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. 2026 Feb 27;129:107–128. doi: 10.3897/mycokeys.129.179192

Taxonomic and phylogenetic analyses reveal two new Comoclathris taxa (Dothideomycetes, Pleosporaceae) from Inner Mongolia, China

Xing-Guo Tian 1,2, Tian-Ye Du 3, An-Dong Ma 3, Rajesh Jeewon 4,5,6, Samantha C Karunarathna 7, Saowaluck Tibpromma 7, Dan-Feng Bao 1,2,✉
PMCID: PMC12966847  PMID: 41798702

Abstract

The Inner Mongolia Autonomous Region, located in northern China, represents a temperate semi-arid ecosystem dominated by steppe vegetation and characterized by rich, yet understudied, fungal diversity. During an investigation of microfungi associated with decaying plant materials in the desert regions of Inner Mongolia, two novel species of Comoclathris were collected and isolated. Detailed morphological examinations and multi-locus phylogenetic analyses based on ITS, LSU, SSU, and rpb2 sequence data support the establishment of these taxa as novel species: Comoclathris desertica and C. xiangshawanensis. These two species are described and illustrated herein, with particular attention to their differences from similar species with respect to ascomatal structures and ascospore morphology. The discovery of these new taxa expands the known diversity and distribution of Comoclathris in China and provides new insights into the saprobic fungal communities of semi-arid regions. This study also highlights the ecological and taxonomic significance of continued surveys on saprobic fungi in grassland and desert ecosystems of northern China.

Key words: 2 new species, desert, Dothideomycetes , Pleosporales , saprobes

Introduction

Comoclathris Clem. is a genus of ascomycetous fungi belonging to Pleosporaceae Nitschke (Pleosporales, Dothideomycetes). The genus was established by Clements (1909), with C. lanata Clem. designated as the type species. Previously, the genus Comoclathris was placed in Diademaceae Shoemaker & C.E. Babc. (Shoemaker and Babcock 1992), but recent morphological and molecular phylogenetic analyses have supported its placement in Pleosporaceae (Zhang et al. 2012; Ariyawansa et al. 2015; Hyde et al. 2024). Currently, 52 epithets of Comoclathris are listed in Index Fungorum (2026), with molecular data available for only 21 species in NCBI. This genus is characterized by ascomata with circular lid-like openings, cylindrical to clavate asci, and muriform ascospores that are reddish-brown to dark brown and applanate (Mattoo et al. 2023). While most Comoclathris species are mainly saprobic on woody substrates, dead stems, and leaves of various plants or soil (Wanasinghe et al. 2015; Thambugala et al. 2017; Brahmanage et al. 2020; Crous et al. 2021), some are endophytes (González-Menéndez et al. 2018; Mattoo et al. 2023) and phytopathogens (Moral et al. 2017). Comoclathris is a widely distributed genus, found in many countries and regions, as detailed in recent publications (Mattoo et al. 2023; Xu et al. 2024). Comoclathris was first reported in China by Xu et al. (2024), who described two novel species from Jilin and Yunnan provinces.

The Inner Mongolia Autonomous Region is located in northern China and lies within a temperate semi-arid zone, with ecological types covering typical steppe, meadow steppe, desert steppe, as well as sandy land and forest-steppe transition zones (Han et al. 2009; Su et al. 2011; Yan et al. 2025). The vegetation of this region is dominated by herbaceous plants of the Asteraceae Bercht. & J.Presl, Fabaceae Lindl., and Poaceae (R. Brown) Barnhart families, and although steppe vegetation predominates, the types are relatively diverse rather than strictly monotypic (Fu et al. 2021; Wang et al. 2024). Despite this lack of vegetation diversity, fungi are important in these regions, as they may harbour new species with significant bioprospecting potential. Recent studies have shown that the fungal communities in Inner Mongolia exhibit considerable diversity. For example, analyses from 63 root samples during a study of ectomycorrhizal (EM) fungi revealed 288 operational taxonomic units (OTUs) spanning 31 lineages, indicating that fungal communities in forest/forest-edge environments may be much more diverse than anticipated (Wang et al. 2021). Further studies on arbuscular mycorrhizal (AM) fungi in grassland environments conducted by Wang et al. (2020) revealed 24 species spanning across eight genera. Despite increasing research on fungal diversity in Inner Mongolia Autonomous Region, detailed data and systematic taxonomic studies on saprobic fungi remain scarce. To date, only a few saprobic fungal novelties have been reported from Inner Mongolia, for example, Diatrype betulaceicola Z.E. Yang and Hai X. Ma, with polysporous asci described by Yang et al. (2022). The unique ecological conditions of the semi-arid grassland ecosystem in this region make saprobic fungi crucial in decomposing plant debris, cycling nutrients, and maintaining soil fertility. Therefore, studying the taxonomy of saprobic fungi is essential to unravel their hidden fungal diversity and potential applications.

Our ongoing studies in this region explore the diversity of saprobic ascomycetes inhabiting dead plant material in the desert regions of Inner Mongolia, specifically the Xiangshawan Scenic Area. During a survey of microfungi from decaying substrates, two species of Comoclathris were collected and examined. Morphological characteristics and phylogenetic analyses based on multi-locus sequence data (ITS, LSU, SSU, and rpb2) were employed to confirm their taxonomic placement within Pleosporaceae and support their establishment as novel species. This study provides new insights into the saprobic fungal diversity of northern China and expands the known distribution and species richness of the genus Comoclathris.

