New and notable taxa of Ascomycota on the Qinghai-Xizang Plateau and its surrounding areas: Filamentous and entomopathogenic fungi

Introduction

Ascomycota is the largest and most species-rich group of fungi (Thiyagaraja et al. 2025), with approximately 111,903 known species worldwide (Catalogue of Life, August 16, 2026). Of these, around 16,474 species have been reported from China, accounting for about one-sixth of the global Ascomycota diversity (Bánki et al. 2025). Ascomycota play a pivotal role in global ecosystems, encompassing both microfungi and macrofungi, and contribute an estimated USD 54.57 trillion to the global economy (Niego et al. 2023a, b). These fungi are extensively distributed across terrestrial and aquatic environments and exhibit a diverse array of lifestyles, including saprotrophic, symbiotic, parasitic, and endophytic modes of existence (Liu et al. 2024a; Liao et al. 2025; Thiyagaraja et al. 2025). Notably, certain micro-ascomycetes are significant plant pathogens, as well as pathogens affecting animals and humans (Jayawardena et al. 2019). Notable examples include Aspergillus fumigatus, Botrytis cinerea, Candida albicans, Colletotrichum gloeosporioides, Fusarium oxysporum, Magnaporthe oryzae and Sclerotinia sclerotiorum (Gordon et al. 2017; Latgé & Chamilos 2019; Jayawardena et al. 2021).

The Qinghai-Xizang Plateau (abbreviated as Q-X Plateau) is bounded to the west by the Pamirs and to the north by the West Kunlun-Qilian-Altun Mountains, and gradually merges with the adjacent Qinling Mountains, Yunnan-Guizhou Plateau and lower mountains in southern Yunnan along the line of Minshan-Daxushan-Yulongshan (Wang et al. 2025b). In the Q-X Plateau, characterized by its challenging environmental conditions that restrict plant growth, fungi emerge as essential players in maintaining ecosystem balance. They contribute significantly by decomposing organic matter and recycling nutrients into the soil, thereby supporting overall ecological health (Bucher et al. 2004). Despite previous studies on fungi on the Q-X Plateau, including a number of published research papers (Wang & Zang 1983; Mao et al. 1993; Zang et al. 1996; He et al. 2024a, b, c, 2025; Xu et al. 2022, 2023, 2024a, b, c, d, e, 2025; Shen et al. 2025) and regional monographs, such as “Notes on the fungi from western Kunlun Mountains” (Zang & Xia 1989), “Economic Macrofungi of Tibet” (Mao et al. 1993), “Economic Macrofungi from Southwestern China” (Ying & Zang 1994), and different volumes of “Flora Fungorum Sinicorum” (Zang 1979, 1980, Yang 1997), the area is home to a vast variety of fungi, many of which require exploration. Therefore, we initiated the series of fungal diversity in the Q-X Plateau to publish fungal species systematically and promptly, and focused on the fungi in a special habitat, viz. macroscopic basidiomycetes in the first paper (Wang et al. 2025b), lignicolous freshwater fungi in the second paper (Xu et al. 2025). In the third paper of this series, we focus on entomopathogenic and filamentous fungi occurring in the pan Q-X Plateau.

Our investigation encompassed comprehensive fieldwork across the pan Q-X Plateau, resulting in the collection of over 14,000 specimens and samples representing diverse ascomycete taxa. By employing an integrative approach that synthesizes morphological assessments, phylogenetic analyses.

Materials and methods

Fungal specimens were collected from woody substrates, insect hosts, leaves and plant litter in terrestrial and freshwater habitats across Gansu, Guizhou, Qinghai, Sichuan, Xizang, Xinjiang and Yunnan, from 2019 to 2025. Morphological observations, macro- and microscopic photography, and isolation were conducted using the methods described by Wang et al. (2020a) and Xu et al. (2025), with necessary adaptations. Voucher specimens, ex-type cultures and living cultures were deposited in the following herbaria and culture collections. HKAS: Herbarium of Cryptogams, Kunming Institute of Botany, Chinese Academy of Sciences, China; KUMCC: Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China; KUNCC: Kunming Institute of Botany Culture Collection, China. DNA extraction was performed using both pure fungal cultures and fruiting bodies when cultures were not available. Procedures for DNA extraction, polymerase chain reaction (PCR), gel electrophoresis, and sequencing were conducted as outlined by Su et al. (2025). The sequence results generated in the study have been submitted to GenBank.

To delineate species boundaries, phylogenetic analyses were conducted using multiple methodologies. Maximum likelihood (ML) analyses were performed using RAxML-HPC2 v.8.2.12 (Silvestro & Michalak 2012; Stamatakis 2014) and the CIPRES Science Gateway portal (Miller et al. 2012), while Bayesian inference (BI) analyses were conducted using MrBayes v3.1.2 (Ronquist et al. 2012), and via the same web portal as in ML. Maximum parsimony (MP) analysis was executed using PAUP* version 4.0b10 (Swofford & Sullivan 2003). Pairwise homoplasy index (PHI) tests were conducted using Split Tree software, as described by Quaedvlieg et al. (2014), to assess the level of recombination among closely related species when necessary. Species delimitation was based on an integrative taxonomic approach combining morphology and multi-gene phylogenetic analyses following the recommendations of Jeewon & Hyde (2016), Chethana et al. (2021a) and Xu et al. (2025). Newly introduced fungal species were registered with Index Fungorum (Index Fungorum, June 30, 2025) and the Faces of Fungi database (Jayasiri et al. 2015).

Results

Checklist of species

This study presents a checklist of filamentous and entomopathogenic fungi from the pan Q-X Plateau, comprising 6 classes, 42 orders, 99 families, 228 genera, and 644 species, and encompassing representative taxa from various ecological niches (Table 1). These fungi are commonly found on substrates such as decaying wood, insect carcasses, leaf litter, soil, freshwater, plant tissues and other fungi, highlighting their broad ecological adaptability in the high-altitude environment. As previous records are somewhat scattered, the taxonomy and phylogenetic placement of certain species remain to be clarified. With the increasing application of high-throughput sequencing technologies, more undocumented fungal taxa are expected to be discovered, providing fundamental data for understanding fungal diversity on the Q-X Plateau.

Taxonomy

The classification followed here is from Hyde et al. (2024). The guidelines of Chethana et al. (2021a), and Maharachchikumbura et al. (2021) are followed when determining if species are novel or new collections.

Ascomycota Caval.-Sm

Dothideomycetes O.E. Erikss & Winka

The class Dothideomycetes, a significant group within the phylum Ascomycota, with approximately 32,621 species worldwide (Pem et al. 2024; Bánki et al. 2025; Catalogue of Life), encompasses a diverse array of fungi characterized by their unique morphological traits and ecological roles. This class is primarily defined by the presence of perithecial ascomata, which are typically dark and globose, facilitating the production of asci that contain ascospores (Hongsanan et al. 2020). In this study, a phylogenetic framework of Dothideomycetes is constructed using four loci (LSU, SSU, rpb2 and tef1-α) from approximately 600 strains. Further details are provided in Supplementary (Fig. 1).

Fig. 1. Maximum likelihood majority rule consensus tree for Acrogenospora using ITS, LSU, SSU, rpb2 and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Minutisphaera aspera (DSM 29478) and M. japonica (HHUF 30098). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Minutisphaerales Raja, Oberlies, Shearer & A.N. Mill.

Acrogenosporaceae Jayasiri & K.D. Hyde

Acrogenospora M.B. Ellis

Acrogenospora was established by Ellis (1971). It is characterized by macronematous, mononematous, and brown conidiophores that frequently proliferate percurrently. The conidiogenous cells are monoblastic, either terminal or intercalary, and its conidia are globose, ellipsoid, or obovoid, ranging from olivaceous to brown (Hughes 1978; Goh et al. 1998c; Bao et al. 2020; Harrington et al. 2022; Li et al. 2024b). Initially, Acrogenospora comprised two species, A. carmichaeliana and A. sphaerocephala, both described solely by their asexual morphs. Goh et al. (1998c) expanded the genus by adding A. ovalia and A. subprolata, while Bao et al. (2020) further refined its taxonomy by accommodating nine species, including seven new taxa and two previously described ones, based on phylogenetic and morphological analyses. Currently, Acrogenospora encompasses 27 species, primarily from freshwater habitats (21 species), and terrestrial habitats (six species) (Bao et al. 2020; Li et al. 2024b; Cao et al. 2025; Xu et al. 2025).

1. Acrogenospora carmichaeliana (Berk.) Rossman & Crous, in Rossman, Crous & Hyde, IMA Fungus 6(2): 509 (2015) Fig. 2

Chinese name

Index Fungorum number: IF 814513; Facesoffungi number: FoF 08065

Saprobic on decaying wood submerged in freshwater stream. Sexual morph: Undetermined. Asexual morph: Colonies effuse on natural substrate, hairy, dark brown, with masses of conidia on conidiophores. Mycelium mostly immersed, composed of grayish-brown, septate, branched, smooth hyphae. Conidiophores 267–354 × 8–10 µm ( =308 × 8 μm, n = 10), mononematous, macronematous, solitary, erect, straight or slightly flexuous, cylindrical, unbranched, brown to dark brown, paler toward apex, septate, smooth-walled. Conidiogenous cells holoblastic, monoblastic, integrated, initially terminal, later becoming intercalary, cylindrical, smooth-walled, pale brown, proliferating percurrently. Conidia 31–38 × 25–31 µm ( = 35 × 28 μm, n = 25), acrogenous or acropleurogenous, solitary, spherical or subspherical, truncate at base, aseptate, hyaline when young, dark brown to black when mature, smooth-walled.

Material examined: CHINA, Xinjiang Autonomous Region, Karamay City, Lengshuigou, 43°58′11.99" N, 85°6′47.30" E, 1785 m a.s.l., on decaying wood submerged in a freshwater stream, 16 July 2021, R.J. Xu, MD-369 (HKAS 136217), living culture KUNCC 24-18045.

Notes: The taxonomic status of Acrogenospora carmichaeliana and its relationship with Farlowiella carmichaeliana have long been complex issues in mycological research. Historically, based on culture studies, Acrogenospora megalospora was considered as the asexual morph of Farlowiella carmichaeliana. Ellis (1971a) reported the asexual morph of F. carmichaeliana as Acrogenospora carmichaeliana, and Goh et al. (1998c) confirmed this relationship and synonymized A. carmichaeliana under Farlowiella carmichaeliana. The asexual-sexual connection between these two genera (Farlowiella and Acrogenospora) was also confirmed by Jayasiri et al. (2018) and Hyde et al. (2019) based on phylogenetic analysis. Therefore, Acrogenospora was given priority over Farlowiella. However, Jayasiri et al. (2018) found that Acrogenospora sphaerocephala clustered with Farlowiella carmichaeliana (the sexual morph of Acrogenospora carmichaeliana), supporting their connection. However, recent studies, Hyde et al. (2019) and Bao et al. (2020), showed that Acrogenospora sphaerocephala forms different clades from Acrogenospora carmichaeliana. Thus, whether A. sphaerocephala was the asexual morph of Farlowiella carmichaeliana and whether Acrogenospora megalospora was wrongly introduced as the asexual morph of Farlowiella carmichaeliana needs further study. In this study, our collection KUNCC 24-18045 is identified as Acrogenospora carmichaeliana based on morphological traits and molecular evidence (Fig. 1).

Pleosporales Luttrell ex M.E. Barr

Corynesporascaceae Sivan.

Corynespora Güssow, J

Corynesporasca, established by Sivanesan (1996) and typified by C. carotae, represents the sexual morph of a Corynespora species found on tropical plants. Corynespora typified by C. mazei and now synonymised with C. cassiicola, comprises about 200 species. The leaf spot fungus Corynespora cassiicola is particularly widespread, infecting numerous plant hosts predominantly in tropical regions (Smith et al. 2007). Due to the extensive usage and historical priority of the name Corynespora, especially in the context of plant pathogenic fungi, it is recommended to continue using this name. Corynespora species are mainly tropical and subtropical, found on a wide range of plant hosts, and known for causing leaf spot diseases. They exhibit notable morphological features such as distinct, determinate or percurrently extending conidiophores and monotretic, integrated, terminal conidiogenous cells producing solitary or sometimes catenate, distoseptate conidia. Liu et al. (2023a) and Zhang et al. (2025) provided a detailed description and explanation of the genus.

2. Corynespora yunnanensis Jing W. Liu & Jian Ma, in Liu, Hu, Luo, Castañeda-Ruíz, Xia, Xu, Cui, Shi, Zhang & Ma, Journal of Fungi 9 (no. 107): 11 (2023) Fig. 4

Chinese name: –

Index Fungorum number: IF 900078; Facesoffungi number: FoF 17390

Saprobic on decaying stems of wood submerged in a freshwater stream habitat. Sexual morph: Undetermined. Asexual morph: Colonies superficial, effuse, velvety, gregarious, pale brown, Mycelium immersed, subhyaline to pale brown, composed of branched, septate hyphae. Conidiophores 406–603 × 12–19 µm ( = 490 × 16 μm, n = 20), macronematous, mononematous, erect, straight or slightly flexuous, solitary or group, tapering towards the apex, septate, unbranched, cylindrical, dark brown or brown, becoming paler towards the apex, with several successive cylindrical extensions. Conidiogenous cells 21–38 × 10–16 µm ( = 29 × 12 μm, n = 20), integrated, terminal, monotretic, cylindrical, pale brown to brown, smooth. Conidia 122–154 × 20–28 µm ( = 134 × 23 μm, n = 25), acrogenous, solitary, obclavate, rostrate, rounded at the apex, straight or slightly curved, multi-distoseptate, olivaceous to brown, smooth-walled, truncate at the base.

Culture characteristics: Conidia germinating on PDA medium within 24 h, and germ tubes produced from both ends. Colonies grown on PDA reached 60 mm diameter at room temperature in two weeks, circular, with velvety, flat, brown mycelium on the surface, from below dark brown with entire margin.

Material examined: CHINA, Xizang Autonomous Region, Shannan Prefecture, Cuona City, Lebugou, 27°52′38.19" N, 91°47′54.64" E, 3560 m a.s.l., on decaying wood submerged in a freshwater stream, 3 August 2023, R.J. Xu, LJN-45 (HKAS 135995), living culture KUNCC 24-17961.

Notes: Corynespora yunnanensis was isolated from dead branches of an unidentified tree in Yunnan Province, China (Liu et al. 2023a). Morphologically, our collection (KUNCC 24-17961) shares identical morphological characters to the holotype of C. yunnanensis but has larger conidia (122–154 × 20–28 vs. 80–128 × 16–19 µm). The phylogenetic position of our strain is unclear (Fig. 3). However, comparisons of ITS sequences demonstrate 100% similarity between the ex-type strain HJAUP M2132 and KUNCC 24-17961. Therefore, based on the current evidence, we tentatively retain our strain as C. yunnanensis. Further collections and additional evidence are needed to clarify their relationship.

Fig. 2. Acrogenospora carmichaeliana (HKAS 136217) a, b Colony on natural substrates. c–f Conidiophores with conidia. g, h Conidia. Scale bars: c–e = 100 μm, f–h = 10 μm.

Fig. 3. Maximum likelihood majority rule consensus tree for Corynespora using ITS and LSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML BS/PP. The scale bar represents the expected number of changes per site. The tree is rooted with Periconia byssoides (H 4600), P. digitata (CBS 510.77) and P. pseudodigitata (KT 1395). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

3. Corynespora xinjiangensis R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 5

Chinese name: –

Index Fungorum number: IF 905026; Facesoffungi number: FoF 19296

Etymology: Refers to Xinjiang Autonomous Region in China, where the type specimen of this species was collected.

Holotype: HKAS 151641

Saprobic on decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum, superficial, effuse, filiform, pale brown. Mycelium mostly immersed, composed of branched, septate, pale brown to brown, smooth-walled hyphae. Conidiophores macronematous, mononematous, fasciculate, filamentous, cylindrical, septate, unbranched, pale brown to brown, sometimes with multiple consecutive proliferations. Conidiogenous cells monotretic, integrated, terminal, cylindrical, pale brown to brown, smooth-walled, often percurrently proliferating. Conidia 63–114 × 6–35 µm ( = 84 × 12 μm, n = 30), acrogenous, solitary, obclavato-cylindrical, straight to slightly flexuous, light olivaceous-brown, truncate at the base, obtuse to rounded at the apex, distoseptate, sometimes proliferates 1 to 3 times at the tip of the conidia.

Culture characteristics: Conidia germinating on PDA medium within 24 hours, with germ tubes arising from both ends of the conidium. Colonies grown on PDA reaching 60 mm diam at room temperature under natural light after two weeks, circular, flat, with velvety, with filiform edge, brown to dark brown from above and below.

Material examined: CHINA, Xinjiang Autonomous Region, Kashi Prefecture, Yecheng City, 37° 45' 8.87" N, 77° 25' 7.71" E, 1490 m a.s.l., on decaying wood submerged in a freshwater habitat, 18 July 2021, R.J. Xu, MD-172A (HKAS 151641, holotype), ex-type culture KUNCC 25-21366.

Fig. 4. Corynespora yunnanensis (HKAS 135995) a Colony on natural substrates. b, c Conidiophores and conidiogenous cells. d–f Conidiogenous cells with young conidia. g–k Conidia. l Germinated conidium. m, n Culture on PDA medium, m from above, n from below. Scale bars: b–c = 100 μm, d–l = 50 μm.

Fig. 5. Corynespora xinjiangensis (HKAS 151641, holotype) a, b Colony on natural substrates. c, d Short conidiophores and conidia. e Conidiogenous cells with young conidia. f–h Conidia. i Germinated conidium. j, k Culture on PDA medium, j from above, k from below. Scale bars: c, d = 100 μm, e–i = 50 μm.

Notes: Corynespora xinjiangensis is morphologically typical of Corynespora, characterized by macronematous, mononematous conidiophores, monotretic conidiogenous cells, and obclavato-cylindrical, distoseptate conidia (Smith et al. 2007). The conidia of C. xinjiangensis are relatively large, measuring 63–114 × 6–35 µm, with an average size of 84 × 12 µm, with 1–3 proliferations at the apex. Corynespora xinjiangensis differs from C. mengsongensis and C. nabanheensis, by having short, multiple consecutive proliferations conidiophores and obclavato-cylindrical conidia (Liu et al. 2023a). Furthermore, a comparison of the ITS nucleotide bases in C. xinjiangensis (KUNCC 25-21366) and C. smithii (MFLU 19-0688) revealed that they differ in 83/869 bp (9.55%) of ITS, and the phylogenetic analyses (Fig. 3) place C. xinjiangensis in a distinct lineage, supporting its separation from the known species.

Dictyosporiaceae Boonmee & K.D. Hyde

Dictyocheirospora M.J. D'souza, S. Boonmee & K.D. Hyde

Dictyocheirospora was established by Boonmee et al. (2016) with D. rotunda as the type species. Dictyocheirospora is distinguished from its close relative genus Dictyosporium, by its unique conidial morphology. The conidia in Dictyocheirospora are cheiroid, non-complanate, or cylindrical, often tightly grouped at the apex, contrasting with the conidial arrangement in Dictyosporium (Boonmee et al. 2016; Wang et al. 2016a; Shen et al. 2022). The asexual morphs of Dictyocheirospora are characterized by dark sporodochial colonies producing aeroaquatic conidia, while its sexual morphs are noted for subglobose ascomata, cylindrical asci, and hyaline, fusiform, uniseptate ascospores, with or without a sheath (Boonmee et al. 2016; Tibpromma et al. 2018; Liu et al. 2023b; Tennakoon et al. 2023)

The phylogenetic placement of Dictyocheirospora within the Dictyosporiaceae family has been validated through analyses employing multiple-gene phylogenies in various studies (Liu et al. 2015a; Tanaka et al. 2015; Boonmee et al. 2016). Dictyocheirospora currently comprises 33 accepted species, identified through both morphological characteristics and phylogenetic analysis (Boonmee et al. 2016; Wang et al. 2016a; Hyde et al. 2017; Hyde et al. 2019; Yang et al. 2018a; Tibpromma et al. 2018; Jayasiri et al. 2019; Phookamsak et al. 2019; Dong et al. 2020; Phukhamsakda et al. 2020; Shen et al. 2022; Tennakoon et al. 2023; Wang et al. 2025c; Sun et al. 2025).

4. Dictyocheirospora luojiensis R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 7

Chinese name: –

Index Fungorum number: IF 903471; Facesoffungi number: FoF 17391

Etymology: Referring to the location “Luoji River” where the holotype of this fungus was collected.

