps. aureonigrescens, collected from Fujian, differs from the other two related species by its bowl-shaped, petal-shaped to irregular apoth- ecium; dull hymenium surface when mature; one-typed external hair, straight to distinctly curved or flexuous (not coiled); and the presence of crystals in the hymenium. Subclade IV comprises Ps. africana, Ps. ericae, Ps. mystica, and Ps. tasmanica, which feature sessile to shortly stipitate fruitbodies, paraphyses MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 56 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa tips not hooked, straight to curved or flexuous external hairs (but not coiled), and ascospores with an eccentrically arranged gelatinous sheath (Van Vooren et al. 2013; Carbone et al. 2014a; Carbone and Rocha 2024; Lin et al. 2024). In Urnula, species are distinctive in their dark, bowl-shaped to funnel-shaped ascomata and smooth, elliptical to allantoid ascospores when mature. Among these species, the dominant diagnostic characters are stipe length; the appear- ance of the external surface or base; the presence of one or two types of ex- ternal hairs; and the shape of the spores. The collection YC2023090928 (IBK), collected from Guangxi, belongs to Urnula. Unfortunately, we did not find any visible ascospores due to the immaturity of the material. Within subclade A, U. auricularioides, collected from Fujian, is charac- terized by a velvety to warty external surface with ridges and a hymenium surface with verrucose bulges to intestinal folds when mature; two types of external hairs composed of strongly encrusted true hairs and smooth, slender, short hairs; and ellipsoid to bean-shaped ascospores. In addition, subclade A is seemingly confined to the Oceania—Asian distribution, shar- ing the characteristics of ellipsoid to allantoid, bean-shaped or reniform-like ascospores and a velvety to warty, ridged to corrugated external surface. According to the concept proposed by Korf (1957a), the species within the monophyletic subclade A are compatible with the morphology of Plectania sect. Curvatisporae. Here, it is worth mentioning that the revision of the type of U. mexicana (Ellis & Holw.) M. Carbone, Agnello, A.D. Parker & P. Alvarado revealed bean-shaped ascospores and one-typed external hairs, and it is so far confined to Mexico (Carbone et al. 2013c), showing morphological simi- larity to the species of subclade A. However, there is currently no sequence available for U. mexicana to confirm its exact phylogenetic position. Molec- ular data from specimens collected at the type locality are still needed. Based on the analysis of morphological characters combined with topo- logical structure, it can be hypothesized that two evolutionary directions of spore shape exist in Sarcosomataceae: from ellipsoid to spherical and from ellipsoid to allantoid. Ellipsoid spores are present in Donadinia, Plectania s. lat., and Sarcosoma. Conversely, as a sister clade to Sarcosoma, Pseudo- plectania displays globose to subglobose spores. Urnula exhibits a mixture of ellipsoid and allantoid spores, with all Urnula species bearing allantoid spores clustered in subclade A. This study also suggests a trend of spore evolution from smooth to ornamented surfaces. As shown by icons in our topology (Fig. 1), species of Pseudoplectania, Sarcosoma, and Urnula uni- formly possess smooth spores, whereas species of Donadinia are char- acterized by verrucose spores. Especially in Plectania s. lat., spore orna- mentation patterns are variable—smooth, finely to obviously verrucose or vermicular, and grooved. Notably, spore surface area is influenced by both shape and ornamentation. These findings indicate two divergent trends in surface area: (1) spores evolving from ellipsoid to allantoid and smooth to ornamented, corresponding with an increase in surface area variation; and (2) spores evolving from ellipsoid to spherical, associated with a decline in surface area variation. These trends may reflect different life strategies. A larger surface area may enhance dispersal and colonization potential (Ja- cobsen et al. 2017; Maurice et al. 2021), whereas a smaller surface area may strengthen water retention capacity (Kauserud et al. 2008; Fan et al. 2023). MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 57 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Moreover, species with sheath-covered spores appear to be confined to Plectania s. lat. and Pseudoplectania, according to our topology. The func- tion of the gelatinous sheath remains poorly understood, but it is probably associated with the protection of developing spores and the liberation of mature spores (Ruddick and Williams 1972; Williams et al. 1972). In conclusion, this study concentrates on updating the diversity and tax- onomy of Sarcosomataceae species in subtropical China. Combined with morphological examination and multi-locus phylogenetic analysis, four new species and two specimens with new distribution records were revealed and described, and keys for elucidating interspecific differences were pro- vided. Moreover, we encourage further systematic studies with the aim of clarifying phylogenetic relationships within Plectania s. lat. and providing new insights into reliable intergeneric criteria to Support a more natural classification and broader concepts within Sarcosomataceae. Acknowledgments We express our gratitude to Shi-En Wang (China) for providing valuable sugges- tions on the manuscript. Guan-Lin Chen (China) is thanked for his companion- ship during the field investigation. Jin-Rong Lu (China) is warmly acknowledged for guiding the examination of microscopic morphology. 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 supported by the following grants: the Fundamental Research Fund of Guangxi Institute of Botany (Grant No. GZY 23009), the Guangxi Natural Science Foun- dation (Grant No. 2025GXNSFBA069485), the Comprehensive Scientific Investigation of Huaping National Nature Reserve, and the Comprehensive Scientific Investigation of Maoershan National Nature Reserve. Author contributions Guang Rong Zhou, Zhou Rong Tan, JianHua Zhang: Specimen collection in subtropical China, field documentation. Jin Rong Liu, Si Ang Chen: Conceptualization, study design, specimen collection, morphological analyses, writing — original draft. Jin Rong Liu, Yan Cheng Zhang, Deng Li: Molecular laboratory work (DNA extraction, PCR amplification, sequencing), data curation, multi-locus dataset assembly. Guang Fu Mou: Phylogenetic analyses, data interpretation, taxonomic key preparation, visualization. Guang Fu Mou, Yan Liu (Corresponding Author): Supervision, funding acquisition, project administration, manuscript review & editing. All authors reviewed and approved the final manuscript. MycoKeys 121: 21-65 (2025), DOI: 10.3897/mycokeys.121.155432 58 Jin Rong Liu et al.: Multigene phylogeny of Sarcosomataceae reveals new taxa Author ORCIDs Jin Rong Liu © https://orcid.org/0000-0003-0843-2097 Deng Li © https://orcid.org/0009-0009-9357-3214 Si Ang Chen © https://orcid.org/0009-0007-7315-8107 Yan Cheng Zhang ® https://orcid.org/0009-0000-5278-1134 Guang Fu Mou @ hitps://orcid.org/0009-0007-231 8-7807 Yan Liu © https://orcid.org/0000-0001-8931-9656 Guang Rong Zhou ® https://orcid.org/0009-0001-6557-7092 Zhou Rong Tan ® hitps://orcid.org/0009-0000-1160-7041 Jian Hua Zhang ® https://orcid.org/0009-0007-971 7-4996 Data availability All of the data that support the findings of this study are available in the main text. 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