Materials and methods

Sample collection, morphological observation, single-spore isolation and preservation

During a fungal resource survey in the desert areas of Inner Mongolia Autonomous Region, China (Fig. 1), dead fallen branches and stems with fungal fruiting bodies were collected. Important information was recorded during the collection process (collector, date, location, and habitat) (Rathnayaka et al. 2024). After that, the samples were placed in dry plastic bags for further observation and experimentation in the laboratory. Morphological structures were examined by using an OPTEC SZ650 dissecting stereomicroscope (Chongqing, China), and the microstructures of fungi were observed and photographed by using an OLYMPUS optical microscope (Tokyo, Japan) with an OLYMPUS DP74 (Tokyo, Japan) digital camera. Micro-morphological structures were measured in the Tarosoft ® Image Framework program v. 1.3. The photo plates were edited in Adobe Photoshop CS3 Extended version 22.0.0 software (Adobe Systems, California, USA).

Figure 1.

Figure 1.

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Vegetation and habitat in the desert area of Xiangshawan Scenic Area.

Fungi were isolated and cultured using the single-spore isolation method described by Senanayake et al. (2020). The experiment was conducted under a sterile environment. After the work surface was disinfected with 75% alcohol, an alcohol lamp was lit. First, fresh fruiting bodies were observed and selected, then cut open with a sterile blade. Spores were picked up with a sterile needle, placed in sterile water and dispersed, and finally transferred to potato dextrose agar (PDA) plates (100 mL/plate) using a pipette. From 6 hours after inoculation to 48 hours, spores were checked promptly after germination with a sterile needle and then placed on a new PDA plate at 23–28 °C. Four germinated spores were transferred to each plate. After colonies formed, the morphology was observed to ensure consistency and confirm the results of the single-spore isolation experiment. The colonies were purified and used for molecular experiments, observed for sporulation, and photographed.

Specimens were deposited at Guizhou Medical University (GMB-W), China. Living cultures are deposited in the Guizhou Medical University Culture Collection (GMBCC), China. Facesoffungi (FoF) numbers were assigned as described by Jayasiri et al. (2015), and MycoBank (MB) numbers were assigned as outlined at https://www.mycobank.org/.

DNA extraction, PCR amplification, and sequencing

Total genomic DNA was extracted from one-month-old fresh fungal mycelium (grown on PDA). The DNA Extraction Kit-BSC14S1 (BioFlux, Hangzhou, P.R. China) was used following the manufacturer’s instructions. Polymerase chain reactions (PCR) were carried out using the following primers: The internal transcribed spacer (ITS) region was amplified with the primers ITS4 and ITS5 (White et al. 1990), 28S nrRNA gene (LSU) was amplified by using the primers LR0R and LR5 (Vilgalys and Hester 1990), 18S ribosomal RNA (SSU) was amplified using the primers NS1 and NS4 (White et al. 1990), and the partial RNA polymerase II subunit (rpb2) region was amplified with primers fRPB2-5F and fRPB2-7cR (Liu et al. 1999). The PCR thermal cycle programs for ITS, LSU, and SSU were as follows: an initialization step of 94 °C for 3 min, followed by 35 cycles of 94 °C for 30 s, an annealing step at 55 °C for 50 s, an elongation step at 72 °C for 1 min and a final extension step of 72 °C for 10 min; and the PCR thermal cycle programs for rpb2 was as follows: an initialization step of 95 °C for 3 min, followed by 40 cycles of 95 °C for 50 s, an annealing step at 57 °C for 50 s, an elongation step at 72 °C for 90 s and a final extension step of 72 °C for 10 min (Du et al. 2025). The DNA amplification procedure was performed by PCR in a 25 μL reaction containing 12.5 μL 2xMaster Mix (mixture of Easy Taq TM DNA Polymerase, dNTPs, and optimized buffer (Beijing Trans Gen Biotech Co., Chaoyang District, Beijing, China)), 8.5 μL ddH2O, 2 μL of DNA template, and 1 μL of each forward and reverse primer (10 pM). Purification and sequencing of PCR products were carried out by Sangon Biotech Co., Kunming, China.

Phylogenetic analyses

A combined dataset of ITS, LSU, SSU, and rpb2 was used for the phylogenetic analyses. The quality of the raw sequences was checked in BioEdit v.7.2.6.1 (Hall 1999), and the forward and reverse sequences were spliced with Geneious 9.1.8 (Kearse et al. 2012). Sequences obtained from this study were searched in the GenBank database (http://blast.ncbi.nlm.nih.gov/) using BLAST to identify the closely related taxa of our strains. The additional sequences included in the analyses were collected from previous publications (Mattoo et al. 2023; Xu et al. 2024) and downloaded from GenBank (Benson et al. 2013). Phylogenetic analyses were carried out with 41 sequences (Table 1). The FASTA file was used for constructing the Randomized Accelerated Maximum Likelihood (RAxML) and Bayesian Inference analyses (BI) was performed using the OFPT (Zeng et al. 2023) with the protocol. Then, the FASTA file was converted to PHYLIP and NEXUS formats in ALTER for RAxML and BI phylogenetic analyses, respectively (Glez-Peña et al. 2010).

Table 1.

Taxa names, strain numbers, and corresponding GenBank accession numbers of the taxa included in the present study. Remarks: The newly generated sequences are indicated in bold, the superscript T indicates ex-type, and “—” indicates information unavailable.