Holotype: HKAS 136148

Saprobic on decaying stems of wood submerged in a freshwater river. Sexual morph: Undetermined. Asexual morph: Colonies sporodochial, punctiform, scattered or in small groups, compactly, water-stained, dark brown to black. Mycelium immersed, subhyaline to pale brown, composed of branched, septate hyphae. Conidiophores macronematous, mononematous, septate, cylindrical, pale brown to subhyaline, smooth-walled, sometimes reduced to conidiogenous cells. Conidiogenous cells holoblastic, cylindrical to subglobose, hyaline to pale brown, smooth-walled. Conidia 56–71 × 22–40 µm ( = 64 × 28 μm, n = 25), acrogenous, solitary, cheiroid, brown or dark brown, closely appressed, composed of seven rows of cells, non-complanate, cylindrical, each row with 8–12 cells. Conidial secession was schizolytic.

Culture characteristics: Conidia germinating on PDA medium within 72 h and germ tubes arising from terminal end of conidium. Colonies grown on PDA reached 30 mm diam at room temperature in one month, circular, with velvety, raised, white mycelium on the surface, from below cream to light brown with entire margin.

Material examined: CHINA, Yunnan Province, Shangri-La City, Luoji River, 27°42′23.30" N, 99°59′31.51" E, 3539 m a.s.l., on decaying wood submerged in a freshwater river, 18 June 2022, R.J. Xu, LTS-25 (HKAS 136148, holotype), ex-type culture KUNCC 24-17999.

Notes: Phylogenetic analysis combining SSU, ITS, LSU and tef1-α sequence data showed that Dictyocheirospora luojiensis clustered within the Dictyocheirospora clade and formed an independent lineage (Fig. 6). Dictyocheirospora luojiensis is closely related to D. lithocarpi, D. multiappendiculata, and D. suae (Fig. 6). Dictyocheirospora luojiensis is morphologically similar to these species in having cheiroid, ellipsoid to cylindrical, not complanate, septate, closely appressed conidia, and with or without appendages (Jayasiri et al. 2019; Shen et al. 2022). However, D. luojiensis can be distinguished from D. lithocarpi by its larger conidia (56–71 × 22–40 µm vs. 35–40 × 12–18 μm) with appendages, and with more rows of cells (7 rows vs. 6 rows) (Jayasiri et al. 2019). Dictyocheirospora luojiensis differs from D. multiappendiculata by having larger conidia (56–71 × 22–40 µm vs. 55–62 × 19–22 µm) and rare appendages, whereas appendages are common in D. multiappendiculata (Shen et al. 2022). Dictyocheirospora luojiensis differs from D. suae by its smaller conidia (56–71 × 22–40 µm vs. 72–79 × 20–25 µm), and fewer cells in each row of cells (8–12 cells vs. 12–15 cells) (Shen et al. 2022). Conidial size is one of the important criteria for distinguishing dictyosporium-like taxa (Goh et al. 1999). Based on the unique morphological characters and phylogenetic distance, we therefore introduce D. luojiensis as a new species here.

Massarinaceae Munk

Helminthosporium Link

Helminthosporium was introduced by Link (1809) with H. velutinum as the type species. Species of Helminthosporium are widely distributed and mainly saprobic on a broad range of natural substrates such as decaying wood, plant stems, bark, dung, and occasionally insects, and some species have been reported as opportunistic human pathogens (Luttrell 1964; Alcorn 1988; Konta et al. 2023; Hyde et al. 2023, 2024). The genus is classified in Massarinaceae (Pleosporales), and is characterized by macronematous, cylindrical, septate, erect conidiophores with polytretic conidiogenous cells and acropleurogenous, clavate or obclavate, distoseptate conidia with a flat, ringed pore at the base (Luttrell 1964; Liu et al. 2022; Hyde et al. 2023). Although over 780 names have been listed under Helminthosporium (Index Fungorum 2026), many were reclassified based on morphological inconsistencies with the characteristics of the genus. In addition, recent phylogenetic analyses based on molecular data indicate that Helminthosporium is polyphyletic (Konta et al. 2021; Chen et al. 2022). In this study, we introduce a new species of Helminthosporium based on morphological characteristics and phylogenetic analysis.

5. Helminthosporium xinjiangense R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 9

Chinese name

Index Fungorum number: IF 905028; Facesoffungi number: FoF 19297

Etymology: Refers to Xinjiang Autonomous Region in China, where the type specimen of this species was collected.

Holotype: HKAS 151642

Saprobic on decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum, superficial, effuse, scattered, hairy, dark brown. Mycelium mostly immersed, composed of branched, septate, brown to dark brown, smooth-walled hyphae. Conidiophores 214–433 × 9–12 μm ( = 329 × 11 μm, n = 20), macronematous, mononematous, solitary or in groups, erect, cylindrical, straight or flexuous, unbranched, dark brown to black, smooth-walled. Conidiogenous cells 47–112 × 5–9 μm ( = 75 × 8 μm, n = 20), polytretic, integrated, terminal and intercalary, cylindrical, brown, smooth-walled. Conidia 55–89 × 8–11 μm ( = 66 × 10 μm, n = 20), acropleurogenous, solitary, obclavate-rostrate, straight or curved, usually curved, tapering towards the rounded apex, brown, subhyaline at the apex, 6–9-distoseptate, smooth-walled, with a conspicuous black scar at the base.

Culture characters: Conidium germinating on PDA within 24 h and germ tubes produced from both ends or middle. Colonies on PDA reaching 10 mm diam., in a week at room temperature, effuse, hairy, mycelium radiating outwards, fimbriate edge, dense, pale. Mycelium superficial and partly immersed, light brown.

Fig. 6. Maximum likelihood majority rule consensus tree for Dictyocheirospora using SSU, ITS, LSU and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Periconia igniaria (CBS: 379.86 and CBS: 845.96). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 7. Dictyocheirospora luojiensis (HKAS 136148, holotype) a, b Colony on natural substrates. c Conidiophores and conidia. d, e Conidiophores and conidiogenous cells. f–j Conidia. k Germinated conidium. Scale bars: c = 50 μm, d–k = 20 μm.

Material examined: CHINA, Xinjiang Autonomous Region, Kashi, Yecheng City, 37° 45' 8.87" N, 77° 25' 7.71" E, 1490 m a.s.l., on decaying wood submerged in a freshwater habitat, 18 July 2021, R.J. Xu, MD-176A (HKAS 151642, holotype), ex-type culture KUNCC 25-21367.

Notes: In phylogenetic analyses, our isolate formed a sister clade with Helminthosporium austriacum and H. yunnanense (Fig. 8). Sequence comparison of the ITS region

Fig. 8. Maximum likelihood majority rule consensus tree for Helminthosporium using SSU, LSU, ITS, rpb2 and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML BS/PP. The scale bar represents the expected number of changes per site. The tree is rooted with Periconia pseudodigitata (CBS 139699). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

showed nucleotide differences of 13.64% (70/513 bp, excluding gaps) and 15.20% (78/513 bp, excluding gaps) between H. xinjiangense and H. austriacum, H. xinjiangense and H. yunnanense, respectively. Morphologically, H. xinjiangense differs from H. austriacum by having shorter conidiophores (214–433 × 9–12 μm vs. 275–700 × 11.5–19 μm), and more slender, usually curved conidia (55–89 × 8–11 μm vs. 35–48 × 13.7–16.5 μm) (Voglmayr et al. 2017), and differs from H. yunnanense in shorter conidiophores (214–433 × 9–12 μm vs. 560–680 × 12.5–15.5 μm) without one cylindrical, enteroblastic percurrent extension, and larger, obclavate-rostrate conidia (55–89 × 8–11 μm vs. 30.5–55.5 × 9–11 μm) with a higher number of distosepta (6–9 vs. 4–7) (Liu et al. 2022). Therefore, we introduce H. xinjiangense as a new species.

Hermatomycetaceae Locq. ex A. Hashim. & Kaz. Tanaka

Hermatomycetaceae was informally introduced by Locquin (1984) and later validated by Hashimoto et al. (2017) to accommodate the monotypic genus Hermatomyces. Members of Hermatomycetaceae are saprobic on various plant substrates. The sexual morph has not yet been observed. Asexual structures include sporodochial, pulvinate conidiomata that are dark brown to black. Conidiophores are pale brown and mononematous, bearing monoblastic, integrated, terminal conidiogenous cells that are cylindrical (Castañeda & Heredia 2000; Ren et al. 2021).

Hermatomyces Speg

Hermatomyces has a wide geographic range, occurring in both tropical and temperate regions, including Brazil, China, Cuba, Japan, Panama, the Philippines, Thailand, and Venezuela (Doilom et al. 2017; Hashimoto et al. 2017b; Koukol et al. 2018; Tibpromma et al. 2018). To date, host specificity within Hermatomyces remains unclear, as species have been identified on hosts from multiple plant families, including Acanthaceae, Arecaceae, Asteraceae, Fabaceae, and Pandanaceae (Doilom et al. 2017; Hashimoto et al. 2017; Koukol et al. 2018; Tibpromma et al. 2018). The sexual-asexual connection in Hermatomyces has not yet been established, since no sexual morph has been documented for any species in the genus (Hashimoto et al. 2017b; Tibpromma et al. 2018). Up to now, 34 species have been accepted within Hermatomyces (Hyde et al. 2024a; Shen et al. 2024; Koukol et al. 2025).

6. Hermatomyces motuoensis R.J. Xu & Q. Zhao, sp. nov. Fig. 11

Chinese name

Index Fungorum number: IF 905256; Facesoffungi number: FoF 19298

Etymology: The epithet refers to Motuo County, located in Xizang of China, where the type specimen was collected.

Holotype: HKAS 151643

Saprobic on decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate effuse, superficial, scattered, conspicuous, flattened, consisting of a sterile mycelial outer zone and a round, glistening, brown to black. Mycelium mostly immersed, composed of branched, septate, brown to dark brown, smooth-walled hyphae. Conidiophores micronematous, mononematous, unbranched, straight or flexuous, hyaline, septate, smooth. Conidiogenous cells 11–20 × 2–7 μm ( = 16 × 4 μm, n = 20), holoblastic, monoblastic, integrated, terminal, cylindrical, hyaline to subhyaline. Conidia 19–33 × 17–27 μm ( = 28 × 22 μm, n = 30), dimorphic, thick-walled, smooth. lenticular conidia, subglobose to ellipsoidal, muriform, smooth, central cells dark brown to black, peripheral cells narrow, flattened, subhyaline to pale brown, obovoid or oblong in side view, cylindrical conidia, 2–3-celled, symmetrical or asymmetrical, distinctly constricted at the septum, guttulate, straight or flexuous, hyaline, the upper cells usually subcylindrical and the apex rounded or slightly flattened, often slightly attenuated and dark brown or black, the lower cells are ellipsoidal.

Cultural characteristics: Conidia germinating on PDA and germ tubes produced from peripheral cells of conidium within 24 h. Colonies on PDA, reaching 30 mm after one month at room temperature, colony circular, edge smooth, surface hyphae grey-brown, fluffy, flocculent, reverse side grey-brown to brown, dense, cracked in the middle.

Fig. 9. Helminthosporium xinjiangense (HKAS 151642, holotype) a–c Colony on natural substrates. d, e Conidiophores and conidiogenous cells. f, g Conidiogenous cells with conidia. h–j Conidia. k Germinated conidium. Scale bars: d = 50 μm, e–k = 20 μm.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, Beibong Township, 29° 14′ 35.76″ N, 95° 10′ 6.94″ E, 1278 m a.s.l., on decaying wood submerged in a freshwater stream, 12 July 2022, R.J. Xu, MD-200A (HKAS 151643, holotype), ex-type culture KUNCC 25-21368.

Notes: Phylogenetic analysis revealed that Hermatomyces motuoensis clustered with H. hongheensis, H. constrictus, H. iriomotensis, H. krabiensis and H. griseomarginatus with 84% ML/0.68 BYPP support (Fig. 10). Morphologically, H. motuoensis displays dimorphic conidia, characterized by both lenticular and cylindrical forms. This morphological trait aligns with the conidial characteristics observed in H. constrictus, H. iriomotensis, and H. krabiensis, thereby indicating a potential phylogenetic relationship among these taxa (Spegazzini 1910; Tibpromma et al. 2016; Hashimoto et al. 2017; Koukol et al. 2018; Ren et al. 2021). However, H. motuoensis can be distinguished from H. constrictus in having larger conidiogenous cells (11–20 μm vs. 4–7 μm), and the cylindrical conidia with more cells (2–3-cells vs. 2-cells) (Koukol et al. 2018). Compared to H. iriomotensis, H. motuoensis has smaller lenticular conidia (28 × 22 μm avg. vs. 33.6 × 23.5 μm, avg.) and fewer septa (1–2-septate vs. 3–7-septate) in cylindrical conidia (Tibpromma et al. 2016). Furthermore, a megablast search of NCBI's GenBank nucleotide database, the best matching result of the ITS sequence is H. bifurcatus (voucher: PMA:116075, sequence ID: LS398263, similarities: 616/656 (94%), 10 gaps. Therefore, based on morphological and molecular sequence evidence, H. motuoensis is herein introduced as a novel species.

Fig. 10. Maximum likelihood majority rule consensus tree for Hermatomyces using ITS, LSU, SSU, tef1-α and rpb2 sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Aquasubmersa japonica (MAFF 245218 and MAFF 245219). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 11. Helminthosporium motuoensis (HKAS 151643, holotype) a, b Colonies on natural substrate. c, d Conidiophores, conidiogenous cells and conidia. e–h Conidia. i Germinated conidium. j, k Culture on PDA medium, j from above, k from below. Scale bars: c, d, i = 50 μm, e–h = 20 μm.

Periconiaceae Nann

Periconia Tode

Periconiaceae was delineated by Nannizzi (1934) and represented by the genus Periconia. The family was resurrected by Tanaka et al. (2015) and placed in Pleosporales based on morphological and phylogenetic evidence. Currently, the family comprises a single genus Periconia. Subsequent taxonomic reassessments by Yang et al. (2022a) led to the synonymization of Bambusistroma and Noosia under Periconia, based on morphological traits observed in sexual reproductive structures and multilocus phylogenetic studies. Additionally, Liao et al. (2024a) suggest that Flavomyces fulophazii should be synonymized with Periconia. However, further corroborative data is essential to affirm the taxonomic placement of F. fulophazii.

Periconia was introduced by Tode (1790) with P. lichenoides as the type species. Periconia was re-classified by Tanaka et al. (2015) based on the multi-locus phylogenetic analysis and placed in Periconiaceae. Species of Periconia are mostly reported as saprobes as their asexual morph, and also have some species as endophytes and pathogens, such as P. endophytica was isolated as an endophytic fungus from asymptomatic leaves of Wurfbainia villosa in China (Liao et al. 2024a). Periconia circinata was isolated as a pathogen causing blackening and rotting of wheat roots and stem bases (Goga 2000). To date, Periconia has 201 epithets listed in Species Fungorum.

7. Periconia xizangensis R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 13

Chinese name

Index Fungorum number: IF 905030; Facesoffungi number: FoF 17397

Etymology: The epithet refers to Xizang, China, where the type specimen was collected.

Holotype: HKAS 136110

Saprobic on decaying wood submerged in a freshwater stream. Sexual morph: Undetermined. Asexual morph: Colonies effuse, hairy, velvety, scattered or in small groups, with glistening, pale brown to brown conidial masses at the apex. Mycelium is mostly superficial, partially immersed, consisting of branched, septate, smooth, subhyaline hyphae. Conidiophores 366–601 × 13–19 μm ( = 467 × 16 μm, n = 20), macronematous, mononematous, solitary, erect, unbranched or occasionally 2–3-branched, septate, straight or flexuous, cylindrical, tapering towards at the apex, with cylindrical to elliptical heads at the apex, often swollen at the base, dark brown at the base, light brown towards the apex. Conidiogenous cells polyblastic, pale brown to brown, terminal, subglobose or elliptical, smooth-walled. Conidia 6–8.5 × 5.6–8 μm ( = 7.3 × 7.3 μm, n = 40), solitary or aggregated in masses, globose, brown, aseptate, verruculose.

Culture characteristics: Conidia germinating on PDA medium within 24 h and germ tubes produced from both ends. Colonies grown on PDA reached 60 mm diam at room temperature in two weeks, filamentous, with velvety, flat, dark brown mycelium, aerial mycelium on the surface, from below dark brown with undulate margin.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°19′43.69" N, 95°21′19.27" E, 798 m a.s.l., on decaying wood submerged in a freshwater stream, 12 July 2022, R.J. Xu, XK-14 (HKAS 136110, holotype), ex-type culture KUNCC 10433.

Notes: Phylogenetic analyses showed that our collection (KUNCC 10433) formed a distinct clade within Periconia, and formed a sister relationship with the ex-type strain of P. thailandica (MFLUCC 17-0065) (Fig. 12). Periconia xizangensis differs from P. thailandica in having conidiophores sometimes with 2–3-branched and cylindrical to elliptical heads (Liu et al. 2017).

Phaeosphaeriaceae M.E. Barr

Phaeosphaeriaceae, typified by Phaeosphaeria, currently comprises 84 genera and remains one of the largest families within Pleosporales (Wanasinghe & Maharachchikumbura 2023). Species in this family exhibit high diversity and are widely distributed in terrestrial environments worldwide, functioning as plant pathogens, saprobes, endophytes, and lichenicolous fungi (Phookamsak et al. 2014). Many species are significant plant pathogens that affect economically important crops, such as Phaeosphaeria and Paraconiothyrium which cause leaf spots on wheat and rice (Phookamsak et al. 2014). Although ongoing research integrating morphological and phylogenetic methods has led to considerable progress in identifying these taxa, the classification is still not fully resolved.

Setophoma Gruyter, Aveskamp & Verkley

Setophoma was originally published to accommodate Phoma terrestris and Pyrenochaeta sacchari (Liu et al. 2019a). Currently, 19 species are listed in Index Fungorum. They were thought to be well-resolved, since many of them are available of molecular data and colour illustrate (Phookamsak et al. 2014). Species in this genus are commonly found in asexual morph, living as plant pathogens (Crous et al. 2014, 2016). Morphologically, they are characterised by phialidic conidiogenous cells, and hyaline, ellipsoidal to subcylindrical, aseptate conidia (Liu et al. 2019a).

8. Setophoma manglietiae H.D. Yang, K.D. Hyde & Q. Zhao, sp. nov. Fig. 15

Chinese name: – 木莲壳霉 (mu lian ke mei)

Index Fungorum number: IF905257; Facesoffungi number: FoF 19299

Etymology: Referring to the host plant Manglietia fordiana (Magnoliaceae).

Holotype: HKAS 154724

Leaf spots amphigenous, angular, brown to grey. Sexual morph: Undetermined. Asexual morph: Colony on PDA grows slowly, reaching 66 mm after 65 days. Upper surface, flat, cottony, white at the beginning, forming pink patches near the centre at the later, undulate to entire edge, lower surface brown with pink plaque, white at the edge. Colony on SNA, upper white, flat, with abundant aerial mycelium, undulate to entire edge, lower surface white. Colony on WA, mycelium sparse, growth zone appears grey. The hyphae measure 1.8–3.3 μm in diameter, are septate, hyaline to lightly pigmented brown, and contain abundant guttules commonly aggregate to form inflated, globose, or bead-like structures.

Material examined: CHINA, Yunnan Province, Kunming City, on leaf of Manglietia fordiana, 14 March 2022, Hong-De Yang14 (HKAS 154724, holotype), ex-type culture KUNCC 23-14872.

Fig. 12. Maximum likelihood majority rule consensus tree for Periconia using ITS, LSU, SSU and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Morosphaeria ramunculicola (KH 220) and M. velatispora (KH 221). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Notes: Setophoma manglietiae exhibits a phylogenetic affinity with S. chromolaenae, S. vernoniae, and S. syzygii (Fig. 14), all of which have been isolated from foliar tissues and associated with leaf pathologies. Our isolate presents shared phenotypic traits, such as the formation of colony patches and an undulate to entire margin; however, it is distinguished by its pigmentation, which varies from white to grey and pink (Quaedvlieg et al. 2013; Crous et al. 2014, 2021). Due to an inability to induce sporulation on various media, including PDA, SNA, and WA, micromorphological characteristics could not be assessed. Consequently, identification was primarily based on colony morphology and molecular data. It is noteworthy that S. chromolaenae, S. vernoniae and S. syzygii are characterized by the production of hyaline, aseptate, subcylindrical conidia (Quaedvlieg et al. 2013; Crous et al. 2014, 2021). Thus, further investigations are warranted to confirm the morphological attributes of our species.

Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray.

Tetraploa Berk. & Broome

Tetraploa (synonym Tetraplosphaeria) was described by Berkeley & Broome in 1850 to accommodate the type species T. aristata. Species of the genus Tetraploa are predominantly isolated from bamboos, various herbaceous plants, and decaying wood across both terrestrial and aquatic ecosystems (Ando 1992; Dong et al. 2020; Hyde et al. 2020b; Phookamsak et al. 2022; Li et al. 2023; Liao et al. 2024b). This genus exhibits both sexual and asexual reproductive strategies. The asexual morph is distinguished by the absence of conidiophores, featuring monoblastic conidiogenous cells, and producing verrucose, short cylindrical to obpyriform conidia arranged in several columns, each adorned with setose appendages at the apex (Tanaka et al. 2009; Hyde et al. 2013). In contrast, the sexual morph is characterized by the presence of immersed to superficial, globose to subglobose ascomata, short-pedicellate asci, and septate, hyaline ascospores that are narrowly fusiform to broadly cylindrical, often enveloped by a mucilaginous sheath-like appendage (Tanaka et al. 2009; Hyde et al. 2013; He et al. 2025). There are about 30 species epithets linked to the genus Tetraploa (Hyde et al. 2024a).

9. Tetraploa hainanensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, in Tang, Jeewon, Lu, Alrefaei, Jayawardena, Xu, Ma, Chen & Kang, MycoKeys 100: 189 (2023) Fig. 17

Chinese name

Index Fungorum number: IF 900952; Facesoffungi number: FoF 14667

Saprobic on the dead stem of Zea mays. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies effuse, brown to dark brown, hairy, gregarious on the host substrate. Mycelium partly superficial, partly immersed, septate, branched and pale brown. Conidiophores absent. Conidiogenous cells integrated, monoblastic, determinate. Conidia 29–44 × 17–31 µm ( = 36 × 25 µm, n = 20), cylindrical with obtuse ends, dark brown, solitary, verrucose, smooth, composed of four columns of cells, sometimes five columns of cells, 2–6-septate in each column, with four apical appendages. Appendages 59–193 × 4–6 μm ( = 131 × 5 μm, n = 30), cylindrical, solitary, straight or slightly flexuous, smooth-walled, unbranched, pale brown to brown, guttulate, 4–8-septate in each column, wide at the base, tapering toward the apex with hyaline.

Material examined: CHINA, Yunnan Province, on a dead stem of Zea mays, 9 September 2022, Yanyan Yang, YYY333 (HKAS 144363), culture KUNCC 25-20221. Yunnan Province, Xishuangbanna, Mengyang Town, on a dead stem of Zea mays, 6 September 2022, Yanyan Yang, YYY333-1 (HKAS 144364), culture KUNCC 25-20222.

Notes: In the integrated phylogenetic analyses encompassing LSU, ITS, and SSU, our isolates exhibited a robust clustering with Tetraploa hainanensis (GZCC 23-0601), attaining statistical support values of 100% ML/1.00 BYPP (Fig. 16). Sequence comparison at the nucleotide level between our new isolate and T. hainanensis (GZCC 23-0601) highlighted divergences of 0.25% (807/809 bp) for LSU, 1.2% (492/498 bp) for ITS, and 0% (780/780 bp) for SSU. Furthermore, both taxa exhibit analogous morphological traits, characterized by cylindrical, verrucose conidia arranged in four columns of cells, accompanied by cylindrical and septate appendages. However, our isolate differs from the T. hainanensis (GZCC 23-0601) in having lower number of septate of appendages (4–8-septate vs. 5–16-septate) (Tang et al. 2023). Tetraploa hainanensis was reported from the unidentified decaying wood in Hainan Province, China. This study provides the first host record of T. hainanensis on a dead stem of Zea mays with evidence of morpho-molecular characterizations.

Fig. 13. Periconia xizangensis (HKAS 136110, holotype) a Colonies on natural substrate. b, c Conidiophores and conidiogenous cells. d–f Conidiogenous cells with developing conidia. g Conidia. Scale bars: b–d = 200 μm, e, f = 20 μm. g = 10 μm.

Fig. 14. IQ-tree based on ITS, LSU, tef1-α, rpb2 and tub2 sequences, SH-aLRT support (greater than 90, left) and Bayesian Posterior Probability (greater than 0.9, right) are shown in nodes. Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Torulaceae Corda

Torula Pers

Torula was introduced by Persoon in 1795, and is characterized by its type species T. herbarum. This genus is primarily recognized through its asexual reproductive structures, which include macronematous, mononematous conidiophores, holoblastic conidiogenous cells that exhibit a clavate to ellipsoidal morphology, and acrogenous, verrucose structures bearing septate conidia (Jayawardena et al. 2022; Hyde et al. 2023; He et al. 2024a, 2025). Most species within the Torula are found inhabiting freshwater and terrestrial environments (Crous et al. 2015a; Li et al. 2017a; He et al. 2024b). Torula is a widely distributed genus, having been documented across various regions including Asia (China, India, Thailand), Europe (France, Germany, Italy), North America (Crous et al. 2015a; Li et al. 2017a; Su et al. 2018; Boonmee et al. 2021; He et al. 2025; Zhang et al. 2025). It holds the distinction of being the largest genus within Torulaceae (He et al. 2024b), with 571 epithets in Index Fungorum (accessed on April 20, 2026). However, it is noteworthy that the majority of these species lack molecular data (Crous et al. 2015a), with only 34 species being corroborated by molecular data (He et al. 2025; Zhang et al. 2025). In this study, based on phylogenetic and morphological analysis and following the guidance of Maharachchikumbura et al. (2021), we introduced a new species, T. motuoensis, from Xizang, China.

Fig. 15. Setophoma manglietiae (HKAS 154724, holotype). a Colony on PDA. b Colony on SNA, HMAS 287387. c Colony on WA. d appearance of leaf spots. e hypha. f, g Chliamydospores. Scale bars: e = 30 μm, f = 20 μm, g = 10 μm.

10. Torula motuoensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Fig. 19

Chinese name

Index Fungorum number: IF 905258, Facesoffungi number: FoF 19300

Etymology: Referring to the location “Motuo County” where the holotype of this fungus was collected.

Holotype: HKAS 150299

Saprobic on dead stems of Ageratina adenophora. Sexual morph: Undetermined. Asexual morph: Colonies effuse, medium dense, hairy, black brown on the host. Mycelium immersed on the substrate, composed of septate, chained, branched, black brown hyphae. Conidiophores macronematous, mononematous, erect, flexuous, branched, verruculose, septate, cylindrical, brown to dark brown, 18.6–54.4 × 6–8.8 μm ( =32.5 × 7.2 μm, n = 20). Conidiogenous cells holoblastic, terminal, erect, ellipsoid, with melanized basal regions, bowl, pale brown to brown. Conidia phragmospores, catenate, acropleurogenous, monilioid, simple, dry, verruculose, guttulate, thick-walled, rough-walled, 0–multi-septate, pale brown to brown, 10–62 × 6–9 μm ( = 38 × 8 μm, n = 30).

Culture characteristics: germinating within 24 h on PDA at 25 °C, reaching 6.4–6.6 cm after 30 days incubation, above colony black, reverse dark brown, edge, entire, undulate, umbonate, surface rough, mycelia dense, no pigment.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°35’N, 95°14′E, 1730 m a.s.l., on dead stems of Ageratina adenophora (Asteraceae), 12 July 2022, Shu-Cheng He, HSC821 (HKAS 150299, holotype), ex-type culture: KUNCC 23-17479. ibid. HSC805, (HKAS 150300), culture: KUNCC 23-17350.

Notes: The phylogenetic analysis indicated that our isolates of T. motuoensis (KUNCC 23-17479 and KUNCC 23-17350) clustered with T. calceiformis (HKAS 125551 and HKAS 125552) (Fig. 18). Torula calceiformis is known to exhibit saprobic associations with unidentified wood samples sourced from Guizhou, China (Hyde et al. 2023). Our isolates were collected from Ageratina adenophora in Linzhi City, Xizang, China. Morphologically, T. motuoensis resembles T. calceiformis in having phragmospores, and monilioid, verruculose, multi-septate conidia. Compared to T. calceiformis, T. motuoensis has longer conidiophores (32.5 × 7.2 vs. 5.5 × 6.5 μm) and conidia (38 × 8 vs. 27.6 × 9.2 μm). Therefore, based on phylogenetic and morphological analyses, we introduce a new species, T. motuoensis, from Linzhi City, Xizang, China.

Fig. 16. Maximum likelihood majority rule consensus tree for Tetraploa using ITS, LSU and SSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML /BYPP. The tree is rooted with Pseudotetraploa bambusicola (CGMCC 3.20939), P. curviappendiculata (HHUF 28582) and P. javanica (HHUF 28596). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 17. Tetraploa hainanensis (HKAS 144363) a, b Colony on natural substrates. c–e Conidiophores with conidia. f Conidiogenous cells with conidia. g, h Conidia. Scale bars: c–e = 100 μm, f–h = 10 μm.

Fig. 18. Maximum likelihood majority rule consensus tree for Torula using ITS, LSU, SSU, rpb2 and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.90 are indicated near branches as ML /BYPP. The tree is rooted with Neotorula aquatica (MFLUCC 15-0342) and N. nujiangensis (HKAS 131234). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Tubeufiales Boonmee & K.D. Hyde

Tubeufiaceae M.E. Barr

Aquaphila Goh, K.D. Hyde & W.H. Ho

Aquaphila was established by Goh et al. (1998b) and typified by A. albicans based on morphological characteristics. Tsui et al. (2007) accepted Aquaphila as a member of Tubeufiaceae based on their phylogenetic analyses. Currently, Aquaphila comprises two species: A. albicans and A. edentata (Lu et al. 2018). Aquaphila albicans is characterized by semi-macronematous, mononematous, hyaline, delicate, septate, simple or branched, flexuous and geniculate conidiophores; integrated, terminal or intercalary, denticulate, monoblastic or polyblastic, proliferation sympodial conidiogenous cells; acrogenous, proliferation, hyaline, fusoid to falcate or sigmoid conidia (Goh et al. 1998b).

11. Aquaphila albicans Goh, K.D. Hyde & W.H. Ho, Mycol. Res. 102(5): 588 (1998) Fig. 21

Chinese name: – 白喜水孢 (bai xi shui bao)

Index Fungorum number: IF 443558; Facesoffungi number: FoF 02356

Saprobic on decaying wood submerged in freshwater habitats. Sexual morph: Undetermined. Asexual morph: Colonies superficial, hairy, velvety, white, conidia arising from aerial hyphae. Mycelium partly immersed in the woody substratum and partly superficial, consisting of septate, branched, hyaline, smooth, repent hyphae. Conidiophores 15–52 × 4–8 μm ( = 30 × 5 μm, n = 10), semi-macronematous, arising singly as lateral branches from the procumbent hyphae, cylindrical, simple or branched, septate, thin-walled, smooth, indistinctly septate, hyaline, flexuous or geniculate. Conidiogenous cells monoblastic or polyblastic, denticulate, sympodial proliferations, cylindrical. Conidia 58–78 × 8–10 μm ( = 65 × 9 μm, n = 30), holoblastic, solitary, borne acrogenously on conidiogenous denticles, becoming lateral after proliferation of the conidiophore, hyaline or very pale yellowish, predominantly fusoid to sickle-shaped, sometimes sigmoid, very rarely straight and obclavate, thin-walled, smooth, 9–12-euseptate, not constricted at the septa, heavily guttulate, conically rounded at both ends slightly broad at the base, basal cell obconical but not pedicellate.

Fig. 19. Torula motuoensis (HKAS 150299, holotype) a–c Colony on natural substrates. d, e Conidiophores. f–i Conidiogenous cells. j–m Conidia. n Germinated conidia on PDA. o, p Culture on the PDA. Scale bars: d-i = 20 μm, j-n = 25 μm.

Culture characteristics: Conidia germinating on PDA medium within 48 h and germ tubes produced from both ends. Colonies grown on PDA reached 30 mm diam at room temperature in two weeks, circular, with velutinous appearance, flat, light brown aerial mycelium on the surface, from below dark brown with entire margin.

Material examined: CHINA, Xizang Autonomous Region, Shigatse City, Yadong County, Xiayadong Township, 27°23′38.03" N, 88°50′15.01" E, 4070 m a.s.l., on decaying wood submerged in a freshwater stream, 26 July 2023, R.J. Xu, LJN-56 (HKAS 135977), culture KUNCC 24-17963.

Notes: Goh et al. (1998b) introduced Aquaphila albicans from submerged wood in a tropical habitat. Our newly isolated strain (KUNCC 24-17963) clusters with A. albicans (MFLUCC 16-0010 and KUMCC 19-0078) with 100% ML/1.00 BYPP support (Fig. 20). Morphologically, our new isolate shares the same morphological characteristics with the holotype (HKU(M) 2856) of A. albicans, thus we identified our new isolate as A. albicans.

Eurotiomycetes O.E. Erikss & Winka

Chaetothyriales M.E. Barr

Herpotrichiellaceae Munk

Cladophialophora Borelli

Cladophialophora was established by Borelli (1980), with C. ajelaloi designated as the type species, now synonymized with C. carrionii (Badali et al. 2008). It is characterized by the production of branched or unbranched chains of conidia through blastic conidiogenesis, a process distinguished by hyaline conidial scars (Sutton et al. 2009). Members of this genus exhibit remarkable ecological adaptability, with approximately 85 accepted species inhabiting a range of environments, from soil and plants in terrestrial ecosystems to unusual substrates such as sports drinks and lichens (Obase et al. 2015; Tang et al. 2017; Wijayawardene et al. 2022; Chang et al. 2023; Thitla et al. 2023; Hyde et al. 2024a). Cladophialophora exhibits a widespread geographical distribution, being found across the continents of Africa, Asia, Europe, North America, Oceania, and South America (De Hoog et al. 1995; Thitla et al. 2023). Species such as C. nyingchiensis and C. tumulicola have been isolated from rocks, demonstrating the genus's ability to colonize various substrates, including aquatic and semi-aquatic environments (Isola et al. 2016; Kiyuna et al. 2017; Sun et al. 2020; Chang et al. 2023; Thitla et al. 2023).

Fig. 20. Maximum likelihood majority rule consensus tree for Tubeufiaceae using ITS, LSU, SSU, rpb2 and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.90 are indicated near branches as ML /BYPP. The tree is rooted with Honghemyces pterolobii (KUMCC 20-0218 and KUMCC 21-0030). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

12. Cladophialophora biguttulata R.J. Xu & Q. Zhao, sp. nov. Fig. 23

Chinese name

Index Fungorum number: IF 903480; Facesoffungi number: FoF 17403

Etymology: the species epithet "biguttulata" refers to the presence of two large guttules within the conidia, which is a characteristic feature of this species.

Holotype: HKAS 136223

Fig. 21. Aquaphila albicans (HKAS 135977) a, b Colony on natural substrates. c, d Conidiophores with conidiogenous denticles. e–g Conidiophores with holoblastic conidia during development. h–k Conida. l Conidia germination. m, n Culture on PDA medium, m from above, n from below. Scale bars: c–l = 20 μm.

Saprobic on decaying wood submerged in freshwater habitats. Sexual morph: Undetermined. Asexual morph: Colonies superficial, effuse, velvety, gregarious, with masses white of conidia on the natural substrate. Mycelium immersed, consisting of hyaline, smooth-walled, branched, septate. Conidiophores semi-macronematous or macronematous, mononematous, cylindrical, straight or slightly flexuous, branched, dark brown or brown, capitate. Conidiogenous cells polyphialidic, with several sympodial proliferations, sometimes with conspicuous denticles, pale brown to subhyaline. Conidia 12–17 × 4–6 μm ( = 16 × 5 μm, n = 25), polyphialidic, reniform or cymbiform, curving, subhyaline, 1-septate, smooth, with two distinct guttules, short subhyaline pedicels.

Culture characteristics: Colonies grown on PDA reached 50 mm diam at room temperature in one month, circular, with dense, viscous, raised, dark brown mycelium on the surface, viscous, immersed mycelium on the surrounding, from below dark brown with entire margin.

Material examined: CHINA, Xinjiang Autonomous Region, Yili, Tacheng City, Dongdatang Scenic Spot, 43°52'42" N, 85°39'7.2" E, 1779 m a.s.l., on decaying wood submerged in a freshwater habitat, 17 July 2021, R.J. Xu, MD-345 (HKAS 136223, holotype), ex-type culture KUNCC 10438.

Notes: Recent morphological studies of Cladophialophora have primarily focused on the asexual forms produced on culture media (Obase et al. 2015; Sun et al. 2020; Boonmee et al. 2021; Cometto et al. 2023). Our collection was isolated from a natural substrate of submerged decaying wood. Morphologically, our collection differs from other species within Cladophialophora in having polyphialidic, with several sympodial proliferations conidiogenous cells and polyphialidic, reniform or cymbiform, with two distinct guttules, short subhyaline pedicels conidia (Sutton et al. 2009; Chang et al. 2023; Thitla et al. 2023). In addition, phylogenetic analyses also reveal that our collection C. biguttulata (KUNCC 10438) forms a separate, basal clade in Cladophialophora (Fig. 22), indicating it represents a new species.

Fig. 22. Maximum likelihood majority rule consensus tree for Cladophialophora using SSU, ITS, LSU and tef1-α sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML /BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Arachnomyces bostrychodes (CBS H-24452) and A. jinanicus (CGMCC 3.14173). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Thysanorea Arzanlou, W. Gams & Crous

Thysanorea was established by Arzanlou et al. (2007), with T. papuana, formerly known as Periconiella papuana, as the type species. Thysanorea is characterized by micro- or macronematous, erect conidiophores that are either simple or apically branched, occasionally proliferating percurrently at the apex. The conidiogenous cells are terminal or intercalary, polyblastic, and exhibit a color gradient, transitioning from brown at the base to a paler shade towards the apex. The conidia are generally pale brown and vary in shape, including oblong, obovoid, and cylindrical forms (Hernández-Restrepo et al. 2020; Yang et al. 2023). Hernández-Restrepo et al. (2020) later transferred species from Minimelanolocus to Thysanorea, citing their congeneric nature based on phylogenetic analyses. However, this reclassification has been debated, with Wan et al. (2021) challenging the move due to differences in the branching pattern of the conidiophores.

Fig. 23. Cladophialophora biguttulata (HKAS 136223, holotype) a Colony on natural substrates. b–d Conidiophores with conidia. e Conidiogenous cells with conidia. f–h Conidia. i Culture on PDA medium. Scale bars: b–d = 20 μm, e–h = 10 μm.

13. Thysanorea linzhiensis R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 25

Chinese name

Index Fungorum number: IF 905033; Facesoffungi number: FoF 19301

Etymology: The epithet refers to Linzhi City, located in the Xizang Autonomous Region of China, where the type specimen was collected.

Holotype: HKAS 136140

Saprobic on decaying submerged wood in a freshwater stream. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrates effuse, hairy, scattered or in small groups, brown, with glistening conidial masses at apex. Mycelium is partly superficial, partly immersed, composed of septate, smooth, pale brown hyphae. Conidiophores 201–320 × 3–9 μm ( = 263 × 6 μm, n = 20), macronematous, mononematous, erect, solitary, straight or flexuous, cylindrical, smooth-walled, septate, unbranched, dark brown, becoming pale brown or subhyaline towards the apex, thick-walled. Conidiogenous cells polyblastic, integrated, terminal, denticulate, sympodial proliferations, cylindrical, slightly constricted at septa, pale brown or subhyaline. Conidia 14–18 × 3–4 μm ( = 16 × 3 μm, n = 20), acrogenous or acropleurogenous, clavate, with a narrow truncate base, truncate at the base, aseptate when young, 1–4-septate when mature, smooth-walled, subhyaline.

Culture characteristics: Conidia germinating on PDA medium within 24 h and germ tubes produced from both ends. Colonies grown on PDA reached 30 mm diam at room temperature in two weeks, circular, with velvety, raised, grayish-green aerial mycelium on the surface, from below grayish-green with entire margin.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Chayu County, 28°39′55.76" N, 97°28′15.71" E, 4118 m a.s.l., on decaying wood submerged in a freshwater stream, 14 July 2022, R.J. Xu, LTS-36 (HKAS 136140, holotype), ex-type culture KUNCC 24-18006.

Notes: Thysanorea linzhiensis resembles T. obscurus in possessing mononematous, macronematous, septate conidiophores, as well as polyblastic, integrated, sympodially proliferating conidiogenous cells, and clavate conidia with a narrow truncate base. However, T. linzhiensis can be distinguished from T. obscurus by its denticulate conidiogenous cells and longer conidiophores (201–320 × 3–9 μm vs. 123–219 × 5–6 μm) (Liu et al. 2015b). In the phylogenetic analysis using combined ITS, LSU, and SSU sequence data, T. linzhiensis formed a distinct clade within Thysanorea (Fig. 24), closely related to but clearly separate from T. obscurus, with a genetic divergence of 2.8% in ITS sequences (16/564 bp difference). The introduction of T. linzhiensis as a new species is supported by both morphological and phylogenetic evidence.

Leotiomycetes O.E. Erikss. & Winka

Helotiales Nannf. ex Korf & Lizoň

Mollisiaceae Rehm

Phialocephala W.B. Kendr.