Taxon name Strain number ITS LSU SSU rpb2 References
Comoclathris acuminata MCC9771T MW205771 OQ547244 — — Mattoo et al. (2023)
C. acuminata MN3-2019 OQ547243 OQ547245 — — Mattoo et al. (2023)
C. ambigua CBS 366.52 KY940748 AY787937 — KT216533 Woudenberg et al. (2017)
C. antarctica WA0000074564T MW040594 MW040597 — — Crous et al. (2021)
C. arrhenatheri MFLUCC 15-0465T KX965737 KY000647 KX986348 KX938346 Thambugala et al. (2017)
C. arrhenatheri MFLUCC 15-0476 KY026595 KY000648 KX986349 — Thambugala et al. (2017)
C. clematidis CCMJ 13076T OQ534243 OQ534239 OQ676454 OQ547800 Xu et al. (2024)
C. clematidis CCMJ 13077 OQ534244 OQ534240 OQ676455 OQ547801 Xu et al. (2024)
C. compressa CBS 156.53 — KC584372 KC584630 KC584497 Woudenberg et al. (2013)
C. compressa CBS 157.53 — MH868679 KC584631 KC584498 Woudenberg et al. (2013)
C. desertica GMBCC2307T PX660512 PX660516 PX660508 PX672979 This study
C. desertica GMBCC2320 PX660513 PX660517 PX660509 PX672980 This study
C. europaeae MFLU 20-0391T MT370396 MT370421 MT370367 MT729650 Brahmanage et al. (2020)
C. flammulae MFLU 20-0397T MT370397 MT370422 MT370368 MT729651 Brahmanage et al. (2020)
C. flammulae MFLU 20-0399 MT370395 MT370420 MT370366 — Brahmanage et al. (2020)
C. galatellae MFLUCC 18-0773T MN632549 MN632550 MN632551 — Hongsanan et al. (2020)
C. incompta CBS 467.76 — GU238087 GU238220 KC584504 Aveskamp et al. (2010)
C. incompta CH-16 KU973716 KU973729 — — Moral et al. (2017)
C. italica MFLUCC 15-0073T KX500109 — — — Tibpromma et al. (2015)
C. lini MFLUCC 14-0968T KR049218 KR049219 KT210389 — Wanasinghe et al. (2015)
C. lini MFLUCC 14-0561 KT591614 KT591615 KT591616 — Wanasinghe et al. (2015)
C. lonicerae MFLU 20-0385T MT370394 MT370419 MT370365 MT729649 Brahmanage et al. (2020)
C. lonicerae MFLU 18-1236 OL744429 OL744433 OL744435 OL771441 Brahmanage et al. (2020)
C. neimengguensis GMBCC2306T PX314703 PX314701 PX314705 PX310456 Unpublished
C. neimengguensis GMBCC2319 PX314704 PX314702 PX314706 PX310457 Unpublished
C. permunda MFLUCC 14-0974 KY659561 KY659564 KY659568 — Vu et al. (2019)
C. pimpinellae MFLUCC 14-1159T KU987665 KU987666 KU987667 — Li et al. (2016)
C. rosae MFLU 15-0203T MG828876 MG828992 MG829103 MG829249 Wanasinghe et al. (2018)
C. rosae MFLU 16-0234 MG828877 MG828993 MG829104 MG829250 Wanasinghe et al. (2018)
C. rosarum MFLUCC 14-0962T MG828878 MG828994 MG829105 MG829251 Wanasinghe et al. (2018)
C. rosigena MFLU 16-0229T MG828879 MG828995 MG829106 MG829252 Wanasinghe et al. (2018)
C. sedi MFLUCC 13-0763T KP334717 KP334707 KP334727 — Ariyawansa et al. (2014)
C. sedi MFLUCC 13-0817 KP334715 KP334705 KP334725 — Ariyawansa et al. (2014)
C. spartii MFLUCC 13-0214T KM577159 KM577160 KM577161 — Crous et al. (2014)
C. typhicola CBS 602.72 MH860592 MH872288 — — Vu et al. (2019)
C. xanthoceratis CCMJ 13078T OQ534245 OQ534241 OQ676456 OQ547802 Xu et al. (2024)
C. xanthoceratis CCMJ 13079 OQ534246 OQ534242 OQ676457 OQ547803 Xu et al. (2024)
C. xiangshawanensis GMBCC2305T PX660510 PX660514 PX660506 PX672977 This study
C. xiangshawanensis GMBCC2318 PX660511 PX660515 PX660507 PX672978 This study
Neocamarosporium betae CBS 523.66 FJ426981 MH870520 EU754080 KT389670 Aveskamp et al. (2009)
N. calvescens CBS 246.79 MH861203 EU754131 EU754032 KC584500 Vu et al. (2019)
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The CIPRES Science Gateway platform was used to carry out the RAxML and BI analyses (Miller et al. 2010). The RAxML tree, generated with 1,000 bootstrap replicates, was analyzed using RAxML-HPC2 on XSEDE (8.2.12) (Stamatakis et al. 2008; Stamatakis 2014) with the GTR+I+G model of evolution and bootstrap support. The BI tree was performed with MrBayes on XSEDE (3.2.7a) (Ronquist et al. 2012) by the Markov Chain Monte Carlo (MCMC) method to evaluate posterior probabilities (BYPP) (Richard and Lippmann 1991; Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002). The best-fit nucleotide substitution models for each dataset were then selected using the Bayesian information criterion (BIC) from 22 common DNA substitution models with rate heterogeneity, as implemented in ModelFinder (Kalyaanamoorthy et al. 2017). The best model for ITS was TIM2e+I+G4, K2P+I for LSU, TPM2u+F+I for SSU, and TN+F+G4 for rpb2. Six simultaneous Markov chains were run for 2,000,000 generations, and a tree was sampled every 100th generation. The phylogenetic tree was visualized in FigTree v.1.4.2 (Rambaut 2012) and edited in Microsoft PowerPoint 2021 and Adobe Photoshop CS3 Extended version 22.0.0 (Adobe Systems, California, USA). All newly generated sequences in this study were deposited in GenBank (https://www.ncbi.nlm.nih.gov/WebSub/?form=history&tool=genbank).

Results

Phylogenetic analyses

The phylogenetic trees obtained from RAxML and Bayesian Inference analyses were essentially similar. The RAxML analysis of the combined dataset yielded the best-scoring tree (Fig. 2), which consisted of 41 taxa, and the final alignment comprised 4029 characters, including gaps (ITS: 1–540, LSU: 541–1922, rpb2: 1923–3083, SSU: 3084–4029). The final ML optimization likelihood value was -14494.444982. The matrix contained 935 distinct alignment patterns, with 34.05% of characters undetermined or missing. Parameters for the GTR+I+G model of the combined ITS, LSU, rpb2, and SSU were as follows: estimated base frequencies A = 0.254158, C = 0.228977, G = 0.268187, T = 0.248678; substitution rates AC = 1.928002, AG = 4.366339, AT = 1.437220, CG = 1.029313, CT = 7.549462, GT = 1.0; proportion of invariable sites I = 0.641653; and gamma distribution shape parameter α = 0.725266. The final RAxML tree is shown in Fig. 2.

Figure 2.

Figure 2.

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Maximum likelihood consensus tree inferred from the combined ITS, LSU, rpb2, and SSU multiple sequence alignments. Bootstrap support values for maximum likelihood (ML, first value) equal to or greater than 60% and Bayesian posterior probabilities from MCMC analyses (BYPP, second 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 Neocamarosporium betae (CBS 523.66) and N. calvescens (CBS 246.79). The new isolates are indicated in red, and the ex-type strains are in bold.

In our phylogenetic analyses, we obtained results consistent with those of recent publications by Mattoo et al. (2023) and Xu et al. (2024). Our new species, Comoclathris desertica (GMBCC2307 and GMBCC2320), was well separated from C. clematidis R. Xu, Phukhams. & Yu Li (CCMJ 13076 and CCMJ 13077) and C. xanthoceratis R. Xu, Phukhams. & Yu Li (CCMJ 13078 and CCMJ 13079) in a distinct lineage with 100% ML/1.00 PP statistical support; Comoclathris xiangshawanensis (GMBCC2305 and GMBCC2318) was well separated from C. antarctica Istel, J. Pawłowska & Wrzosek (WA0000074564, ex-type) with 87% ML/0.98 PP statistical support.

Taxonomy

Comoclathris desertica

X.G. Tian, T.Y. Du & D.F. Bao sp. nov.

74F9D643-66EA-5A91-8FAD-FF66B536FE4B

MB861793

Facesoffungi Number: FoF19021

Fig. 3

Figure 3.

Figure 3.

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Comoclathris desertica (GMB-W1518, holotype). a, b. Appearance of ascomata on the substrate; c. Section through an ascoma; d. Peridium; e-g, i. Asci; h. Pseudoparaphyses; j–n. Ascospores (m, n surrounded by mucilaginous sheath); o. A germinated ascospore; p, q. Colonies on PDA from above and below after one week. Scale bars: 100 µm (c); 10 µm (d, h); 50 µm (e-g, i); 20 µm (j–o).

Etymology.

Named after the desert habitat from where the holotype was collected.

Holotype.

GMB-W1518.

Description.

Saprobic on dead branches of an unidentified wood. Sexual morph: Ascomata 90–200 × 70–190 μm (x̄ = 115 × 116 μm, n = 5), solitary, scattered or aggregated, immersed, not obvious on the surface of the host, subglobose, black, ostiole not obvious. Peridium 10–30 μm wide, comprising thin-walled cells of textura angularis, dark brown to black. Hamathecium comprising numerous, 2 μm wide, filamentous, septate, branched, cellular pseudoparaphyses, hyaline, embedded in a gelatinous matrix, extending above the asci. Asci 65–85 × 18–23 μm (x̄ = 76 × 20 μm, n = 30), bitunicate, fissitunicate, 8-spored, cylindrical-clavate, short pedicellate, apically rounded. Ascospores 16–23 × 8–10 μm (x̄ = 19 × 9 μm, n = 30), 1-seriate, slightly overlapping, fusiform, initially yellowish, becoming brown to dark brown, muriform, with 3 transversely septa and 1 vertical septum, consisting of 6 cells, upper end conical, lower end tapered gradually, apical cell mostly undivided, straight or slightly curved, smooth-walled, surrounded by a mucilaginous sheath. Asexual morph: Undetermined.

Culture characteristics.

Ascospores germinated on PDA within 12 h at 28 °C, and germ tubes were produced around spores. Colonies on PDA reached 2 cm diam. after one week at 28 °C. Colonies obverse: circular, white-cream to grey, with sparsely hairy edge; grey from below, the active hyphae at the edge are white.

Material examined.

China • Inner Mongolia Autonomous Region, Ordos City, Xiangshawan, on dead branches of an unidentified wood in the desert, 01 October 2023, J.Y. Zhang, BTD24 (GMB-W1518, holotype), ex-type, GMBCC2307, other living culture, GMBCC2320.

GenBank numbers.

GMBCC2307: ITS = PX660512, LSU = PX660516, SSU = PX660508, rpb2 = PX672979; GMBCC2320: ITS = PX660513, LSU = PX660517, SSU = PX660509, rpb2 = PX672980.

Notes.