Phialocephala was introduced by Kendrick (1961) to accommodate P. dimorphospora (type species) and P. bactrospora, the latter was recently transferred to Parasporendocladia (Wu & Diao 2022). Phialocephala is a polyphyletic genus throughout Mollisiaceae (Tanney & Seifert 2020; Visagie et al. 2024). Tanney & Seifert (2020) defined the Phialocephala sensu stricto which contains 14 species. Phialocephala is characterized by macronematous, mononematous, brown conidiophores, monophialidic, discrete conidiogenous cells arranged penicillately, and often catenate, subglobose to ellipsoidal, hyaline or pigmented, aseptate conidia (Ellis 1971b; Onofri et al. 1994; Tanney et al. 2016; Tanney & Seifert 2020; Senanayake et al. 2023). Phialocephala also has diplococcium-like synasexual morph, e.g., P. catenospora, and synnematous synasexual morph, e.g., P. oblonga (Tanney et al. 2016).

14. Phialocephala guizhouensis J. Ma & Y.Z. Lu, sp. nov. Fig. 27

Chinese name

Index Fungorum number: IF 573589; Facesoffungi number: FoF 19302

Etymology: the species epithet "guizhouensis" refers to the location where this species was collected

Holotype: GZAAS 23-0033

Saprobic on decaying wood in a terrestrial habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate effuse, black, hairy. Mycelium mostly partly immersed, partly superficial, composed of brown, branched, septate, smooth hyphae. Conidiophores up to 220 μm long, macronematous, mononematous, erect, cylindrical, brown to reddish-brown, septate, thick-walled. Conidiogenous cells polytretic, integrated, terminal and intercalary, brown to reddish-brown. Conidia 6–13 × 3–5 μm ( = 8.5 × 4 μm, n = 30), catenate, formed in acropetal, branched chains, oblong with obtuse ends, brown to slight dark brown, 1–2-septate, darkened and slightly constricted at the septa, smooth.

Culture characteristics: Conidia germinate on water agar within 24 hrs. Germ tubes are produced from the apex or base. Colonies superficial, irregularly circular, flat, with entire edge, from above grey to whitish-grey in the center, brownish-gray at the edge, and from below, pale grey in the middle, brownish-grey at the edge.

Material examined: CHINA, Guizhou Province, Qiannan Buyi Autonomous Prefecture, Longli County, on decaying wood in a terrestrial habitat, 2 September 2020, Jian Ma, LLZB3 (GZAAS 23-0033, holotype), ex-type cultures GZCC 24-0159.

Notes: Our collection morphologically resembles the synasexual morph of Phialocephala catenospora, which has oblong and brown conidia (Tanney et al. 2016). However, the conidia of P. catenospora are larger than those of ours, (6–13 × 3–5 vs. 12–22.5 × 5–6 μm). Besides, P. catenospora (DAOMC 250108) is phylogenetically distant from our strain LLZB3 in the analyses (Fig. 26). In the phylogenetic tree, our strain LLZB3 formed an internal subclade with P. amethystea (DAOMC 251552) and P. compacta (CBS 507.94) with 99% ML/1.00 BYPP support (Fig. 26). ITS comparison shows there are 7.11% (53 out of 746 bp, including 7 gaps) and 4.97% (37 out of 745 bp, including 1 gap) differences with the strains DAOMC 251552 and CBS 507.94, respectively. Therefore, we introduce our collection as a new species.

Fig. 24. Maximum likelihood majority rule consensus tree for Thysanorea using ITS, LSU and SSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Brycekendrickomyces acaciae (CBS 124104) and Metulocladosporiella musae (CBS 113863). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 25. Thysanorea linzhiensis (HKAS 136140, holotype) a Colony on natural substrates. b, c Conidiophores with conidia. d–g Conidiogenous cells with developmental conidia. h–r Conidia. s Germinated conidium. t, u Culture on PDA medium, t from above, u from below. Scale bars: b–e = 50 μm, f, g = 20 μm, h–k = 10 μm.

Fig. 26. ML tree (−lnL = 16244.186) based on the combined ITS-LSU-LNS2-7 rpb1-TOP1 sequences. The combined dataset comprises 36 strains, including the new collection. The alignment comprises 3,919 characters (ITS: 1–659, LSU: 660–1870, LNS2: 1871–2151, rpb1: 2152–2965, TOP1: 2966–3919) including gaps. Among them, number of constant sites is 2,853, and number of parsimony informative sites are 676. Bootstrap support values for ML greater than 75% and PP greater than 0.95 are given near nodes as ML-BS/PP. The tree is rooted with Leotia lubrica (AFTOL-ID 1). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 27. Phialocephala guizhouensis (GZAAS 23-0033, holotype) a Colonies on natural substrate. b, c Conidiophores and conidia. d Conidiogenous cells and conidia. e–h Conidia. Scale bars: b–c = 50 μm, d = 30 μm, e–i = 10 μm.

Pezizellaceae Velen

Chalara (Corda) Rabenh

Chalara was originally introduced by Corda (1837) as a subgenus of Torula to accommodate the hyphomycete Chalara fusidioides, characterized by a granular, effuse stroma, simple conidiophores, lageniform, cylindrical conidiogenous cells, and cylindrical, hyaline conidia. Rabenhorst (1844) elevated Chalara to the generic level. The genus Chalara comprises a group of hyphomycetes characterized by the presence or absence of stroma and setae; conidiophores are scattered or aggregated, erect, simple, subcylindrical to cylindrical, smooth or verrucose, septate or aseptate; conidiogenous cells ampulliform, lageniform, obclavate, ellipsoidal, or subcylindrical, with a venter and a collarette; conidia cylindrical to obclavate, hyaline, aseptate, and extruded in short chains (Nag Raj & Kendrick 1975; Wu & Diao 2023). Based on morphological and phylogenetic analyses, Wu & Diao (2023) revised the genus Chalara, typified by C. fusidioides, into a narrowly concept monophyletic genus to accommodate seven species that formed a distinct clade separate from other chalara-like fungi in their phylogenetic reconstruction. These species include Chalara africana, C. bambusicola, C. clidemiae, C. cylindrosperma, C. longiphora, C. platanicola, and C. qinlingensis (Wu & Diao 2023). In addition to these phylogenetically supported species, Chalara sensu stricto also includes nine other species for which molecular data are currently lacking (Wu & Diao 2023). Species of Chalara s. str. are saprobic and occur on various plant substrates and in soil (Nag Raj & Kendrick 1975; Kirk 1986; Christiansen 1993; McKenzie et al. 2002; Wu 2004; Pratibha et al. 2005; Wu & Diao 2023)

15. Chalara aquatica H.W. Shen, K.D. Hyde & Z.L. Luo, sp. nov. Fig. 28

Chinese name

Index Fungorum number: IF 905259; Facesoffungi number: FoF 19303

Etymology: The specific epithet “aquatica” refers to the aquatica habitat, which is where this species was collected.

Holotype: HKAS 151651

Saprobic on submerged decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate, superficial, effuse, scattered, hairy, dark brown, solitary or in small groups. Mycelium partly immersed and partly superficial, composed of branched, septate, pale brown hyphae. Stroma absent. Setae absent. Conidiophores (85–)115–171(–183) × 5.3–8.5 µm ( = 143 × 7.2 µm, n = 20), macronematous, mononematous, cylindrical, straight or slightly flexuous, unbranched, septate, basal cells slightly swollen, brown to dark brown, smooth-walled. Conidiogenous cells (30–)36–42(–44) × 7.4–9.3 µm ( = 39 × 8.3 µm, n = 25), integrated, terminal, lageniform, pale to medium brown, consisting of the wider venter and the narrower collarette, transition from venter to collarette abrupt. Venter 10–21 × 7–9 µm, ellipsoid to subcylindrical, medium brown. Collarette 18–26 × 4–6 µm, cylindrical, pale to medium brown. Conidia (3.5–)5.2–7.8(–8.6) × 3.5–5 µm ( = 6.5 × 4.2 µm, n = 30), endogenous, subglobose to barrel-shaped to rectangular, truncate or obtuse at both ends, hyaline, aseptate, guttulate, smooth-walled.

Culture characteristics: Conidia germinated on PDA within 12 h and germ tubes produced from around the conidium. Colonies on PDA reaching 2–3 cm diameter after one month at room temperature (around 22 °C) in the dark, dense, velvety, gray to brown, with smooth margins and bulge center from above, dense, gray to brown, with smooth margins from below.

Material examined: CHINA, Yunnan Province, Lijiang City, on the unidentified decaying wood, October 2021, L.L. Li, L944 (HKAS 151651, holotype), ex-type culture KUNCC 23-14304.

Notes: Phylogenetic analysis revealed that Chalara aquatica is closely related to C. cylindrosperma, C. longiphora, and C. platanicola, forming a robustly supported clade within the phylogenetic framework (Fig. 29). Morphologically, C. aquatica is similar to all three species, but can be distinguished from C. cylindrosperma by its larger venter (10–21 × 7–9 µm vs. 10–12.5 × 5–7 µm) and collarette (18–26 × 4–6 µm vs. 17.5–20 × 2.5–3.5 µm), as well as by its smaller conidia (5.2–7.8 × 3.5–5 µm vs. 10–17 × 1.5–2.5 µm) (Wu & Diao 2023). Chalara aquatica differs from C. longiphora in having a larger collarette (18–26 × 4–6 µm vs. 14–17 × 2.5–2.7 µm), wider conidia (3.5–5 µm vs. 2–2.5 µm), and subglobose to barrel-shaped or rectangular conidia, whereas the conidia of C. longiphora are cylindrical (Wu & Diao 2023). Compared to C. platanicola, C. aquatica has longer conidiogenous cells (36–42 µm vs. 26–27 µm), venter (10–21 × 7–9 µm vs. 10–12.5 × 5.7–6.5 µm), and collarette (18–26 × 4–6 µm vs. 17–18 × 2.8–3.3 µm), but produces shorter and wider conidia (5.2–7.8 × 3.5–5 µm vs. 9–10.5 × 2.2–2.5 µm) (Wu & Diao 2023). Based on both phylogenetic distinction and morphological differences, we introduce C. aquatica as a new species.

16. Chalara guttulata H.W. Shen, K.D. Hyde & Z.L. Luo, sp. nov. Fig. 30

Chinese name:

Index Fungorum number: IF 905260; Facesoffungi number: FoF 19304

Etymology: The specific epithet “guttulata” refers to the guttulate conidia of this species.

Holotype: DLU 3129

Saprobic on submerged decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate, superficial, effuse, scattered, hairy, dark brown, solitary or in small groups. Mycelium partly immersed and partly superficial, composed of branched, septate, pale brown hyphae. Stroma absent. Setae absent. Conidiophores 82–88 × 4.3–6 µm (x̄ = 85 × 5 µm, n = 15), macronematous, mononematous, cylindrical, straight or slightly flexuous, unbranched, 3–septate, basal cells slightly swollen, brown to dark brown, smooth-walled. Conidiogenous cells 50–60 × 5–6.4 µm ( = 56 × 5.5 µm, n = 15), integrated, terminal, long-lageniform, pale to medium brown, consisting of the wider venter and the narrower collarette, transition from venter to collarette abrupt. Venter 21–31 × 4.5–6 µm, subcylindrical to long-clavate, medium brown. Collarette 25.5–32 × 2.8–3.1 µm, cylindrical, pale to medium brown. Conidia 7.8–12 × 1.8–2.5 µm ( = 10 × 2.2 µm, n = 40), endogenous, cylindrical, truncate or obtuse at both ends, hyaline, aseptate, guttulate, smooth-walled.

Material examined: CHINA, Yunnan Province, Dali City, on undetermined decaying wood, May 2021, S. Luan, S3129 (DLU3129, holotype), ex-type culture DLUCC 3129.

Notes: Phylogenetic analysis placed Chalara guttulata within the Chalara clade, where it formed a distinct and with 100% ML/1.00 BYPP support (Fig. 29). Morphologically, C. guttulata resembles C. versicolor in having cylindrical, unbranched, septate conidiophores, integrated, terminal, lageniform conidiogenous cells, consisting of the wider venter and the narrower collarette, and cylindrical, hyaline, aseptate conidia, truncate or obtuse at both ends (Wu & Diao 2023). However, it differs by its longer conidiophores (82–88µm vs. 15–50 µm), longer conidiogenous cells (50–60 µm vs. 47–50 µm), and smaller conidia (7.8–12 × 1.8–2.5 µm vs. 6.5–15 × 2.5–3 µm) (Wu & Diao 2023). A BLASTn search of the ITS sequence against NCBI GenBank revealed the closest match to be Chalara sp. (Specimen voucher: HAY-F-011218, Sequence ID: PV742674.1), with 98% identity (485/496 bp, including 2 gaps). The closest match for the LSU sequence was Endoradiciella communis (Strain: P2333, Sequence ID: NG_149040.1), showing 99% identity (857/864 bp, including 1 gap). Based on both morphological characteristics and molecular phylogenetic evidence, we introduce C. guttulata as a novel species.

Sordariomycetes O.E. Erikss. & Winka

Sordariomycetes is one of the largest classes within Ascomycota, widely distributed worldwide and predominantly comprising terrestrial taxa, although several members also occur in aquatic habitats (Luo et al. 2019; Hyde et al. 2020c). In the present study, a phylogenetic framework of Sordariomycetes is reconstructed based on sequence data from three loci (LSU, rpb2 and tef1-α) using approximately 1,000 strains. Further details are provided in Supplementary (Fig. 2).

Fig. 28. Chalara aquatica (HKAS 151651, holotype) a, b Colonies on woody substrates. c–e Conidiophores. f Conidiogenous cell and conidium g–i Conidiogenous cells with developing conidia. j–l Conidia. h, i Conidia. m Germinated conidium. n Culture on PDA, left from surface, right from back. Scale bars: c–e = 40 μm, f–i = 15 μm, j–m = 10 μm.

Fig. 29. Phylogram generated from maximum likelihood analysis based on combined ITS and LSU. Thirty-seven strains were included in the combined sequence analyses, which comprised 3817 characters with gaps (ITS = 464 and LSU = 825). Single gene analyses were also performed, and topology and clade stability were compared from the combined gene analyses. Lareunionomyces loeiensis (CGMCC3.23405 and NN047769) strains were used as the outgroup taxa. The final ML optimization likelihood is -27926.145972. The matrix included 1676 distinct alignment patterns, with 21.83% undetermined characters or gaps. Estimated base frequencies were obtained as follows: A = 0.234861, C = 0.271796, G = 0.258425, T = 0.234918; substitution rates AC = 1.379110, AG = 3.330310, AT = 1.229619, CG = 1.122043, CT = 5.312755, GT = 1.000000; gamma distribution. Bootstrap support values for ML (first set) equal to or greater than 70%, BYPP equal to or greater than 0.95 are given above or below the nodes. Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández

Chaetosphaeriaceae Réblová, M.E. Barr & Samuels

Chaetosphaeriaceae was established by Réblová et al. (1999a) to accommodate Chaetosphaeria with the closely related genera including: Ascocodinaea, Melanochaela, and Sporoschisma. Chaetosphaeriaceae is a major family within Sordariomycetes, known for its complex pleomorphism and diverse asexual morphologies. Previous, the phylogenetic analyses have revealed that both Stanjehughesia and Linkosia are polyphyletic (Shenoy et al. 2006; Su et al. 2016a; Hyde et al. 2020c; Hsieh et al. 2021), and the type species S. hormiscioides lacks molecular data, leaving the phylogenetic position of the genus unresolved. However, Réblová et al. (2024) re-evaluated species previously classified in the genera Exserticlava, Phaeostalagmus, Phialocephala, and several Chalara and Stanjehughesia-like fungi, and reassigned them to Chaetosphaeriaceae based on living cultures and DNA sequence data obtained from numerous type species. Currently, 101 genera are accepted in Chaetosphaeriaceae and phylogenetic analysis based on combined ITS, LSU, and tef1-α sequence data (Hyde et al. 2024a; Réblová et al. 2024).

Chloridium Link

Chloridium is known for its cosmopolitan distribution, thriving on decaying plant tissues in terrestrial and freshwater habitats, notably on wood buried in moist soil or leaf litter. Chloridium was introduced with C. virescens as the type species, characterized by pigmented, unbranched, erect conidiophores with globose conidia forming green masses (Link 1809; Réblová et al. 2022). The taxonomy of Chloridium has been complex, with early classifications based on morphological features and later, molecular data. Recent molecular analyses have further refined the understanding of phylogeny of Chloridium, revealing its paraphyletic nature and close relationships with other genera such as Adautomilanezia and Sporoschisma (Réblová et al. 2022).

Fig. 30. Chalara guttulata (DLU3129, holotype) a Colonies on woody substrates. b–d Conidiophores. e Conidiogenous cell f, g Conidia. Scale bars: b–e = 20 μm, f, g = 10 μm.

17. Chloridium fuscum (Corda) Réblová & Hern. -Restr., Stud. Mycol. 103: 176 (2022) Fig. 32

Chinese name

Index Fungorum number: IF 555386; Facesoffungi number: FoF 04950

Saprobic on decaying wood submerged in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies effuse, dark brown, olivaceous brown to olivaceous grey, with long hairy mycelium. Mycelium partly immersed, partly superficial, consisting of branched, septate, brown hyphae. Conidiophores 158–241 × 3–6 µm ( = 195 × 5 μm, n = 15), macronematous, solitary or crowded, erect, straight, septate, unbranched, cylindrical, 4–6- septate, dark brown, pale brown to subhyaline towards the apex, slightly tapering towards the apex. Conidiogenous cells 11–31 × 3–5 µm ( = 21 × 4 μm, n = 20), monophialidic, discrete, phialide, sometimes extending percurrently, subulate to cylindrical, pale brown to subhyaline, hyaline towards the apex, in 5–10 cylindrical located in the two lower thirds of the conidiophore in the nodes at intervals. Conidia 4–5 × 3–4 µm ( = 5 × 3 μm, n = 20), acrogenous, ellipsoidal, subhyaline to pale brown, aseptate, smooth,

Culture characteristics: Conidia germinating on PDA medium within 24 h and germ tubes produced from both ends, Colonies grown on PDA reached 50 mm diam at room temperature in two weeks, circular, with velvety, flat, dark brown mycelium on the surface, from below dark brown to grayish-brown with entire margin.

Material examined: CHINA, Xizang Autonomous Region, Rikaze City, Yadong County, 27° 21' 11" N, 88° 58' 10" E, 3061 m a.s.l., on decaying wood submerged in a freshwater habitat, 01 July 2022, R.J. Xu, LTS-8 (HKAS 136133), culture KUNCC 24-17992. Linzhi City, Motuo County, Dexing Township, 29°21′14.97" N, 95°8′53.19" E, 2172 m a.s.l., on decaying wood submerged in a freshwater stream, 10 August 2023, R.J. Xu, LJN-42 (HKAS 135976), culture KUNCC 24-17960. Shannan City, Luozha County, 28°11′3.15" N, 90°59′57.88" E, 3491 m a.s.l., on decaying wood submerged in a freshwater habitat, 08 July 2022, R.J. Xu, XK-68 (HKAS 136102), culture KUNCC 24-18124. Linzhi City, Chayu County, Guyu Township, 29°19′31.89" N, 97°8′52.91" E, 3858 m a.s.l., on decaying wood submerged in a freshwater habitat, 12 August 2023, R.J. Xu, LJN-72 (HKAS 135984), culture KUNCC 24-17972.

Notes: Chloridium fuscum was initially described by Corda (1837) as Chaetopsis fusca. It was later synonymized with Gonytrichum and subsequently reclassified under Chloridium based on comprehensive phylogenetic analyses (Saccardo 1880; Hughes 1951; Réblová et al. 2022). Synonyms of this species include Mesobotrys simplex and Chl. aseptatum. Chl. fuscum is closely related to Chl. volubile, sharing similar conidiophore characters, but can be distinguished based on specific morphological differences (Réblová et al. 2022).

Chloridium fuscum is a widely distributed saprobic species that can be found in both terrestrial and freshwater environments, including soil, decaying wood and bark, thriving on a variety of plant remnants. This species exhibits a broad geographic presence, recorded in regions such as Australia, Canada, China, the Czech Republic, Germany, Italy, Korea, Puerto Rico, South Africa, Thailand, the United Kingdom, and the USA (Réblová et al. 2022). Morphologically, our collection fits well with the species concept of Chl. fuscum, and phylogenetic analyses showed that our collections of Chlo. fuscum clustered closely with Chl. fuscum (CBS 195.60) (Fig. 31). Consequently, we identified the four strains as Chl. fuscum and reported as a new collection from Xizang, China.

Reticulascus was established by Réblová et al. (2011a), based on a combined of ITS, LSU and SSU phylogeny and to accommodate two holomorph Chaetosphaeria species. Whereafter, Réblová et al. (2016a) given its widespread use, priority, and greater number of names, therefore recommend the use of Cylindrotrichum rather than Reticulascus. The genus is characterized by brown to black, hairy, effuse colonies in vivo. Setae are absent. Conidiophores are macronematous, mononematous, cylindrical, and straight. Conidiogenous cells are usually monophialidic, rarely polyphialidic with up to two lateral enteroblastic openings, with collarette hyaline to subhyaline. Conidia are cylindrical, slightly tapering, rounded at the apex, obtuse at the base, 1-septate, not constricted at the septum, hyaline, guttulate, and smooth-walled.

18. Cylindrotrichum clavatum W. Gams & Hol. -Jech., Stud. Mycol. 43: 54. 1976 Fig. 34

= Reticulascus clavatus Réblová & J. Fourn., in Réblová, Gams & Seifert, Stud. Mycol. 68: 181 (2011)

Chinese name

Index Fungorum number: IF 555386; Facesoffungi number: FoF 04950

Saprobic on decaying wood submerged in a freshwater river. Sexual morph: Reticulascus clavatus. Asexual morph: Colonies superficial, effuse, glistening, hairy or velvety, pale brown to brown. Mycelium immersed, consisting of hyaline, smooth-walled, branched, septate. Conidiophores 105–182 × 3–5 μm ( = 153 × 4 μm, n = 15), macronematous, mononematous, solitary or 2-group, erect, unbranched, 4–8-septate, straight or flexuous, cylindrical, dark brown at the base, light brown towards the apex. Conidiogenous cells 22–27 × 4–5 μm ( = 24 × 4 μm, n = 20), monophialidic, polyphialidic, collarette hyaline to subhyaline. Conidia 7–11 × 3–4 μm ( = 9 × 3 μm, n = 25), acrogenous, cylindrical or clavate, rounded at apex, slightly tapering, obtuse at base, 0–1-septate, hyaline, smooth-walled.

Culture characteristics: Conidia germinating on PDA medium within 24 h and germ tubes produced from both ends. Colonies grown on PDA reached 30 mm diam at room temperature in two weeks, irregular, with velvety, flat, aerial mycelium on the surface, from below dark brown with entire margin.

Material examined: CHINA, Xinjiang Autonomous Region, Aksu City, Aksu River, 41° 8' 13.8" N, 80° 8' 42.8" E, 1112 m a.s.l, on decaying wood submerged in a freshwater river, 02 August 2021, R.J. Xu, MD-306 (HKAS 136202), culture KUNCC 24-18038.

Notes: Réblová et al. (2011a) introduced the new holomorph genus Reticulascus for two species, based on multi-gene phylogenetic analyses. This included the new species R. clavatus, with Cylindrotrichum clavatum identified as its anamorph. However, Réblová et al. (2016a) later advised using the name Cylindrotrichum instead of Reticulascus. This recommendation was based on the discovery that the type species of Cylindrotrichum, Cy. oligospermum, is the asexual morph of Chaetosphaeria tulasneorum, which is the type species of Reticulascus (Réblová 1999; Réblová et al. 2011a). Consequently, Cylindrotrichum and Reticulascus are synonymous.

Cylindrotrichum clavatum, originally found on submerged wood in freshwater habitats in France, was later reported by Maharachchikumbura et al. (2018) in the Lancang River, Yunnan Province. Our phylogenetic analyses showed that the isolate (KUNCC 24-18038) clusters with Cy. clavatum (DLUCC 0575, DLUCC 0572, CBS 125296, CBS 125297, CBS 125239 and CBS 428.76) (Fig. 33). The morphology of our strain's conidiophores, conidiogenous cells, and conidia aligns with those of Cy. clavatum, leading to its identification as a new collection from Xinjiang, China.

Falholtia W.P. Wu & Y.Z. Diao

Falholtia is a monotypic genus established by Wu & Diao (2022) to accommodate a synnematous Stanjehughesia-like taxon previously described as Stanjehughesia kaohsiungensis (Hsieh et al. 2021). This genus is morphologically distinct within Chaetosphaeriaceae by producing cylindrical, septate conidiophores aggregated into conspicuous synnemata, and determinate, monoblastic conidiogenous cells. The conidia are holoblastic, obclavate-rostrate, multi-euseptate (up to 27 septa), and brown to dark olivaceous, with the base truncate and the apex paler and often slightly curved. These features, particularly the well-developed synnemata and the unique conidial morphology, clearly distinguish Falholtia from Stanjehughesia, in which conidiophores are usually solitary or in clusters, and conidia are less complex.

Phylogenetically, Falholtia forms a distinct, well-supported clade within Chaetosphaeriaceae (Hsieh et al. 2021; Wu & Diao 2022), separated from Stanjehughesia and other Sporidesmium-like genera such as Ellisembia, Morrisiella, and Paliphora (Réblová et al. 2021; Luo et al. 2019). Falholtia represents one of the few genera in Chaetosphaeriaceae with non-phialidic, holoblastic conidiogenous cells. It adds to the diversity of non-phialidic anamorphs in the family, which are otherwise represented mainly by Sporidesmium-like fungi. Other related genera with similar conidiogenesis include Paliphora and Chaetosphaeria, though their conidial and conidiophore structures are notably different (Sivanesan & Sutton 1985; Luo et al. 2019).

Fig. 31. Maximum likelihood majority rule consensus tree for Chloridium using ITS, LSU and SSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Sporoschisma aquaticum (DLUCC 0628) and S. atroviride (GZCC 20-0490). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 32. Chloridium fuscum (HKAS 136133) a Colonies on natural substrate. b, c Conidiophores with conidiogenous cell. d–f Conidiogenous cell with developing conidia. g–j Conidia. k, l Culture on PDA medium, k from above, l from below. Scale bars: b, c = 50 μm, d–f = 10 μm, g–j = 1 μm.

Fig. 33. Maximum likelihood majority rule consensus tree for Cylindrotrichum using ITS, LSU and SSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML /BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Peethambara spirostriata (BPI843537). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Chinese name

Index Fungorum number: IF 905035; Facesoffungi number: FoF 19305

Etymology: the term “linzhiensis” refers to the locality where the type specimen was found.

Holotype: HKAS 141644

Saprobic on decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum, superficial, effuse, scattered, hairy, dark brown. Mycelium partly immersed and partly superficial, composed of branched, septate, brown to dark brown, smooth-walled hyphae. Conidiomata 480–1128 × 34–50 μm ( = 640 × 40 μm, n = 20), synnematous, solitary or in group, erect, cylindrical, dark brown to black, tapering towards the apex, composed of closely compacted conidiophores and hyphae. Conidiophores 16–23 × 4–6 μm ( = 19 × 5 μm, n = 20), short, 1(2)-septate, cylindrical, unbranched, dark brown to black, smooth-walled. Conidiogenous cells monoblastic, integrated, terminal, determinate, cylindrical, brown to dark brown, smooth-walled. Conidia 73–152 × 11–15 μm ( = 98 × 13 μm, n = 20), acrogenous, solitary, straight to slightly flexuous, obclavate-rostrate, tapering towards the rounded apex, truncate at the base, brown to dark olivaceous brown, hyaline at the apex, base cells cuneiform, 8–15-euseptate, slightly constricted at the septa, sometimes with a sheath at the apex.

Fig. 34. Cylindrotrichum clavatum (HKAS 136202) a Colony on natural substrates. b, c Conidiophores with conidia. d, e Conidiogenous cells with developmental conidia. f–i Conidia. j, k Culture on PDA medium, j from above, k from below. Scale bars: b–c = 50 μm, d–e = 10 μm, f–i = 5 μm.

Culture characteristics: Conidia germinate on PDA within 48 h, with germ tubes emerging from a subtruncate base. Colonies on PDA reach a diameter of 3 cm within one month at room temperature. These colonies are circular with irregular margin, raised in the middle, cream-colored mycelium that covers the entire colony.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Chayu County, 28° 29′ 39.27″ N, 96° 59′ 35.25″ E, 1537 m a.s.l., on decaying wood submerged in a freshwater stream, 14 July 2022, R.J. Xu, MD-152A (HKAS 141644, holotype), ex-type culture KUNCC 25-21369.

Notes: Falholtia linzhiensis shares the generic features of Falholtia by having compact synnemata composed of cylindrical, septate conidiophores, terminal, determinate, monoblastic conidiogenous cells, and holoblastic, obclavate-rostrate, euseptate conidia with a truncate base and paler apex (Hsieh et al. 2021; Wu & Diao 2022). However, F. linzhiensis is morphologically distinct from the type species F. kaohsiungensis in having shorter (16–23 μm vs. 13–34 μm), and conidia with fewer septa (8–15 vs. up to 27), often bearing an apical sheath, this feature not reported in the type. Additionally, F. linzhiensis forms conidiomata that are often scattered and hairy on the substrate surface, differing from the more compact and erect synnemata of F. kaohsiungensis.

Phylogenetic analysis based on ITS and LSU sequences places F. linzhiensis within Chaetosphaeriaceae, forming a well-supported and independent clade sister to F. kaohsiungensis (Fig. 35), thus supporting its recognition as a distinct taxon within the genus.

Fig. 35. Maximum likelihood consensus tree inferred from the combined ITS, LSU, and rpb2 multiple sequence alignments. ML support equal to or greater than 75% and BYPP value equal to or greater than 0.95 are given near the nodes. The scale bar indicates expected changes per site The tree is rooted to Tracylla eucahpti (CPC 31777). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 36. Falholtia linzhiensis (HKAS 141644, holotype) a, b Synnemata on natural substrates. c, d Synnemata, conidiophores and conidia. e Conidiophores with conidia. f conidiophores. g–j Conidia. k Germinated conidium. l, m Culture on PDA, l from above, m from below. Scale bars: b–f = 50 μm, g–l = 20 μm.

Phialosporostilbe Mercado & J. Mena

Phialosporostilbe was established by Sierra & Portales (1985) to accommodate P. turbinata. Members of Phialosporostilbe are characterized by the production of synnematous conidiophores bearing integrated, monophialidic or annellidic conidiogenous cells at the apices. The conidia are typically acrogenous, phialidic, catenate, and range from turbinate to cuneiform or cordiform in shape, colourless to pale brown, smooth-walled, and amerosporous, often with short subapical or apical appendages, such as setulae.

To date, seven species have been recognized worldwide: Phialosporostilbe catenata, P. dendroidea, P. gregariclava, P. scutiformis, P. setosa, P. turbinata and P. yadongensis, described from various habitats and geographic regions including Asia and the Americas (Sierra & Portales 1985; Bhat & Kendrick 1993; Shirouzu & Harada 2004; Sureshkumar et al. 2005; Wu & Zhang 2009; Yang et al. 2018b). Although similar in conidial shape and conidiogenesis to species of Nawawia, Phialosporostilbe is distinguishable by its synnematous rather than mononematous conidiophores, which has been the key feature supporting its separation at the generic level. Nonetheless, due to the similarities in conidial morphology and development, some authors previously considered Phialosporostilbe and Nawawia to be congeneric (Yang et al. 2018b), warranting further phylogenetic investigation to clarify their relationships.

20. Phialosporostilbe gaoligongensis R.J. Xu, Y. Li & Q. Zhao, sp. nov. Fig. 38

Chinese name: – 高黎贡瓶梗孢柱菌 (gao li gong ping geng zhu bao jun)

Index Fungorum number: IF905261; Facesoffungi number: FoF 19307

Etymology: The term “gaoligongensis” refers to the locality where the type specimen was found.

Holotype: HKAS 149332

Saprobic on decaying wood submerged in a freshwater stream. Sexual morph: Undetermined. Asexual morph: Colonies effuse, scattered or in small groups, dark brown, with masses of conidia on the apex of conidiophores. Mycelium immersed, consisting of pale brown, smooth-walled, branched, septate, smooth, subhyaline hyphae. Conidiophores 131–253 × 4–7 μm ( = 175 × 5 μm, n = 20), macronematous, mononematous, straight or slightly flexuous, solitary or 2–3-group, cylindrical, smooth-walled, 4–6-septate, unbranched, dark brown or brown, becoming paler towards the apex, rounded at the apex. Conidiogenous cells 26–44 × 4–7 μm ( = 35 × 6 μm, n = 20), monophialidic, terminal, integrated, pale brown, subhyaline towards the apex, cylindrical to cylindric-clavate, with an inconspicuous apical collarette. Conidia 8–12 × 6–10 μm ( = 10 × 8 μm, n = 30), acrogenous, solitary, round-tetrahedral, pyramidal, smooth, aseptate, thin-walled, guttulate, with 3-4 radiating from each corner appendages 1.6–3.3 μm ( = 2.4 μm, n = 30) long.

Culture characteristics: Conidia germinating on PDA medium within 48 h and germ tubes arising from terminal end of conidium. Colonies grown on PDA reached 30 mm diam at room temperature in three weeks, irregular, with velvety, dry, raised, grayish-white mycelium on the surface, from below cream to light brown, producing cream pigments, with entire margin.

Material examined: CHINA, Yunnan Province, Tengchong City, Gaoligong Mountain, 25° 12' 36.00" N, 98° 19' 48.00" E, 1852 m a.s.l., on decaying wood submerged in a freshwater stream, 12 November 2024, Y. Li, ly42 (HKAS 149332, holotype), ex-type culture KUNCC 25-19643.

Notes: Phialosporostilbe gaoligongensis possess its conidial characteristics that fit with Phialosporostilbe (Sierra & Portales 1985; Wu & Zhang 2009; Yang et al. 2018b), but differs from P. catenate, P. dendroidea, P. scutiformis, P. setosa, P. turbinata and P. yadongensis in having macronematous, mononematous, unbranched conidiophores. Phialosporostilbe gaoligongensis resembles P. gregariclava in having macronematous, mononematous, conidiophores, acrogenous, solitary, pyramidal conidia, with 3-4 radiating subapical appendages. However, P. gregariclava differs from P. gaoligongensis in having conidiophores of P. gregariclava sometimes with percurrently proliferate, and more septa (5–15 septa vs. 4–6 septa).

Phylogenetic analysis showed that Phialosporostilbe gaoligongensis (KUNCC 25-19643) isolates formed a distinct lineage within the genus, and sister clade with P. scutiformis (MFLU 18-1502 and MFLUCC 18-1288) (Fig. 37). but it differs from P. scutiformis in having unbranched conidiophores and shorter appendages (1.6–3.3 μm vs. 3.5–7.5 μm) (Yang et al. 2018b).

Sporoschisma Berk. & Broome

Sporoschisma was introduced by Berkeley (1847), with S. mirabile as the type species. Significant revisions of the genus were made by Hughes (1966) and Goh et al. (1997), who provided historical collection details and comprehensive illustrations. Sporoschisma is characterized by scattered, capitate setae and straight, smooth, unbranched brown conidiophores, typically consisting of a cylindrical stipe, a swollen venter, and a long cylindrical neck. The conidiogenous cells are phialidic, while the conidia are cylindrical, often truncate at both ends, and form in chains endogenously in basipetal succession (Seifert & Gams 2011; Wu & Diao 2022; Yang et al. 2023; Xu et al. 2024f).

21. Sporoschisma longicatenatum Jing Yang, Jian K. Liu & K.D. Hyde, in Yang, Liu, Hyde, Bhat, Gareth Jones & Liu, Phytotaxa 289(2): 152 (2016) Fig. 40

Chinese name

Index Fungorum number: IF 552182; Facesoffungi number: FoF 02243

Saprobic on decaying wood submerged in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies effuse, black, hairy, chain-like conidia at the apex. Mycelium immersed, composed of pale to dark brown hyphae. Setae 105–156 × 5–7 µm ( = 127 × 6 μm, n = 10), scattered or in groups mixed with conidiophores, capitate, usually at the swollen apex, straight or flexuous, septate. Conidiophores 293–359 × 12–15 µm ( = 318 × 14 μm, n = 15), macronematous, mononematous, smooth, dark brown to black, paler at the torn apex, straight or slightly flexuous, solitary or in groups, setae, arising from dark brown to black bulbous base, composed of a cylindrical stipe and a swollen venter with a long cylindrical neck, erect. Conidiogenous cells monophialidic, percurrent, integrated, terminal, determinate, brown, lageniform, frayed at the apex. Conidia 66–87× 19–25 µm ( = 74 × 21 μm, n = 20), cylindrical, 5-septate, hyaline when young, olivaceous to brown at maturity, with hyaline to pale brown end cells, which are much shorter than the four inner cells, conspicuously darkened at the septa, rounded at both ends.

Culture characteristics: Conidia germinating on PDA medium within 72 h and germ tubes produced from both ends. Colonies grown on PDA reached 60 mm diam at room temperature in two weeks, irregular, with velvety, flat, dark brown mycelium on the surface, from below dark brown with undulate margin.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°19'43" N, 95°21'19" E, 677 m a.s.l., on decaying wood submerged in a freshwater habitat, 13 July 2022, R.J. Xu, MD-109 (HKAS 136183), living culture KUNCC 24-18020.

Fig. 37. Maximum likelihood consensus tree inferred from the combined ITS, LSU, and rpb2 multiple sequence alignments. ML support equal to or greater than 75% and PP value equal to or greater than 0.95 are given near the nodes. The scale bar indicates expected changes per site. The tree is rooted to Helminthosphaeria clavariarum (SMH 4605), Echinosphaeria canescens (SMH 4791), Ruzenia spermoides (SMH 4606) and Synaptospora plumbea (SMH 3962). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Notes: Sporoschisma longicatenatum was identified in a freshwater habitat in Thailand by Yang et al. (2016b), based on phylogenetic analyses and morphological comparisons. Subsequently, Yang et al. (2016a) and Bao et al. (2021) also reported the presence of S. longicatenatum in Thailand. Phylogenetic analyses showed that our isolated of S. longicatenatum (KUNCC 24-18020) clustered together with S. longicatenatum (MFLUCC16-0180 and GZCC15-0072) with strong statistical support (100% ML/1.00 BYPP) (Fig. 39). Therefore, we reported a new record for S. longicatenatum on the Q-X Plateau, China.

Fig. 38. Phialosporostilbe gaoligongensis (HKAS 149332, holotype) a Specimen. b Colonies on woody substrate. c–e Conidiophores. f, g Conidiogenous cell with conidia. h Conidia. i Germinated conidium. j, k Culture on PDA medium, j from above, k from below. Scale bars: c–e = 50 μm, f–i = 10 μm.

Stanjehughesia Subram

Stanjehughesia was established by Subramanian (1992) during his taxonomic revision of the heterogeneous Sporidesmium, in which he segregated several genera based on morphological features such as the presence or absence of conidiophores, the nature of conidial septation (euseptate vs. distoseptate), and percurrent proliferation. Species within Stanjehughesia are characterized by lacking well-developed conidiophores and producing obclavate to obclavate-rostrate, euseptate conidia, which may be smooth or verrucose, directly on simple, monoblastic conidiogenous cells (Subramanian 1992; Wu & Zhuang 2005; Seifert et al. 2011). To date, approximately 21 species have been reported in this genus (Hsieh et al. 2021). Cultural studies have linked the sexual morphs Umbrinosphaeria caesariata syn. Chaetosphaeria caesariata and Miyoshiella larvata linked to Stanjehughesia (Réblová 1999b). Morphologically similar genera such as Linkosia also possess reduced conidiophores and comparable conidial features. However, Linkosia can be distinguished by its distoseptate conidia (Hernández-Gutiérrez & Sutton 1997). Phylogenetic analyses have revealed that both Stanjehughesia and Linkosia as distinct linages in Chaetosphaeriaceae, Chaetosphaeriales (Hyde et al. 2024a; Réblová et al. 2024).

Fig. 39. Maximum likelihood consensus tree inferred from the combined ITS, LSU, and rpb2 multiple sequence alignments. ML support equal to or greater than 75% and PP value equal to or greater than 0.95 are given near the nodes. The scale bar indicates expected changes per site. The tree is rooted to Adautomilanezia caesaloiniae (LAMIC 010212). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

22. Stanjehughesia motuoensis R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 42

Chinese name

Index Fungorum number: IF 905037; Facesoffungi number: FoF 19308

Etymology: the term “motuoensis” refers to the locality where the type specimen was found.

Holotype: HKAS 151645

Saprobic on decaying wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on wood effuse, scattered or in small groups, hairy, dark brown, glistening. Mycelium mostly immersed, composed of septate, branched, hyaline to brown, smooth-walled hyphae. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–6 × 3–4 μm ( = 5 × 3 μm, n = 20), monoblastic, integrated, terminal, determinate, erect, solitary or fasciculate, straight, cylindrical or lageniform, aseptate, slight olivaceous green, brown to dark brown, smooth-walled. Conidia 30–110 × 9–12 μm ( = 90 × 11 μm, n = 20), acrogenous, solitary, lanceolate, rostrate, straight or slightly curved, 14–19-euseptate, brown to dark brown with olive green tint, pale brown to subhyaline at the apex, guttulate, tip swollen and gradually tapers towards the base part when young, the lower middle part swollen and gradually tapers towards both ends when mature, rounded at the apex and truncate at the base, smooth-walled.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°19′43.6″ N, 95°21′19.2″ E, 798 m a.s.l., saprobic on submerged decaying wood in a freshwater habitat, 12 July 2022, R.J. Xu, MD-381A (HKAS 151645, holotype), ex-type culture KUNCC 10488.