Comoclathris desertica clustered with C. clematidis (CCMJ 13076, ex-type and CCMJ 13077) and C. xanthoceratis (CCMJ 13078, ex-type and CCMJ 13079) in the phylogenetic tree with 100% in ML and 1.00 in BYPP statistical support (Fig. 2). A nucleotide base pair differences between our new strain (GMBCC2307, ex-type) and C. clematidis (CCMJ 13076, ex-type) in ITS, LSU, SSU, and rpb2 (without gaps) showed a 2.70% (14/519 bp) difference in ITS, 1.33% (9/679 bp) difference in LSU, 0.32% (2/628 bp) difference in SSU, and 5.58% (47/842 bp) difference in rpb2. Between our new strain (GMBCC2307, ex-type) and C. xanthoceratis (CCMJ 13078, ex-type), there was a 2.35% (12/510 bp) difference in ITS, 1.18% (8/676 bp) difference in LSU, 0.32% (2/628 bp) difference in SSU, and 6% (52/866 bp) difference in rpb2.

Morphologically, C. desertica is similar to C. clematidis and C. xanthoceratis, both of which have ascospores with 3 transverse septa and a vertical septum, surrounded by a thick mucilaginous sheath. But C. desertica is different from C. clematidis and C. xanthoceratis in having immersed ascomata, and dark brown ascospores, 1-seriate, slightly overlapping, smooth-walled; while C. clematidis has immersed to erumpent ascomata, and brown ascospores, with verrucose or echinulate wall (Xu et al. 2024); while C. xanthoceratis has immersed to semi-immersed ascomata, and brown to dark brown ascospores, 1–2-seriate (Xu et al. 2024). And our collection displays comparatively smaller ascospores than these two species (viz., C. desertica: 19 × 9 μm, C. clematidis: 30 × 14 μm, C. xanthoceratis: 37 × 16 μm).

Therefore, C. desertica is introduced as a new species from China based on morphological and molecular evidence as recommended by Jeewon and Hyde (2016).

Comoclathris xiangshawanensis

X.G. Tian, T.Y. Du & D.F. Bao sp. nov.

E45D5D8A-CF79-5DB2-A8EB-EBFFC06FD720

MB861794

Facesoffungi Number: FoF19022

Fig. 4

Figure 4.

Figure 4.

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Comoclathris xiangshawanensis (GMB-W1526, holotype). a, b. Appearance of ascomata on the substrate (arrows indicate black ascomata); c, d. Section through ascomata; e. Peridium; f. Pseudoparaphyses; g–k. Asci; l–q. Ascospores (arrow indicates mucilaginous sheath); r. A germinated ascospore; s, t. Colonies on PDA from above and below after one week. Scale bars: 100 µm (c, d); 10 µm (e, f); 50 µm (g–k); 20 µm (l–r).

Etymology.

Named after the type locality, “the desert region of Xiangshawan, China”.

Holotype.

GMB-W1526.

Description.

Saprobic on dead branches of an unidentified wood. Sexual morph: Ascomata 100–200 × 90–200 μm (x̄ = 132 × 118 μm, n = 10), solitary, scattered or aggregated into small groups, semi-immersed, subglobose, black, ostiole not obvious. Peridium 10–20 μm wide, comprising thin-walled cells of textura angularis, dark brown to black. Hamathecium comprising numerous, 2 μm wide, filamentous, septate, branched, cellular pseudoparaphyses, hyaline, embedded in a gelatinous matrix, extending above the asci. Asci 65–85 × 15–23 μm (x̄ = 73 × 21 μm, n = 30), bitunicate, fissitunicate, 8-spored, cylindrical-clavate, short pedicellate, apically rounded. Ascospores 17–23 × 7–11 μm (x̄ = 19 × 8.5 μm, n = 30), 1–2-seriate, broadly fusiform, initially yellowish, becoming brown, muriform, with 5(–6) transversely septa and 1–2 vertical septa, consisting of 9–14 cells, with conical or obtuse ends, straight or slightly curved, smooth-walled, surrounded by a mucilaginous sheath. Asexual morph: Undetermined.

Culture characteristics.

Ascospores germinated on PDA within 12 h at 28 °C and germ tubes were produced around spores. Colonies on PDA reached 2 cm diam. after one week at 28 °C. Colonies obverse: circular, white-cream, with sparsely hairy edge; cream to light yellow from below.

Material examined.

China • Inner Mongolia Autonomous Region, Ordos City, Xiangshawan, on dead branches of an unidentified wood in the desert, 01 October 2023, J.Y. Zhang, BTD16 (GMB-W1526, holotype), ex-type, GMBCC2305, other living culture, GMBCC2318.

GenBank numbers.

GMBCC2305: ITS = PX660510, LSU = PX660514, SSU = PX660506, rpb2 = PX672977; GMBCC2318: ITS = PX660511, LSU = PX660515, SSU = PX660507, rpb2 = PX672978.

Notes.

Comoclathris xiangshawanensis clustered with C. antarctica (WA0000074564, ex-type) and C. acuminata Mattoo, Nonzom & A. Ghosh (MCC9771, ex-type and MN3-2019) in the phylogenetic tree (Fig. 2). A comparison of nucleotide base pair differences between our new strain (GMBCC2305, ex-type) and C. antarctica (WA0000074564, ex-type) in ITS and LSU, the result (without gaps) showed a 1.65% (8/484 bp) difference in ITS, 0.24% (2/818 bp) difference in LSU. Between our new strain (GMBCC2305, ex-type) and C. acuminata (MCC9771, ex-type) in ITS and LSU, the result (without gaps) showed a 1.86% (9/484 bp) difference in ITS, 0.25% (2/807 bp) difference in LSU (C. antarctica (WA0000074564, ex-type) and C. acuminata (MCC9771, ex-type) have no available SSU and rpb2 sequences in NCBI and hence cannot be compared).