Fig. 40. Sporoschisma longicatenatum (HKAS 136183) a, b Colony on wood with long conidia chains. c–e Conidiophores with setae. f, g Conidiogenous cell with conidia. h–k Conidia. l, m Culture on PDA medium, l from above, m from below. Scale bars: c–e = 100 μm, f–k = 20 μm.

Notes: Stanjehughesia motuoensis was collected from decaying wood in a freshwater habitat in Q-X Plateau, China. In the multi-gene phylogenetic analysis based on combined LSU, SSU, ITS, and tef1-α sequence data, S. motuoensis forms a distinct, well-supported clade within Chaetosphaeriaceae (Fig. 41), clearly separated from other known species of the genus (Wu & Diao 2022). Although S. motuoensis shares the typical lanceolate, multi-septate conidia characteristic of Stanjehughesia, it is readily distinguished from S. hormiscioides by its longer conidia (30–110 μm vs. 30–60 μm), higher septation (14–19 vs. 7–14), and more pronounced tapering at both ends. Compared to S. kaohsiungensis, which has shorter (35–70 μm), less septate (7–13) conidia and a broader apex, S. motuoensis also exhibits a distinctive rostrate shape and a darker pigmentation with an olive tint. Stanjehughesia motuoensis is introduced herein as a new species based on its unique morphological characteristics and distinct phylogenetic placement within Stanjehughesia.

Glomerellales Chadef. ex Réblová, W. Gams & Seifert

Reticulascaceae Réblová & W. Gams

Kylindria DiCosmo, S.M. Berch & W.B. Kendr.

DiCosmo et al. (1983) established Kylindria to accommodate Cylindrotrichum triseptatumK. triseptata, type species), along with four other Cylindrotrichum species. The asexual morphs are characterized by solitary, unbranched, multi-septate, dark brown conidiophores, monophialidic, integrated, terminal, ampulliform conidiogenous cells with narrow apertures at the apex and with or without a conspicuous collarette, and unicellular or euseptate, hyaline conidia with or without an excentric lateral or basal hilum (DiCosmo et al. 1983; Zhang et al. 2010). Kylindria species are predominantly saprobic fungi that flourish on decomposing organic matter, including dead branches, decaying leaves, rotting rachides, and desiccated twigs, found in both aquatic and terrestrial ecosystems (Maharachchikumbura et al. 2018; Tian et al. 2024a). Currently, 18 species epithets are listed in Index Fungorum (accessed on 16 July 2025), however, sequence data are available for only 6 species.

Fig. 41. Maximum likelihood consensus tree inferred from the combined ITS, LSU, and rpb2 multiple sequence alignments. ML support equal to or greater than 75% and PP value equal to or greater than 0.95 are given near the nodes. The scale bar indicates expected changes per site. The tree is rooted to Amphisphaeria sorbi (MFLUCC 13-0721) and A. thailandica (MFLU 18-0794). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 42. Stanjehughesia motuoensis (HKAS 151645, holotype) a–c Colony on natural substrates. d Conidiogenous cells with conidia. e–g Conidia. h, i Conidiogenous cells with developmental conidia. j–l Conidia. m Germinated conidium. Scale bars: b–c = 50 μm, d–e = 10 μm, f–i = 5 μm.

23. Kylindria cryptomeriae W.H. Tian, K.D. Hyde & Maharachch., Mycosphere 15 (1): 1861 (2024) Fig. 44

Chinese name

Mycobank number: MB 851891; Facesoffungi number: FoF 16281

Saprobic on decaying submerged wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate, superficial, effuse, gregarious, brown to dark brown, hairy, with white glistening conidial masses at the apex. Mycelium partly immersed, composed of branched, septate, brown to dark brown, smooth hyphae. Conidiophores 204–305 × 9–16 µm ( = 268 × 12 µm, n = 20), mononematous, macronematous, straight to slightly curved, erect, unbranched, multi-septate, dark brown at base, paler toward the apex, thick-walled. Conidiogenous cells 41–53 × 9–15 µm ( = 46 × 13 μm, n=20), phialidic, terminal, integrated, ampuliform, broadest above the middle, tapering near the apex, with an inconspicuous apical collarette. Conidia 24–34 × 9–14 µm ( = 30 × 11 μm, n = 30), arranged in a slimy head, solitary, oblong with obtuse ends, hyaline, 0–3-septate, with a slimy mucilaginous coating around.

Culture characteristics: Conidia germinating on PDA medium within 24 hours, with germ tubes arising from polar cells of the conidium. Colonies grown on PDA reached a diameter of 12 mm at room temperature under natural light in one month, edges regular, flat, white to pale yellow on the surface, pale yellow on the reverse.

Material examined: CHINA, Guizhou Province, Liupanshui City, Shuicheng County, Yushe National Forest Park, N 26°27′17″ N, 104°48′26″ E, 2232 m a.s.l., on decaying submerged wood in a freshwater habitat, 27 February 2023, Xingjuan Xiao, YSS89 (HKAS 145867), culture GZCC 24-0088.

Note: Phylogenetically, our new collection is closely allied with Kylindria cryptomeriae (UESTCC 23.0231) with 96% ML/1.00 BYPP support (Fig. 43). Comparison of the ITS, LSU, and rpb2 sequences between the new collection and K. cryptomeriae (UESTCC 23.0231) showed 99% identity (481/483 bp, 1 gap), 100% identity (801/801 bp), and 99% identity (1019/1023 bp, 1 gap), respectively. Morphologically, the new collection displays nearly identical characteristics, including conidiophore dimensions (204–305 × 9–16 µm vs. 180–310 × 6–11 µm) and conidia (24–34 × 9–14 µm vs. 22–32 × 9–12 µm) with the holotype (HKAS 131279) of K. cryptomeriae (Tian et al. 2024a). Consequently, we classify our collection as K. cryptomeriae, marking the first report of this species from a freshwater habitat.

Conioscyphales Réblová & Seifert

Conioscyphaceae Réblová & Seifert

Conioscypha Höhn

Conioscypha was established by von Höhnel (1904), with C. lignicola as the type species. Species within this genus are widely distributed across terrestrial and freshwater habitats globally. The sexual morphs of Conioscypha are characterized by inconspicuous ascomata, which are either superficial or immersed, ostiolate, and subhyaline to pale orange. These ascomata contain hyaline, 8-spored, unitunicate asci with a refractive J-apical annulus, and fusiform, 3–7-septate ascospores that lack mucilaginous sheaths or appendages (Réblová & Seifert 2004; Zelski et al. 2014; Réblová et al. 2016b; Hyde et al. 2020b). The asexual morphs are defined by micronematous, mononematous, erect, hyaline, and smooth-walled conidiophores. The conidiogenous cells are enteroblastic, terminal, or intercalary, and exhibit cyathiform to doliiform shapes with multilayered, cup-like collarettes. The conidia are dark brown to black, ranging from globose to subglobose or elliptical, with smooth surfaces (Shearer & Motta 1973; Goh & Hyde 1998a; Chuaseeharonnachai et al. 2017; Luo et al. 2019). Currently, Conioscypha comprises 29 accepted species (Index Fungorum 2026; Liu et al. 2019b; Xu et al. 2024e, 2025; Li et al. 2024b; Sun et al. 2025).

24. Conioscypha aquatica (Z.L. Luo, K.D. Hyde & H.Y. Su) R.J. Xu, Q. Zhao & K.D. Hyde, comb. nov. Fig. 46

Chinese name

Index Fungorum number: IF 905262; Facesoffungi number: FoF 05450

Synonym: Conioscypha submersa Z.L. Luo, K.D. Hyde & Hong Y. Su, Fungal Diversity 99: 560 (2019)

Saprobic on decaying wood submerged in a freshwater habitats. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate effuse, scattered, gregarious, punctiform, dark brown or black. Mycelium immersed, composed of hyaline, septate, branched, smooth hyphae. Conidiophores micronematous, mononematous, hyaline, often reduced to conidiogenous cells. Conidiogenous cells sessile or on short conidiophores, and arising directly from the hyphae. Conidia 17–23 × 14–22 μm ( = 20 × 18 μm, n = 40), acrogenous, globose, subglobose, or obovoid, aseptate, pale brown when young, dark brown to black when mature, smooth-walled, subtruncate at base.

Culture characteristics: Conidia germinating on PDA within 48 h and germ tubes produced from base cell. Colonies on PDA reaching 5–10 mm diam, at one month at room temperature, circular, with sparse dark brown mycelium in the middle, sparse pale brown mycelium on the outer ring with irregular margin. Sporulated conidia in cultures. Conidiophores micronematous, branched, sometimes reduced to a moniliform conidiogenous cell, hyaline to pale brown. Conidiogenous cells monoblastic, integrated, cylindrical, hyaline. Conidia 9–17 × 8–14 μm ( =12 × 11 μm, n = 40), acrogenous, subglobose or elliptical, olivaceous or pale brown.

Material examined: CHINA, Xizang Autonomous Region, Rikaze City, Nyalam County, 27° 59' 13.09" N, 85° 59' 0.93" E, 2236 m a.s.l., on decaying wood submerged in a freshwater stream, 06 July 2022, R.J. Xu, MD-680 (HKAS 136019), culture KUNCC 10475. Xizang Autonomous Region, Rikaze City, Dingjie County 27° 53' 8.72" N, 87° 27' 36.21" E, 3042 m a.s.l., on decaying wood submerged in a freshwater stream, 05 July 2022, R.J. Xu, MD-681 (HKAS 136027), culture KUNCC 10473.

Notes: Conioscypha aquatica and C. submersa were initially described by Luo et al. (2019), based on comprehensive phylogenetic analyses and morphological assessments. Our multigene phylogenetic analyses, coupled with meticulous morphological comparisons, indicate that our collected specimens (KUNCC 10475 and KUNCC 10473) group together with ex-type of C. aquatica (MFLUCC 18–1333) and ex-type of C. submersa (DLUCC 0904) (Fig. 45). Despite the observed differences in their ITS sequences, which amount to 7/528 bp (1.3%, excluding gaps), these intraspecific variabilities do not satisfy the standard thresholds commonly employed for species delineation (Jeewon & Hyde 2016; Chethana et al. 2021a). Furthermore, morphological analysis reveals a notable congruence among these specimens regarding conidiophore architecture and the morphology, dimensions, and shape of the spores, thereby reinforcing the hypothesis that they constitute a singular species.

Given these findings, we consolidate Conioscypha aquatica and C. submersa into a single species. In accordance with the International Code of Nomenclature for algae, fungi, and plants (ICN), we recommend the retention of the name C. aquatica, as it is supported by the earlier collection date of its type specimen (2015) in contrast to that of C. submersa (2016). Additionally, we have updated the molecular phylogenetic data for this species and provided a comprehensive description of its colonies cultivated on potato dextrose agar (PDA) medium.

Fig. 43. Maximum likelihood consensus tree inferred from the combined ITS, LSU, and rpb2 multiple sequence alignments. ML support equal to or greater than 75% and PP value equal to or greater than 0.95 are given near the nodes. The scale bar indicates expected changes per site. The tree is rooted to Colletotrichum asianum (LC0037) and C. fructicola (LC0033). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 44. Kylindria cryptomeriae (HKAS 145867, new habitat records) a, b Colonies on dead wood. c–f Conidiophores and conidia. g, h Conidiogenous cells and young conidium. i–l Conidia, m Germinating conidia. n, o Colonies on PDA medium, n from above, o from below. Scale bars: c–f = 50 μm, g–m = 20 μm.

Coniochaetales Huhndorf, A.N. Mill. & F.A. Fernández

Cordanaceae Nann.

Cordana Preuss, Linnaea 24: 129 (1851)

Cordana was established by Preuss (1851) to include three species C. polyseptata, C. pauciseptata, and C. pedunculata, though no type species was initially designated (Hernández-Restrepo et al. 2014; Hyde et al. 2020c). Cordana is defined by macronematous, mononematous conidiophores with polyblastic, inflated, integrate or discrete conidiogenous cells that are terminal or intercalary, nodose, and swollen or umbellate at the apices. Its conidia are acropleurogenous, ovoid to ellipsoid, and typically bear a protuberant hilum (Castañeda Ruiz et al. 1999; Ai et al. 2019; Yang et al. 2022b).

Members of Cordana are globally distributed and commonly found on various substrates, including soil, plant debris, and other fungal organisms (De Hoog et al. 1983; Cai 2004; Soares et al. 2005). Currently, 21 species are recognized in the genus, with some species, such as C. abramovii, C. aquatica, C. reniformis, C. uniseptata and C. xizangensis also identified in freshwater environments (Cai 2004; Hernández-Restrepo et al. 2014; Zelski et al. 2014; Luo et al. 2019; Xu et al. 2024e, 2025).

Fig. 45. Maximum likelihood majority rule consensus tree for Conioscyphaceae using ITS, LSU, SSU and rpb2 sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML BS/PP. The scale bar represents the expected number of changes per site. The tree is rooted with Ascotaiwania sawadae (SS00051). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 46. Conioscypha aquatica (HKAS 136019) a, b Colony on natural substrates. c–e Conidia. f Germinated conidium. g, h Colony on PDA. i–k Conidia on PDA. Scale bars: c–k = 10 μm.

25. Cordana pohuashanensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Fig. 48

Index Fungorum number: IF905263; Facesoffungi number: FoF 19309

Chinese name

Etymology: Name reflects the host, Sorbus pohuashanensis, from which the fungus was collected.

Holotype: HKAS 144626

Saprobic on decaying of Sorbus pohuashanensis (Rosaceae). Sexual morph: Not observed. Asexual morph: Hyphomycetous. Colonies on the host surface, effuse, medium sparse, dark brown. Mycelium immersed, septate, branched, smooth. Conidiophores macronematous, mononematous, erect, straight or slightly flexuous, branched, rough-walled, thick-walled, septate, cylindrical, 0-10–inflated, brown to hyaline, conidiophores have multiple conidiogenous loci, 51–260 × 4–7.7 μm ( = 160 × 6 μm, n = 20). Conidiogenous cell polyblastic, integrated, terminal. Conidia didymospores, solitary, acropleurogenous, simple, dry, oblong with obtuse ends, smooth, thin-walled, 1-septate, guttulate when immature, constricted at septate, darker at septum hyaline, hyaline to brown, 8.7–12 × 3.8–5.7 μm ( = 10.5 × 5 μm, n = 30).

Culture characteristics: Conidia germinating within 12 h on PDA media at 25 °C, reaching 2.2–2.4 cm after 15 days incubation, colony surface pale olivaceous-grey, reverse pale black, dense, circular, slightly raised, smooth, entire, hairy, pigmentation not produced.

Material examined: CHINA, Xizang, Linzhi City, Motuo County, 29°15’N, 95°15′E, 1663 m a.s.l., on the Sorbus pohuashanensis (Rosaceae), June 27, 2022, collected by Yun-Wei Zhao, ZYW221 (HKAS 144626, holotype), ex-type culture, KUNCC24-19008.

Notes: Cordana aquatica was introduced by Luo et al. (2019), based on phylogenetic and morphological analyses, and was collected from aquatic environments in Yunnan, China. Our specimen (HKAS 144626) was collected in Xizang, China, from a terrestrial environment of Sorbus pohuashanensis. Morphologically, our specimen (HKAS 144626) are similar to C. aquatica, with conidiophores that are macronematous, mononematous, and cylindrical, and conidia that are didymospores, acropleurogenous, and 1-septate (Luo et al. 2019). However, the conidiophores of our samples are shorter (160 × 6 μm vs. 209 × 6 μm), and the conidia are larger (10.5 × 5 μm vs. 9 × 4 μm). Through base pair comparison, the ITS and LSU sequence of C. aquatica differs from that of our specimen (HKAS 144626) in 17/482 (3.5 %) and 7/759 (1 %) respectively, and phylogenetic analysis showed that C. pohuashanensis sister group with C. reniformis (Fig. 47). Therefore, we introduce a new species, C. pohuashanensis, from Xizang, China, based on phylogenetic analysis, morphology, and habitat.

Fig. 47. Maximum likelihood majority rule consensus tree for Cordanaceae using LSU and ITS sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Humicola limonispora (LC 5707) and Teracosphaeria petroica (ICMP 15111). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 48. Cordana pohuashanensis (HKAS 144626, holotype) a Host. b Colonies on the substrate surface. c-g Conidiophores. h, i Conidiogenous cell. j Conidium. k Germinated conidia. l-m Culture on PDA. Scale bars: c-g = 50μm, h-k = 10 μm.

26. Cordana tumida R.J. Xu, Q. Zhao & K.D. Hyde, sp. nov. Fig. 49

Chinese name:

Index Fungorum number: IF 903488; Facesoffungi number: FoF 17412

Etymology: The epithet "tumida" refers to the swollen conidiophores.

Holotype: HKAS 136033

Saprobic on decaying wood submerged in a freshwater stream. Sexual morph: Undetermined. Asexual morph: Colonies superficial, effuse, scattered, hairy, dark brown. Mycelium immersed, consisting of pale brown, smooth-walled, branched, septate. Conidiophores 197–357 × 4–6 μm ( = 273 × 5 μm, n = 10), macronematous, mononematous, solitary, erect, straight or flexuous, unbranched, multi-septate, with swollen, fertile, intercalary nodes in the upper middle, cylindrical, slightly constricted at septa, dark brown at the base, lighter towards the apex. Conidiogenous cells 30–66 × 5–7 μm ( = 36 × 6 μm, n = 20), polyblastic, integrated, terminal, becoming intercalary, percurrent, sympodial, denticulate, usually swollen at the conidiogenous loci. Conidia 13–17 × 7–10 μm ( = 14 × 9 μm, n = 30), acropleurogenous, calabash-shaped, or cylindrical, smooth, 1-septate, pale brown, apparent constricted at the septa, elliptical at the apex, rounded at the base, guttulate.

Culture characteristics: Conidia germinating on PDA medium within 48 h and germ tubes arising from terminal end of conidium. Colonies grown on PDA reached 30 mm diam at room temperature in three weeks, circular, with velvety, flat, grayish-white mycelium on the surface, from below cream with entire margin.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°19'36.31" N, 95°21'22.18" E, 1481 m a.s.l., on decaying wood submerged in a freshwater stream, 13 July 2022, R.J. Xu, MD-676 (HKAS 136033, holotype), ex-type culture KUNCC 10467. Linzhi City, Chayu County, 28°29'39.27" N, 96°59'35.25" E, 1537 m a.s.l., on decaying wood submerged in a freshwater stream, 14 July 2022, R.J. Xu, MD-676-2 (HKAS 136032, paratype), culture KUNCC 10468.

Notes: Among the known species of Cordana, only C. aquatica, C. lignicola, C. lushanensis and C. verruculose resemble C. tumida in having intercalary nodes (Hernández-Restrepo et al. 2014; Ai et al. 2019; Luo et al. 2019). However, C. aquatica differs from C. tumida by its smaller conidia (8–10 × 3–5 μm vs. 13–17 × 7–10 μm), and mostly oblong, sometimes ellipsoid conidia (Luo et al. 2019). Cordana lignicola differs from C. tumida by its narrower conidia (3.5–4.5 μm vs. 7–10 μm). Cordana lushanensis differs from C. tumida by its branched conidiophores and aseptate conidia (Ai et al. 2019). Cordana verruculose differs from C. tumida by its aseptate and verruculose conidia (Hernández-Restrepo et al. 2014).

In the phylogenetic analysis, Cordana tumida was grouped with C. linzhiensis and C. crassa forming a distinct clade separate from other species within Cordana (Fig. 47). However, C. tumida differs from C. linzhiensis in having smaller conidia (13–17 × 7–10 vs. 20–28 × 11–15 μm) and in the presence or absence of intercalary nodes in the conidiophores. Additionally, the conidia of C. linzhiensis are characterized by a hyaline mucoid sheath (Xu et al. 2024e). By contrast, C. crassa differs from C. tumida in that the conidiophores of the former absence intercalary nodes. In addition, a comparison of the ITS nucleotide bases in C. tumida (KUNCC 10467) and C. crassa (KUNCC 24-17964) revealed that they differ in 10/491 bp (2%) of ITS. Therefore, the new species C. tumida is introduced based on the recommendation of Jeewon & Hyde (2016) and the polyphasic approach recommended by (Chethana et al. 2021a).

Diaporthales Nannf

Lamproconiaceae Norph., T.C. Wen & K.D. Hyde

Lamproconiaceae was established to accommodate Lamproconium and Hercospora by Norphanphoun et al. (2016) based on both morphological and phylogenetic evidence. This family includes both pathogenic and saprobic fungi, typically found on dead herbaceous twigs (Norphanphoun et al. 2016; Crous et al. 2020). The sexual morph is characterized by prosenchymatous stromata surrounding the perithecia, externally demarcated by a greenish-black, dense pseudoparenchymatous layer, while the interior is whitish, composed of interwoven hyphae intermixed with host tissue, each stroma generally contains 3–5 perithecia. The asexual morph produces pycnidial conidiomata that are solitary, partially embedded in host tissues, and either uniloculate, multiloculate, or convoluted, conidiogenous cells are holoblastic, cylindrical to subcylindrical, forming single conidia at the apex, or annellidic, and may appear colorless to olivaceous with smooth walls, conidia are aseptate, fusiform to ellipsoid, thick-walled, containing granular contents (Norphanphoun et al. 2016). Currently this family contains 27 epithets in three genera, viz., Hercospora, Lamproconium, and Neolamproconium (Index Fungorum 2025; Hyde et al. 2024a).

27. Zangmuomyces R.J. Xu, K.D. Hyde & Q. Zhao, gen. nov.

Chinese name

Index Fungorum number: IF 905264; Facesoffungi number: FoF 19310

Etymology: The genus name Zangmuomyces is established to honor Professor Zang Mu for his outstanding contributions to the field of mycology in China.

Saprobic on decaying unidentified wood in terrestrial habitat. Sexual morph: Undetermined. Asexual morph: Conidiomata 378–481 µm high, 660–858 µm diam ( =428 × 740 μm, n = 15), pycnidial, solitary, scattered, semi-immersed or superficial, black, uni-loculate, ellipsoid or irregular when viewed from above, like a raised cylindrical barrel when viewed from the front, a large opening in the middle, glabrous. Peridium 69–120 μm wide, composed of brown to dark brown, irregular, thick-walled cells arranged in a textura angularis, the side walls are significantly thicker than the bottom, fused with the host tissue. Paraphyses interspersed within conidiophores. Conidiophores 15–78 × 2–4 μm ( = 47 × 3 μm, n = 20), arising from the outermost wall layer at the basal of pycnidium, filiform or cylindrical, hyaline, aseptate, unbranched, smooth-walled. Conidiogenous cells cylindrical to subcylindrical, filiform, hyaline, thin-walled, becoming swollen towards the apex. Conidia 26–35 × 10–13 μm ( = 31 × 12 μm, n = 25), hyaline, ellipsoidal to long ellipsoidal, rounded at both ends, straight or slightly curved, 3-septate, slightly constricted at septum, hyaline sheath around conidia, smooth-walled, guttulate.

Fig. 49. Cordana tumida (HKAS 136033, holotype) a, b Colony on natural substrates. c–e Conidiophores with conidia. f–h Conidiogenous cells with conidia. i Swollen, intercalary conidiogenous cells. j–l Conidia. m, o Culture on PDA medium. m from above, o from below. Scale bars: c–e = 50 μm, f–i = 20 μm, j–l = 10 μm.

Type species: Zangmuomyces xizangensis R.J. Xu, K.D. Hyde & Q. Zhao (HKAS 151647)

Notes: In the phylogenetic analyses, strains of Zangmuomyces were clustered with Hercospora, Lamproconium, and Neolamproconium in the Lamproconiaceae. In morphology, Z. xizangensis shows unique features: conidiomata like a raised cylindrical barrel when viewed from the front, conidiogenous cells becoming swollen towards apex, and conidia ellipsoidal to long ellipsoidal, rounded at both ends, 3-septate. It can be distinguished from other three genera in Lamproconiaceae. Therefore, this study introduces Zangmuomyces as a new genus to accommodate Z. xizangensis.

28. Zangmuomyces xizangensis R.J. Xu, K.D. Hyde & Q. Zhao, sp. nov. Fig. 51

Chinese name

Index Fungorum number: IF 905043; Facesoffungi number: FoF 19311

Etymology: refers to Xizang Autonomous Region in China, where the type specimen was collected.

Holotype: HKAS 151647

Saprobic on decaying unidentified wood in a freshwater habitat. Sexual morph: Undetermined. Asexual morph: Conidiomata 378–481 µm high, 660–858 µm diam ( = 428 × 740 μm, n = 15), pycnidial, solitary, scattered, semi-immersed or superficial, black, uni-loculate, ellipsoid or irregular when viewed from above, like a raised cylindrical barrel when viewed from the front, a large opening in the middle, glabrous. Peridium 69–120 μm wide, composed of brown to dark brown, irregular, thick-walled cells arranged in a textura angularis, the side walls are significantly thicker than the bottom, fused with the host tissue. Paraphyses filamentous, aseptate, interspersed within conidiophores. Conidiophores 15–78 × 2–4 μm ( = 47 × 3 μm, n = 20), arising from the outermost wall layer at the basal of pycnidium, filiform or cylindrical, hyaline, aseptate, unbranched, smooth-walled. Conidiogenous cells cylindrical to subcylindrical, filiform, hyaline, guttulate, thin-walled, becoming swollen towards apex. Conidia 26–35 × 10–13 μm ( = 31 × 12 μm, n = 25), hyaline, ellipsoidal to long ellipsoidal, rounded at both ends, straight or slightly curved, 3-septate, slightly constricted at septum, hyaline sheath around conidia, smooth-walled, guttulate.

Material examined: CHINA, Xizang Autonomous Region, Linzhi City, Motuo County, 29°19′43.69″ N, 95°21′19.27″ E, 677 m a.s.l., on decaying wood submerged in a freshwater stream, 13 July 2022, R.J. Xu, MD-308A (HKAS 151647, holotype), ex-type culture KUNCC 25-21370.

Notes: In the phylogenetic tree, Zangmuomyces xizangensis clustered with strains of Hercospora, Lamproconium, and Neolamproconium with 99% ML/1.00 BYPP statistical support (Fig. 50). Morphologically, Zangmuomyces and other three genera of Lamproconiaceae show differences on conidiomata, conidiogenous cells, and conidia. The conidiomata of Z. xizangensis are like a raised cylindrical barrel when viewed from the front, conidiogenous cells become swollen towards the apex, and conidia are 3-septate, ellipsoidal to long ellipsoidal, often with a hyaline sheath, and rounded at both ends, while Hercospora has conidiomata with a superficial cap of sterile tissues, elongate conidiophores (conidiogenous cells not mentioned), and ovoid to ellipsoid, one-celled conidia (Norphanphoun et al. 2016). Lamproconium has dark blue conidiomata, with a raised centre, conidiogenous cells with flared periclinal thickenings in the collarette zone, and conidia are fusiform, ellipsoid, with narrowly rounded ends, aseptate, bluish to glistening dark blue (Norphanphoun et al. 2016). The stromata of Neolamproconium are characterized by aggregated rosettes of up to 15 pycnidia, phialidic conidiogenous cells, with prominent periclinal thickenings, and conidia are dimorphic, with macroconidia 1(–3)-septate, and microconidia aseptate (Crous et al. 2020).

Therefore, based on its obvious branching in the phylogenetic tree and unique morphological structure, this study introduces Zangmuomyces xizangensis as a new species of a new genus Zangmuomyces.

Hypocreales Lindau

Bionectriaceae Samuels & Rossman

Clonostachys

The asexual morph-typified genus Clonostachys, described by Corda (1839), is anchored by its type species, Clo. araucaria, distinguished by its penicillate conidiophores and imbricate conidia organized in columns (Schroers et al. 1999). Clonostachys is typified by its penicillate, sporodochial, or dimorphic conidiophores and phialidic conidiogenous cells, which generate hyaline conidia (Schroers 2001). The teleomorph, originally described as Bionectria (Spegazzini 1919), is characterized by ascomata typically situated on a pseudoparenchymatous stroma or directly arising from the substrate, ascomata exhibit a color range from white, pale yellow, to orange and dark brownish-orange, remaining unchanged in appearance when exposed to 3% KOH or lactic acid (Schroers 2001; Lechat & Fournier 2018; Perera et al. 2023). Currently, 136 records have been listed in the Index Fungorum (Index Fungorum Name Search, April 18, 2026).

29. Clonostachys habaensis Y.B. Wang, B.Z. Chen & Zhu L. Yang, sp. nov. Fig. 53

Chinese name

Index Fungorum number: IF 905265; Facesoffungi number: FoF 19312

Etymology: The epithet habaensis refers to Haba Snow Mountain (Haba Xueshan) in Shangri-La City, Yunnan Province, China, where the type specimen was collected.

Diagnosis: Clonostachys habaensis produces slightly smaller ellipsoidal conidia compared to its close relatives.

Holotype: HKAS 135029

Endophytic from the root of Rhododendron decorum. Sexual morph: Undetermined. Asexual morph: Conidiophores broadly penicillate, arising from the agar surface or aerial mycelium ter- to quinquiesverticillate, aggregated in pustules or sporodochia, phialides slightly divergent or adpressed, in whorls of 1–3, flask-shaped, widest in the lower third or almost cylindrical, slightly tapering in the upper part towards the tip, without a visible collarette, (11.3–) 11.4–18.9 (–19.2) × (1.9–) 2–3.2 (3.2) μm ( 15.5 × 2.7 μm, n = 20). Conidia aseptate, hyaline, smooth-walled, ellipsoid, some slightly curved or asymmetric with one more flattened side, with a laterally displaced hilum, 5.1–7.6 (–9) × (2.9–) 3–5.2 (–5.8) μm ( 6.3 × 4 μm, n = 30).

Fig. 50. Maximum likelihood majority rule consensus tree for Diaporthales using ITS and LSU sequence data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities greater than 0.95 are indicated near branches as ML/BYPP. The scale bar represents the expected number of changes per site. The tree is rooted with Magnaporthe salvinii (CBS 243.76) and M. grisea (GAD1). Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 51. Zangmuomyces xizangensis (HKAS 151647, holotype) a–c Conidiomata on host. c–e Conidiophores with conidia. d, e Cross section of a conidioma. f–i Conidiogenous cells with attached conidia. j–m Conidia. n Germinated conidium. Scale bars: f–i = 20 μm, j–n = 10 μm.

Culture characteristics: Colonies on PDA reaching 54–61 mm diameter after 8 days at 25 °C, flat, with entire margin, aerial mycelium abundant, cottony, pale yellow, reverse yellowish.

Material examined: CHINA, Yunnan Province, Diqing, Shangri-La City, Haba Snow Mountain, isolated as an endophyte from Rhododendron decorum root, 15 March 2023, collected by Q. Fan (HKAS 135029, holotype, dried culture on PDA, ex-type culture KUNCC 11234), other collection: Ibid., Q. Fan (culture KUNCC 11233).

Notes: Clonostachys habaensis is closely related to Clo. pseudostriata and Clo. palmae, as strongly supported by multi-gene phylogenetic inference. However, it shows clear sequence divergence from its sister species Clo. pseudostriata (CBS 120.87) (Fig. 52), with 99.4% identity in ITS (2 bp differences), 99.8% in LSU (1 bp), 98.7% in rpb2 (9 bp), 98.4% in tef1-α (12 bp), and 95.6% in tub (39 bp). Morphologically, Clo. habaensis produces ellipsoidal conidia that are slightly smaller than those of Clo. palmae (Schroers 2002; Zhao et al. 2023).

Fig. 52. The maximum likelihood tree using IQ-Tree (Web-IQ-TREE) inferred from combined LSU, tef1-α, ITS, tub and rpb2 sequences. Stephanonectria chromolaenae R.H. Perera & K.D. Hyde and Ste. keithii (Berk. & Broome) Schroers & Samuels were designated as the outgroup taxa. Values at the nodes before and after the backslash are ML bootstrap proportions (ML-BP> 70%) and BI posterior probabilities (BI-PP> 0.70), respectively. Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Clavicipitaceae Rogerson

Clavicipitaceae was initially treated as a subfamily of Hypocreaceae (Earle 1901). It was later placed within Hypocreales by Spatafora & Blackwell (1993). Sung et al. (2007) further divided it into three monophyletic families: Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae. Xiao et al. (2023) introduced a fourth family, Polycephalomycetaceae. Currently, Clavicipitaceae includes over 40 genera, such as Aciculosporium, Claviceps, Epichloë, and Metarhizium (Kepler et al. 2014). Many species are entomopathogenic, often parasitizing insects or forming symbiotic relationships with plants (Kepler et al. 2012; Khonsanit et al. 2021). The sexual morph features dark or brightly colored, fleshy to tough stromata or subiculum, with perithecial ascomata that can be superficial or immersed and are arranged in ordinal or oblique patterns, containing cylindrical asci with thickened apices and typically cylindrical, multiseptate ascospores that may disarticulate into part-spores (Sung et al. 2007; Kepler et al. 2014). The asexual morphs include genera like Aschersonia, Ephelis, Metarhiziopsis, Metarhizium, Neotyphodium, Nomuraea, Pochonia, and Rotiferophthora (Sung et al. 2007; Kepler et al. 2012; Mongkolsamrit et al. 2020).

Fig. 53. Clonostachys habaensis (HKAS 135029, holotype) a, b Colonies on PDA, front and back. c, d Conidiophores and Phialides. e–g conidia. Scale bar: a = 1 cm, c–g = 5 μm.

Metarhizium Sorokin

Metarhizium is of paramount significance in the study of entomopathogenic fungi, catalyzing two molecular phylogenetic revisions that have robustly validated and refined the morphological classification delineated by Rombach et al. (1987). The species classified within Metarhizium are acknowledged as entomopathogens, effectively targeting insect hosts and resulting in their mortality or incapacitation. Furthermore, these fungi have been documented to promote plant growth and enhance biomass accumulation (Elena et al, 2011; Khan et al, 2012; Mongkolsamrit et al. 2020).

30. Metarhizium luteum Y.B. Wang, C.Y. Wei, B.Z. Chen & Zhu L. Yang, sp. nov. Fig. 54

Chinese name:

Index Fungorum number: IF 905266; Facesoffungi number: FoF 19313

Etymology: The stromata are earthy yellow.

Diagnosis: Different from all other Metarhizium members by color of its stromata.

Holotype: HKAS 126260

Parasitic on larvae of Scarabaeidae. Sexual morph: Stromata from the head of the Scarabaeidae (Coleoptera) buried in soil, solitary, occasionally branched, fleshy, glabrous, 7–10 cm long. Stipes cylindrical, easily bent, white to orange, 2–4 mm wide. Fertile parts terminal, cylindrical, orange, tapering upwards, contains infertile tip, 70–100 × 2–4 mm. Perithecia obliquely immersed, dense, ovate, 600–800 × 100–400 µm. Asci linear, (100–) 150–300 (–350) × 3.5–5 (–5.5) µm ( 207.6 × 4.3 µm), apical caps prominent, hemispherical, (3–) 3.5–6 (–6.5) × 1.5–2 (–2.5) µm ( = 4.9 × 1.9 µm). Ascospores hyaline, filiform multiseptate, breaking into secondary ascospores, 15–25 (–30) × (0.5–) 1–1.5 µm ( 22.5 × 1.2 µm) Secondary ascospores cylindrical, rounded at both ends, (8.5-) 8.6 - 10.4 (-10.5) × 1 - 1.5 µm (x = 9.5 × 1.1 µm). Asexual morph: Conidiophores smooth-walled, cylindrical, apically branched. Phialides smooth-walled, cylindrical with semi-papillate apices, arranged in whorls of 3–5 at the apex of conidiophores, (6–) 6.5–10.5 (–11) × (1–) 1.5–3 (–3.5) µm ( 8.3 × 2.3 µm). Conidia hyaline, cylindrical, (6.5–)7–10(–10.5) × (2–)2.5–3.5(–4.0 μm ( = 8.3 × 2.9 μm).

Culture characteristics: Colonies on PDA attaining a diam of 23 mm in 14 d, white, flocculent, high mycelial density, intact edges, reverse white cream. Center of the colony is dark green due to the production of conidia. Hyphae smooth-walled, colorless, septate.

Material examined: CHINA, Yunnan Province, Baoshan City, parasite on larvae of Scarabaeidae buried in soil, 2 August 2022, C.Y. Wei, (HKAS 126260 holotype, ex-type culture KUNCC 10831), other collection: Ibid., C.Y. Wei (HKAS 126261, culture KUNCC 10832; HKAS 126262, culture KUNCC 10833).

Notes: As shown in the phylogenetic tree, Met. luteum forms a distinct terminal lineage (Fig. 57), strongly supported by multi-locus phylogenetic analyses. Metarhizium luteum shares similarity with Met. candelabrum and Met. cercopidarum in the production of cylindrical phialides with rounded ends (Mongkolsamrit et al. 2020). However, Met. luteum parasitize scarabaeidae larvae while Met. candelabrum and Met. cercopidarum infect audlt leafhopper. In Met. luteum the conidia on PDA are wider than in Met. candelabrum and Met. cercopidarum. Further, Met. luteum is distinguished from Met. candelabrum and Met. cercopidarum by growth rate on PDA, and Met. luteum is significantly faster than Met. candelabrum and Met. cercopidarum.

Moelleriella Bres

Moelleriella, an entomopathogenic genus within the Clavicipitaceae (Hypocreales, Ascomycota), is distinguished from related genera like Conoideocrella, Hypocrella and Samuelsia by unique features such as disarticulating ascospores and fusiform conidia produced by its anamorph (Bresadola 1897; Chaverri et al. 2008). Moelleriella species engage in a dual parasitic role, acting as necrotrophic parasites of scale insects and whiteflies, and biotrophic parasites of plants, often presenting as vibrant stromata on insect hosts (Chaverri et al. 2008; Mongkolsamrit et al. 2015). Molecular analyses, especially of LSU, tef1-α and rpb1 genes, have clarified their taxonomy, ecological roles, and evolutionary relationships, enhancing our understanding of fungal biodiversity and ecosystem dynamics (Khonsanit et al. 2021).

31. Moelleriella yusheensis Y. Yang, K.D. Hyde & Y.P. Xiao, sp. nov. Fig. 56

Chinese name

Index Fungorum number: IF 903156; Facesoffungi number: FoF 17061

Etymology: in reference to the place where the fungus was collected, China, Guizhou Province, Liupanshui City, Shuicheng County, Yushe forest Park.

Holotype: HKAS 132266

Sexual morph: ( = 296 × 110 µm, n = 30). Ostioles pale yellow to light orange. Asci hyaline, cylindrical, 130–180 × 5–11 μm ( = 162 × 7 µm, n = 50). Asci caps inconspicuous. Ascospores hyaline, disarticulating into part-spores. Secondary spores oblong to cylindrical with round or blunt ends, 7.2–9.7 × 2.3–3.5 μm ( = 8.6 × 2.9 µm, n = 50), smooth, hyaline, thin-walled, guttulate. Asexual morph: undetermined.

Material examined: CHINA, Guizhou Province, Liupanshui City, Shuicheng County, Yushe forest Park. Parasitism on scale insects (Coccidae), found on the upper surfaces of dicotyledonous leaves, 1 August 2022, Yu Yang SC186 (HKAS 132266, holotype).

Notes: Moelleriella yusheensis clusters with Moe. gracilispora and Moe. kanchanaburiensis in the phylogenetic tree, with 100% MLBP, 0.99 BYPP support (Fig. 55). The sequences of six loci of Moe. gracilispora and Moe. kanchanaburiensis share 97–98% identity with Moe. yusheensis in LSU, 94–95% in tef1-α, and 91–92% in rpb1. Moe. gracilispora infects whitefly nymphs (Hemiptera), while Moe. yusheensis and Moe. kanchanaburiensis both infect scale insects (Coccidae) (Yuan et al. 2020; Khonsanit et al. 2021). Morphologically, Moe. yusheensis can be easily distinguished from other Moelleriella species by its flat to umbonate, orange to dark orange teleomorphic stromata, numerous embedded perithecia, hyaline asci, and cylindrical thin-walled, and guttulate secondary spores. In this study, we introduce a novel species, Moe. yusheensis, from China.

Papiliomyces Luangsa-ard, Samson & Thanakitp. 2020

Papiliomyces was introduced by Mongkolsamrit et al. (2020) within the Clavicipiteae, to accommodate Pap. liangshanensis and Pap. shibinensis. The teleomorph of Papiliomyces encompass solitary to multiple (branched and robust to wiry) stromata, superficial to entirely immersed perithecia within the stroma, and ampulliform, ellipsoid to ovoid asci, with ascospores that are whole, exhibiting septation or fragmentation into cylindrical part-spores (Kepler et al. 2012a; Kepler et al. 2012b; Mongkolsamrit et al. 2020; Wang et al. 2021).

Fig. 54. Metarhizium luteum (HKAS 126260, holotype) a Stroma arising from a larva of Coleoptera. b Fertile part. c Perithecia. d, e Colonies on PDA. f–i Asci. j Ascospore. k, l Secondary ascospores. m–q Conidiophores, phialides and conidia on PDA. r Conidia on PDA. Scale bars: a = 1 cm, b = 1 mm, c = 200 μm, d, e = 2 cm, f–h = 20 μm, i–r = 10 μm.