Morphologically, C. xiangshawanensis differs from C. antarctica and C. acuminata in having brown ascospores with 5(–6) transversely septa and 1–2 vertical septa, consisting of 9–14 cells, surrounded by a mucilaginous sheath; while C. antarctica has ascospores with 6–8 transvers septa, consisting of 10–17 cells (Crous et al. 2021); while C. acuminata has dark brown ascospores, with 3–8 transverse septa, 3–5 vertical septa, consisting of 15–21 cells, with a beak-like extension (Mattoo et al. 2023). In addition, our collection is saprobic on dead branches in desert areas in China, while C. antarctica is isolated from soil sample in Antarctica, and C. acuminata was isolated as an endophyte from the stem of Ephedra gerardiana Wall. in India (Mattoo et al. 2023).

Therefore, C. xiangshawanensis is introduced as a new species from China based on morphological and molecular evidence. To provide a better overview of the different morphologies associated with different Comoclathris species, we also compiled ascospore morphology and other ecological data in a table (Table 2).

Table 2.

Morphological characteristics of ascospore and other ecological data of Comoclathris species. NA: Information not available.

Species Ascospore Hosts Collection locations References
Color Septa Sheath
Comoclathris acuminata Dark brown 3–8 transverse septa and 3–5 vertical septa N/A Ephedra gerardiana India Mattoo et al. (2023)
Comoclathris antarctica Yellow to pale brown 6–8 transverse septa N/A Soil sample Antarctica Crous et al. (2021)
Comoclathris arctica Mid to dark reddish-brown 6 transverse septa and 1 vertical septum With a sheath 2–3 μm wide Festucae brachyphyllae, Puccinellia angustata Canada Shoemaker and Babcock (1992)
Comoclathris arrhenatheri Yellow to pale brown 4 transverse septa and 2–3 vertical septa Surrounded by a thick mucilaginous sheath Arrhenatherum elatius Italy Thambugala et al. (2017)
Comoclathris bougainvilleae N/A N/A N/A Bougainvillea spectabilis India Patil and Ramesh (1987)
Comoclathris cichoriacearum N/A N/A N/A Hieracium villosum N/A Nograsek (1990)
Comoclathris clematidis Brown 3 transverse septa and 1 vertical septum Surrounded by a thick mucilaginous sheath Clematis sp. China Xu et al. (2024)
Comoclathris compressa Dark reddish-brown 3 transverse septa and 1 vertical septum With a uniform sheath 2–3 μm wide Agastache urticifolia, Agoseris glauca, Aquilegia leptocera, Arternisia pedatzjidi, Balsatnorrhiza deltoidea, Brickellia grandijlora, Coleosanthus grandijlorus, Castilleja miniata, Clematis hirsutissirna, Helianthella sp., Helianthus sp., Juncus balticus, Ligusticum purpureum, Lupinus parvijlorus, Penstemon sp., Polygonum alpina, Rudbeckia sp., Senecio thapsoides, Valeriana sp., Viguiera multijlora, Wyethia mollis, Wyethia sp., Xerophyllum tenux USA Shoemaker and Babcock (1992)
Comoclathris depressa Mid yellowish-brown 3 transverse septa and 1 vertical septum With a sheath 2–3 μm wide Arabis sp., Lotnatiutn dissectum USA Shoemaker and Babcock (1992)
Comoclathris emodi Dark yellowish-brown 4 transverse septa and 1 vertical septum With a sheath 1.5–4 μm wide Festucae lucidae India Shoemaker and Babcock (1992)
Comoclathris europaeae Brown 7 transverse septa and 1 vertical septum Surrounded by amucilaginous sheath Olea europaea Italy Brahmanage et al. (2020)
Comoclathris extrema Dark reddish-brown 4 transverse septa and 1 vertical septum With a sheath 2–3 μm wide Lini perenni India Shoemaker and Babcock (1992)
Comoclathris flammulae Dark brown 6 transverse septa and 1–2 vertical septa Mucilaginous sheath Clematis flammula, Colutea arborescens Italy Brahmanage et al. (2020)
Comoclathris galatellae Brown or pale brown 2–4 transverse septa and 1–2 vertical septa Without sheath Galatella villosa Russia, Ukraine Hongsanan et al. (2020)
Comoclathris harperi Mid reddish-brown 7 transverse septa and 1 vertical septum With a sheath 1–2 μm wide Carex rupestris, Hierochloe alpina Sweden Shoemaker and Babcock (1992)
Comoclathris incompta N/A N/A N/A Olea europaea Greece, Italy Ariyawansa et al. (2015)
Comoclathris indica N/A N/A N/A Arachis hypogaea India Patil and Pawar (1987)
Comoclathris italica Brown 6–8 transverse septa and 1–2 vertical septa Mucilaginous sheath Dactylis glomerata Italy Thambugala et al. (2017)
Comoclathris lanata Brown to reddish-brown 4–5 transverse septa and 1–2 vertical septa Surrounded by a distinct, hyaline, mucilaginous 3–8 μm wide sheath Leptotaenia multifida USA Ariyawansa et al. (2015)
Comoclathris lini yellowish-brown 3–5 transverse septa and 1–2 vertical septa With a thick mucilaginous sheath Linum sp. Italy Wanasinghe et al. (2015)
Comoclathris lonicerae yellowish-brown 3–5 transverse septa and 1–2 vertical septa With a thick mucilaginous sheath Lonicera sp. Italy Brahmanage et al. (2020)
Comoclathris magna Dark reddish-brown 3 transverse septa and 1 vertical septum With a sheath 2–4 μm wide Anemone occidentalis, Chlorogalum pomeridianum, Helianthella californica, Linum perenne, Lomatium nudicaule, Penstemon gracilentus, Sphenosciadium capitellatum, umbellifer USA Shoemaker and Babcock (1992)
Comoclathris miliarakisii N/A N/A N/A Sclerochorton junceum Greece Petrak (1938)
Comoclathris persica Mid reddish-brown 4 transverse septa and 1 vertical septum With a uniform sheath 3–4 μm wide Dianthus orientalis Iran Shoemaker and Babcock (1992)