32. Papiliomyces aurantiacus Y.B. Wang, C.Y. Wei, B.Z. Chen & Zhu L. Yang, sp. nov.Fig. 58

Chinese name:

Index Fungorum number: IF 905267; Facesoffungi number: FoF 19314

Etymology: Fertile parts are orange

Diagnosis: Different from all other Papiliomyces members by color of its fertile parts.

Holotype: HKAS 126283

Parasitic on larvae of Scarabaeidae. Sexual morph: Stromata from the head of lepidopteran larvae buried in the soil, solitary, occasionally branched, 30–45 mm long. Stipe cylindrical, white to orange-yellow, 1.5–3 mm wide. Fertile parts cylindrical, orange, 5–10 × 2.5–3 mm. Perithecia embedded, dense, ovate or long oval, 445–503 × 176–239 µm ( 465.5 × 214.7 µm). Asci transparent, linear, 114–202 × 4.8–6.7 µm ( 152.7 × 5.5 µm), apical caps prominent, hemispherical, (4–) 4.5–5.5 (–6) × 2.5–3.5 (–4) µm ( 4.9 × 3 µm). Ascospores columnar, multiseptate, composed of numerous cells, 162–175 × 2.4–3.2 µm ( 168.8 × 2.8 µm), not breaking into secondary ascospores. Asexual morph: Conidiophores smooth-walled, cylindrical, top branch. Phialides smooth-walled, colorless, flask-shaped, base thinning upward, arising singly from the hyphae or in whorls of 3-4 at the top of the conidiophores, (11–) 11.5–15 (–15.5) × (0.5–) 1–2 (–2.5) µm ( 13.4 × 1.6 µm). Conidia smooth-walled, unicellular, globose to subglobose, surface tooth teeth, often arranged into chains, and 3.5–4.5 (–5) × 3.5–4 (–4.5) µm ( 4.2 × 4.1 µm).

Culture characteristics: Colonies on PDA attaining a diam of 30 mm in 14 d, dense, pulvinate, white, intact edges. Hyphae smooth-walled, colorless.

Material examined: CHINA, Yunnan Province, Nujiang, parasite on larvae of Hepialidae buried in soil. 12 August 2023, C.Y. Wei (HKAS 126283 holotype, ex-type culture KUNCC 10852), other collection: Ibid., C.Y. Wei (HKAS 126282).

Fig. 55. Maximum likelihood consensus tree inferred from the combined LSU, tef1-α, and rpb1 multiple sequence alignments. Bootstrap support values for maximum likelihood (ML, first value) equal to or greater than 75% and Bayesian posterior probabilities from MCMC analyses (BYPP, second value) equal to or greater than 0.90 are given above the nodes. The scale bar indicates expected changes per site. The tree is rooted to Ophiocordyceps aphrophoridarum YTS19061912 and O. aphrophoridarum YTS19061907. Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 56. Moelleriella yusheensis (HKAS 132266, holotype) a Habitat. b-e Telemorphic stroma. f Perithecia. g-i Asci. j Apical cap of asci. k part of Ascospores. l Secondary spores. Scale bars: d = 0.1 cm, e = 0.5 mm, f = 200 μm, g–i = 50 μm, j–k = 20 μm, l = 10 μm.

Notes: As shown in the phylogenetic tree, Pap. aurantiacus represents a distinct terminal lineage (Fig. 57), strongly supported by multi-locus phylogenetic analyses. Papiliomyces aurantiacus exhibits similarities to Pap. shibinensis, as both species share the same host, namely the larvae of Lepidoptera. However, they differ in the size and

color of fertile parts and ascopores. Papiliomyces aurantiacus has orange fertile parts, whereas Pap. shibinensis is faint yellow (Mongkolsamrit et al. 2020). In Pap.aurantiacus (162–175 × 2.4–3.2 µm), the fertile parts are wider than in Pap. shibinensis (120–170 × 1.4–2.1 μm).

Fig. 57. The maximum likelihood tree using IQ-Tree (Web-IQ-TREE) inferred from combined LSU, SSU, tef1-α, rpb1 and rpb2 sequences. Pleurocordyceps sinensis (Q.T. Chen, S.R. Xiao & Z.Y. Shi) Y.J. Yao, Y.H. Wang, S. Ban, W.J. Wang, Yi Li, Ke Wang & P.M. Kirk was designated as outgroup taxon. Values at the nodes before and after the backslash are ML bootstrap proportions (ML-BP> 70%) and BI posterior probabilities (BI-PP> 0.70), respectively. Sample labels of the newly obtained sequences are provided in the order of herbarium no., province name, and county/city name.

Fig. 58. Papiliomyces aurantiacus (HKAS 126283, holotype) a, b Stromata arising from lepidopteran larvae. c Fertile parts, d Colonies on PDA. e Perithecia. f–i Asci. j, k Ascospores. l–o Conidiophores, phialides and conidia on PDA. Scale bars: a–c = 1 cm, d = 2 cm, e, f = 100 μm, g, h = 20 μm, i = 10 μm, j = 20 μm, k–o = 10 μm.

Petchia Thanakitp., Mongkols. & Luangsa-ard 2020

Petchia was proposed by Mongkolsamrit et al. (2020) within the Clavicipiteae. Petchia is distinguished from related genera by stromata arising from the egg cases of praying mantis (Mantidae), multiple, mostly erect up to 2 cm high, fertile region is located at the apical end of the stroma, exhibiting a pale cream coloration, with a globose morphology measuring 1.5–1.8 cm in diameter, perithecia are narrowly ovoid, exhibiting a brown hue, and are immersed with dense mycelial encasement around each perithecium (Mongkolsamrit et al. 2020). Currently, only two species in the genus, all are saprotrophic, on the egg cases of praying mantis (Mongkolsamrit et al. 2020).

33. Petchia siamensis Thanakitp., Mongkols. & Luangsa-ard, in Thanakitpipattana, Tasanathai, Mongkolsamrit, Khonsanit, Lamlertthon & Luangsa-ard, Persoonia 44: 154 (2020) Fig. 59

Chinese name:

Index Fungorum number: IF 830168; Facesoffungi number: FoF 14494

Diagnosis: The specimen examined in this study shares the same phylogenetic position as Petchia siamensis.

Parasitic on the ootheca of praying mantis (Mantidae). Sexual morph: Stromata grow from the ootheca of praying mantis (Mantidae), several produced on a single host, each stroma unbranched, easily bent, length 5-7 mm. Stipe cylindrical, curved, smooth, glabrous, light yellow, with a diameter of 0.8-1 mm. The fertile part is globose apical, light yellow, measuring 2 × 1.5 mm. Perithecia are immersed, oval, brown, (550–)578–624(–650) × (400–)413–434(–450) µm, ( 596 × 434 µm). Asci are cylindrical, slender, (170–)179–250 × 3–4.8(–5) µm ( 203 × 4 µm). The apical cap is prominent, hemispherical, thickened, measuring 3–3.5 × 1.5–2 µm, ( 3.1 × 1.8 µm). Ascospores are transparent, thread-like, multi-septate, (200–)215–275(–300) × 1–1.5 µm, ( 246 × 1.3 µm), and break into secondary ascospores when mature. Secondary ascospores are short-cylindrical, (5.5–)5.7–9.8(–10) × (0.5–)0.7–1.2(–1.5) µm, ( 8.1 × 0.9 µm). Asexual morph: Hyphae smooth-walled, colorless, branched. Conidiophores smooth-walled, cylindrical, branched at the apex. Phialides smooth-walled, base cylindrical and gradually tapering upwards, bottle-shaped, whorled at the top of the conidiophores, 12.5–18.8(–20) × (1–)1.3–3.3(–3.5) µm, ( 14.9 × 1.9 µm). Conidia unicellular, colorless, ellipsoidal, 3–5 × (1.5–)1.7–2.8(–3) µm, ( 4.4 × 2.3 µm).

Culture characteristics: Colonies on PDA reach 25 mm in diameter within 14 days, tightly attached, flat, cushion-like, fuzzy, gray-purple, clear edges.

Material examined: CHINA, Yunnan Province, Nujiang, Gongshan County, on the ootheca of praying mantis (Mantidae), August 3, 2024, J.L. Jia (HKAS 135119, culture KUNCC 11022).

Notes: Thanakitpipattana et al. (2020) discovered a fungus parasitizing the oothecae of praying mantises in Khao Yai National Park, Thailand. Morphologically, this fungus showed similarities to Cor. mantidicola, but exhibited significant differences in the size of its perithecia and the shape of its ascospores. Based on a combination of multigene phylogenetic analysis and morphological data, researchers confirmed that it belongs to the family Clavicipitaceae and is distinct from other known genera. Consequently, Thanakitpipattana et al. (2020) established the new genus Petchia, with Pet. siamensis as the type species. In this study, phylogenetic analyses (Fig. 57), clearly demonstrated that the newly collected specimen belongs to the same species as Pet. siamensis were observed emerging from various parts of the host’s body, and their microscopic characteristics were highly consistent with those previously reported. This specimen represents a new distribution record for the genus Petchia, specifically for the species Pet. siamensis.

Cordycipitaceae Kreisel ex G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora

Akanthomyces Lebert 1858

Akanthomyces, described by Lebert in 1858, is typified by Aka. aculeatus, a species originally identified on a moth in Europe (Mains, 1950). Specific insect targeted by Akanthomyces include Hemiptera (Aka. lecanii), Coleoptera (Aka. neocoleopterorum), Lepidoptera (Aka. pistillariiformis), and Orthoptera (Aka. fragilis) (Hodge et al, 2003; Mongkolsamrit et al, 2018; Chen et al, 2020). Overall, the host range of Akanthomyces remains consistent for both its teleomorph and anamorph forms. Additionally, the genus encompasses Cordyceps tuberculata, which parasitizes adult moths and is taxonomically linked to the anamorphic stage Aka. pistillariiformis. Furthermore, Akanthomyces holds taxonomic precedence over Lecanicillium, a member of the family Cordycipitaceae characterized by its verticillium-like morphology.

34. Akanthomyces phariformis Khons., Thanakitp. & Luangsa-ard, in Khonsanit, Thanakitpipattana, Mongkolsamrit, Kobmoo, Phosrithong, Samson, Crous & Luangsa-ard, Fungal Syst. Evol. 14: 285 (2024) Fig. 60

Chinese name

Index Fungorum number: IF 842455; Facesoffungi number: FoF 19315

Diagnosis: The molecular data showed a high level of identity with Akanthomyces phariformis.

Parasitic on the adult of moth clinging to the stone. Sexual morph: Stromata numerous, arising from the head, thorax and abdomen of the insect host, 3.6 mm long, 0.6 mm wide. Stipes cylindrical to clavate, white. Fertile parts terminal, slightly curved, yellow, 2 mm long, 1.2 mm wide. Perithecia superficial, ovoid, 424–678 × 217–401 μm. Asci cylindrical, 8-spored, 176–433 × 4–9 μm. Ascal cap hemispherical, 3.6–4.9 × 2.2–3.5 μm. Ascospores hyaline, entire, cylindrical, 39–187 × 0.6–1.8 μm, with 10–16 septa. Secondary ascospores cylindrical, 5–10 μm long, smooth, hyaline. Asexual morph: Hyphae smooth-walled, hyaline, measuring 1.4–4.5 μm in width. Phialides 11.2–21.2 μm long, arising singly from vegetative hyphae; base cylindrical, middle portion swollen, tapering gradually or abruptly towards the apex; base 1.9–4.8 μm wide, neck 0.2–1.5 μm wide. Conidia 4.3–6.5 × 2.4–3.8 μm, smooth-walled, hyaline, globose to ellipsoidal.

Culture characteristics: Colonies on PDA attaining 15–30 mm in diameter after 14 days at 25 ℃, with dense, centrally elevated mycelia, margins radiating and creeping. Colony center pale yellow, margins white, reverse white to pale yellow.

Material examined: CHINA, Yunnan Province, Nujiang, Fugong County, on the adult of moth clinging to the stone, 5 September 2023, M. Tang, (HKAS 135021, culture KUNCC 11410).

Fig. 59. Petchia siamensis (HKAS 135119) a Stromata arising from ootheca of praying mantis. b, c Fertile parts. d Perithecia. e Colonies on PDA. f–h. Asci, i Ascospore. j Secondary ascospores. k–m Conidiophores, phialides and conidia on PDA. Bars: b, c = 2 mm, d = 200 μm, e = 2 cm, f = 50 μm, g = 20 μm, h–m = 10 μm.

Notes: Akanthomyces phariformis is characterized by numerous stromata, with a terminal, slightly curved, yellow fertile part, superficial ovoid perithecia, and cylindrical asci bearing filiform ascospores. Phylogenetic analyses placed the specimen within Akanthomyces and showed that it is closely related to Aka. taiwanicus. Morphologically, Aka. phariformis can be distinguished from Aka. taiwanicus by its phialides. In Aka. phariformis, phialides arise singly from vegetative hyphae, with a cylindrical base, a swollen middle portion, and a tapering apex that narrows gradually or abruptly, whereas these features are absent in Aka. taiwanicus. In addition, Aka. phariformis has a wider phialide base (1.9–4.8 μm) compared to that of Aka. taiwanicus (2.5–3 μm) (Chuang et al. 2024). Phylogenetic analysis of the present specimen showed that its clusters with Aka. phariformis, and morphological observations are consistent with the original description. Therefore, Aka. phariformis is reported here as a new record from China (Fig. 63).

Fig. 60. Akanthomyces phariformis (HKAS 135021) a Fungus on the adult of Lepidoptera. b Fertile part. c Section of perithecia. d–e Asci. f Ascocarp caps. g–h Ascospores. i Secondary ascospores. j–k Colony on PDA, front and back. l–q Phialides and conidia. Scale bars: a–b = 1000 μm, c–i = 10 μm, j–k = 1 cm, l–q = 10 μm.

Beauveria Vuill. 1912

Beauveria was proposed by Vuillemin in 1912, with B. bassiana (Bals.-Criv.) Vuill. as the type species. Since its inception, numerous investigations have been conducted to elucidate and delineate the distinct taxa within Beauveria (Petch 1926; MacLeod 1954; De Hoog 1972; Rehner & Buckley 2005). The identification of species within Beauveria utilizing solely ITS as a DNA barcode has reached a resolution threshold. Recently, there has been a discernible shift towards employing an integrative multi-locus delimitation framework for closely related species. To date, 81 records in the Index Fungorum (Index Fungorum July 18, 2025), and 17 species have been accepted in Beauveria (Imoulan et al. 2017).

35. Beauveria lanceolata Y.B. Wang, B.Z. Chen & Zhu L. Yang, sp. nov. Fig. 62

Chinese name:

Index Fungorum number: IF 905268; Facesoffungi number: FoF 19316

Etymology: The specific epithet “lanceolata” refers to the narrowly elongated and tapering stroma, resembling a spearhead.

Diagnosis: Beauveria lanceolata is distinguished by narrowly elongated, spearhead-shaped stroma arising from coleopteran larva, pale yellow to orange, with superficial, densely arranged ovoid to ellipsoidal perithecia.

Holotype: HKAS 135020

Parasitic on the larva of Coleoptera buried in soil and adult of Dermaptera clinging to the fallen leaves. Sexual morph: Stroma 2–5 cm long, solitary, unbranched, arising from larvae of Coleoptera, pale yellow to orange. Fertile part 1–4 cm long, orange, cylindrical or flattened, 2–10 mm wide. Perithecia 200–540 × 160–320 µm, superficial, densely aggregated, and ovoid to ellipsoidal. Asci 120–250 × 2–3 µm, cylindrical, slender. Ascospore caps 2–3.5 × 1.5–2.5 µm, hemispherical. Asexual morph: Hyphae 0.5–4 µm wide, hyaline, septate, smooth-walled. Conidiophores 8–12.5 × 1–2 µm, smooth, hyaline, cylindrical, branched at the apex. Phialides 4–18 × 1–2 µm, smooth-walled, cylindrical or flask-shaped, produced in clusters at the tips of conidiophores or directly from hyphae, with zigzag-shaped apices. Conidia 3–6 × 2–4 µm, single-celled, hyaline, smooth, ovoid to ellipsoidal, produced at the tips of phialides.

Culture characteristics: Mycelia emerge from the entire host body, forming white to pale yellow powdery conidia. Colonies on PDA reach 30–40 mm in diameter after 14 days at 25 °C, surface white to pale yellow, darkening outward, with smooth margins and a dense or slightly floccose texture, reverse side darker, centrally pale yellow and fading toward the edge.

Material examined: CHINA, Yunnan Province, Kunming City, 12 September 2023, on the adult of Dermaptera clinging to the fallen leaves, W.B Zeng (HKAS 135020 holotype, ex-type culture KUNCC 11286). CHINA, Yunnan Province, Kunming City, on the larva of Coleoptera buried in soil, 12 September 2023, Y.B. Wang (culture KUNCC 8665).

Notes: In the multigene phylogenetic analyses based on tef1-α, rpb1, rpb2, and Bloc sequences, B. lanceolata clusters closely with B. pseudobassiana within the same clade but is clearly separated with strong support (Fig. 61). Morphologically, B. lanceolata is distinct from B. pseudobassiana and B. varroae in producing ovoid to ellipsoidal conidia, whereas the latter species bear globose to broadly ellipsoidal conidia (Rehner et al. 2011).

Cordyceps Fr.

Cordyceps, a fascinating group of fungi, has garnered significant attention within scientific and medical communities due to its unique parasitic lifestyle and potential therapeutic properties. The diversity within Cordyceps is vast, encompassing over 300 identified species, with ongoing research continually broadening our understanding of this intriguing fungal group (Wang et al. 2020a). To date, 607 records have been listed in the Index Fungorum (Index Fungorum July 18, 2025).

36. Cordyceps biclavata Y.B. Wang, B.Z. Chen, M. Tang & Zhu L. Yang, sp. nov. Fig. 64

Chinese name

Index Fungorum number: IF905269; Facesoffungi number: FoF 19317

Etymology: The epithet “biclavata” refers to the two clavate stromata emerging from the head of lepidopteran pupa.

Diagnosis: Cordyceps biclavata differs from related species by producing two clavate stromata from the head of lepidopteran pupa, and the verticillium-like asexual morph.

Holotype: HKAS 135022

Parasitic on the pupae of Lepidoptera. Sexual morph: Stromata arising from the head of lepidopteran pupa buried in soil, two in number, unbranched, 30–35 mm long and 3–8 mm wide, orange to orange-yellow, cylindrical or flattened at the base and cylindrical toward the apex. Fertile parts cylindrical to clavate, 15–20 mm long and 2–6 mm wide, orange-yellow. Perithecia semi-immersed, ovoid, 280–499 × 134–379 μm. Asci eight-spored, hyaline, cylindrical. Ascal cap hemispherical, 2.8–4 × 1.3–3.3 μm. Asexual morph: Hyphae 1.5–3.3 μm wide, smooth, hyaline. Phialides 15.7–47.2 μm long, typically solitary or arising laterally from vegetative hyphae or terminally from conidiogenous cells; base cylindrical, tapering gradually toward the apex; basal width 1.3–3 μm, neck 0.3–1.2 μm wide. Conidia 3.4–5.2 × 1.8–3.3 μm, hyaline, ellipsoidal. Two types of conidia are observed, all are one-celled, occurring singly or more commonly aggregated into subglobose to ellipsoidal heads at the apex of phialides.

Culture characteristics: Colonies on PDA at 25 °C for 14 days grow rapidly, reaching 30–35 mm in diameter, with a floccose texture, white to pale yellow.

Material examined: CHINA, Yunnan Province, Baoshan City, the host is a pupa of Lepidoptera found underground. 5 September 2023, M. Tang, (HKAS 135022 holotype, ex-type culture KUNCC 11366), other collection: Ibid., M. Tang (culture KUNCC 11367).

Notes: Cordyceps biclavata is phylogenetically closely related to Cor. militaris, Cor. shuifuensis, and Cor. changbaiensis (Fig. 63), but forms a distinct and well-supported clade within this lineage, indicating that it represents a separate species. Morphologically, the stromata of Cor. biclavata are cylindrical or flattened at the base and cylindrical toward the apex, differing from the solitary, cylindrical to slightly clavate stromata observed in Cor. militaris and Cor. shuifuensis (Wang et al. 2020a). Among these related species, Cor. biclavata is most similar to Cor. militaris, sharing characteristics such as cylindrical to slightly clavate stromata with yellowish to reddish-orange coloration, superficial perithecia, and a verticillium-like asexual morph. However, it can be distinguished by its overall smaller size and the absence of an isaria-like asexual morph, which is present in Cor. militaris (Wang et al. 2020a). In contrast, Cor. changbaiensis differs significantly from Cor. biclavata in having longer stromata covered with white mycelia and in parasitizing coleopteran larvae rather than lepidopteran hosts (Hu et al. 2021). These phylogenetic and morphological distinctions support the recognition of Cor. biclavata as a novel species within the genus Cordyceps.