Comoclathris pimpinellae Yellow to light brown 3 transverse septa and 2 vertical septa Surrounded by a thick, hyaline, a mucilaginous sheath Pimpinella tragium Russia Li et al. (2016)
Comoclathris platysporioides Dark reddish-brown 5 transverse septa and 1 vertical septum With a uniform sheath, 1–2 μm wide Selaginella wrightii USA Shoemaker and Babcock (1992)
Comoclathris pseudoverruculosa N/A N/A N/A Carex firma N/A Nograsek (1990)
Comoclathris pyrenophoroides N/A N/A N/A Festuca and Bromus Italy Checa (1998)
Comoclathris quadriseptata Mid reddish-brown 4 transverse septa and 1 vertical septum With a uniform sheath 1–2 μm wide Agropyron divergens, Avena pratensis, Dianthus orientalis, Matthiola sp., Poaceae, Sporobolus cryptandrus Canada Shoemaker and Babcock (1992)
Comoclathris rosae Pale brown 4–7 transverse septa and 1–2 vertical septa Surrounded by a thick mucilaginous sheath Rosa canina Italy Wanasinghe et al. (2018)
Comoclathris rosarum Brown 6–7 transverse septa and 2–4 vertical septa Surrounded by a thick mucilaginous sheath Rosa canina Italy Wanasinghe et al. (2018)
Comoclathris rosigena Pale brown 5–7 transverse septa and 1 vertical septum Surrounded by a thick mucilaginous sheath Rosa canina Italy Wanasinghe et al. (2018)
Comoclathris salsolae Mid yellowish-brown 4 transverse septa and 1 vertical septum With a uniform sheath 2–3 μm wide lpomoealeptoplzylla, Salsola sp. USA Shoemaker and Babcock (1992)
Comoclathris sedi Brown to reddish-brown 4–5 transverse septa and 1–2 vertical septa Surrounded by a distinct, hyaline, mucilaginous 5–9 μm wide sheath Clematis vitalba, Rosa sp., Sedum sp. Italy Ariyawansa et al. (2015)
Comoclathris sisyrinchii Mid yellowish-brown 3 transverse septa and 1 vertical septum N/A Carpha alina, Lomatium californicuin, Sisyrinchium junceum, Sisyrinchium middletoni USA Shoemaker and Babcock (1992)
Comoclathris sororia Dark reddish-brown 3 transverse septa and 1 vertical septum N/A Dianthus orientalis Turkey Shoemaker and Babcock (1992)
Comoclathris spartii Yellow to pale brown Muriform Surrounded by a mucilaginous sheath Spartium junceum Italy Crous et al. (2014)
Comoclathris subdeflectens N/A N/A N/A Ranunculus montanus N/A Nograsek (1990)
Comoclathris typhicola Brown 3 transverse septa and 1 vertical septum N/A Typha latifolia, Typha angustifolia Austria, Denmark, England, Germany, India, Iran, Netherlands, Pakistan, Poland, Portugal, Russia, Switzerland, and USA Cooke (1872); Webster and Lucas (1959); Shoemaker and Babcock (1992); Boerema et al. (2004); Ahmadpour et al. (2024)
Comoclathris utahensis Dark reddish-brown 3 transverse septa and 1 vertical septum With a sheath 1–3 μm wide Arternisia scopulorurn, Brickelliaun bellata, Castilleja rniniata, Eupatoriurn occidentale, Linurn perenne, Linum perenne L., Polygonurn arnphibiurn, Senecio sp., Silene lnontana USA Shoemaker and Babcock (1992)
Comoclathris verrucosa Dark reddish-brown 3 transverse septa and 1 vertical septum With a uniform sheath 2–3 μm wide Juncus balticus USA Shoemaker and Babcock (1992)
Comoclathris verruculosa Mid reddish-brown 5 transverse septa and 1 vertical septum With a uniform sheath 1.5–3.5 μm wide Deschampsia caespitosa, Poa glauca Finland, Norway, Switzerland Shoemaker and Babcock (1992)
Comoclathris wehmeyeri N/A N/A N/A Crossandra infundibuliformis India Pande (1980)
Comoclathris xanthoceratis Brown to dark brown 3 transverse septa and 1 vertical septum Surrounded by a thick mucilaginous sheath Xanthoceras sorbifolium China Xu et al. (2024)
Comoclathris xerophila Mid reddish-brown 3 transverse septa and 1 vertical septum N/A Scleropogon brevifolius Argentina Shoemaker and Babcock (1992)
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Discussion

Comoclathris is a widely distributed genus, recorded on all five continents viz. Asia, Europe, Africa, the Americas, and Antarctica (Mattoo et al. 2023; Xu et al. 2024). Recent studies in China and other regions have shown that Comoclathris species are distributed in a variety of environments and substrates, from humid forests to semi-arid deserts (Mattoo et al. 2023; Xu et al. 2024; this study), suggesting that their diversity and ecological adaptability may be underestimated. The discovery of C. desertica and C. xiangshawanensis in desert wood in Inner Mongolia further expands the known distribution of this genus, providing valuable data for the global taxonomic system of Comoclathris. Furthermore, the thick, gelatinous sheaths enclosing the ascospores of these two species highlight the adaptability to extremely arid environments of these species. This is in line with the general characteristics of this genus, with obvious mucilaginous sheaths surrounding the ascospores. Based on morphological information currently available for this genus (Table 2), most species have mucilaginous sheaths, with only seven species having no record of mucilaginous sheaths (viz., C. acuminata, C. antarctica, C. galatellae D. Pem, Bulgakov & K.D. Hyde, C. sisyrinchii (Speg.) Shoemaker & C.E. Babc., C. sororia (Bubák) Shoemaker & C.E. Babc., C. typhicola (Cooke) H.A. Ariy. & K.D. Hyde, and C. xerophila (Speg.) Shoemaker & C.E. Babc.) (Cooke 1872; Shoemaker and Babcock 1992; Hongsanan et al. 2020; Crous et al. 2021; Mattoo et al. 2023).

Although 52 epithets are listed in Index Fungorum (2026), only 21 species have molecular data in NCBI, and the naming of many historical species is based solely on morphological characteristics. The lack of DNA sequences, type specimen revisions, and living cultures hinder phylogenetic analysis and species delineation. Therefore, future work should collect more specimens and supplement them with molecular data of extant species to clarify species boundaries and evolutionary relationships.

Inner Mongolia is a typical semi-arid region where long-term grazing and desertification have led to the degradation of grasslands and soils. Fungi play a crucial role in the decomposition of organic matter and nutrient cycling in fragile ecosystems, but the diversity of saprobic fungi remains poorly understood. The discovery of two new Comoclathris species in the Xiangshawan desert demonstrates that even degraded or arid habitats can foster rich and unique fungal communities. These fungal groups reflect the potential ecological resilience and adaptive evolution of microfungi under extreme temperature and humidity fluctuations. This study not only enriches the known fungal diversity in northern China but also highlights the importance of ongoing mycological surveys in arid grassland and desert ecosystems for taxonomic progress and ecological conservation.

Supplementary Material

XML Treatment for Comoclathris desertica
XML Treatment for Comoclathris xiangshawanensis

Acknowledgements

Xing-Guo Tian thanks the Science and Technology Foundation of Guizhou Province (Qian Ke He Pingtai ZSYS [2025] 029) and the Science and Technology Program of Guizhou Province (No. Qian Ke He Jichu QN [2025] 215). Dan-Feng Bao thanks the Postdoctoral Fellowship Program of CPSF under Grant Number GZC20240346 and Guizhou Provincial Basic Research Program (Natural Science) (No. QianKeHe Basic -MS [2025] general program 666) for their support. Rajesh Jeewon would like to thank the University of Mauritius, the New Drug Discovery and Evaluation Center for international cooperation and disciplinary innovation (“111 Center”) at Zunyi Medical University, and the PIFI (2026PVB0047) by CAS, Kunming Institute of Botany, China. Samantha C. Karunaratha and Saowaluck Tibpromma thank the High-Level Talent Recruitment Plan of Yunnan Province (High-End Foreign Experts program), the Key Laboratory of Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River, and the National Natural Science Foundation of China (Number 32260004) for their support. Shaun Pennycook is thanked for his assistance in selecting species epithets for the new species. Li-Juan Zheng and Jun-Yuan Zhang were thanked for their assistance in sampling.

Citation

Tian X-G, Du T-Y, Ma A-D, Jeewon R, Karunarathna SC, Tibpromma S, Bao D-F (2026) Taxonomic and phylogenetic analyses reveal two new Comoclathris taxa (Dothideomycetes, Pleosporaceae) from Inner Mongolia, China. MycoKeys 129: 107–128. https://doi.org/10.3897/mycokeys.129.179192

Funding Statement

The Science and Technology Program of Guizhou Province (No. Qian Ke He Jichu QN [2025] 215), the Postdoctoral Fellowship Program of CPSF under Grant Number GZC20240346, Guizhou Provincial Basic Research Program (Natural Science) (No. QianKeHe Basic -MS [2025] general program 666)

Additional information

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

The Science and Technology Program of Guizhou Province (No. Qian Ke He Jichu QN [2025] 215), the Postdoctoral Fellowship Program of CPSF under Grant Number GZC20240346, Guizhou Provincial Basic Research Program (Natural Science) (No. QianKeHe Basic -MS [2025] general program 666).

Author contributions

Conceptualization: XGT. Data curation: ADM, TYD. Formal analysis: XGT, RJ. Funding acquisition: DFB. Investigation: TYD. Methodology: XGT, TYD. Project administration: DFB, XGT. Resources: TYD. Software: ADM, TYD. Supervision: RJ, SCK, ST. Validation: RJ, DFB. Visualization: TYD, XGT. Writing – original draft: XGT. Writing – review and editing: RJ, ST, SCK.

Author ORCIDs

Xing-Guo Tian https://orcid.org/0000-0002-8124-1281

Tian-Ye Du https://orcid.org/0000-0003-2105-1803

An-Dong Ma https://orcid.org/0009-0005-5981-3182

Rajesh Jeewon https://orcid.org/0000-0002-8563-957X

Samantha C. Karunarathna https://orcid.org/0000-0001-7080-0781

Saowaluck Tibpromma https://orcid.org/0000-0002-4706-6547

Dan-Feng Bao https://orcid.org/0000-0002-5697-4280

Data availability

All of the data that support the findings of this study are available in the main text.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Comoclathris desertica
mycokeys.129.179192-treatment1.xml (18.9KB, xml)
XML Treatment for Comoclathris xiangshawanensis
mycokeys.129.179192-treatment2.xml (92.8KB, xml)

Data Availability Statement

All of the data that support the findings of this study are available in the main text.

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