Abstract
Xylariales (Sordariomycetes, Ascomycota) comprise a wide range of species that exhibit considerable variation in stromatic characteristics, including conspicuous to inconspicuous perithecia and unitunicate asci. Most known species are endophytes and saprobes, recognized for producing secondary metabolites of fundamental importance in the pharmaceutical and chemical industries. The main objectives of this study were to identify novel species, document new host and geographical records within the families Diatrypaceae, Hypoxylaceae, and Xylariaceae in northern and central Thailand, and explore the bioactive properties of secondary metabolites produced by selected Xylariales species. Taxa were identified through morphological examination, supported by phylogenetic analyses using maximum likelihood and Bayesian inference based on LSU, ITS, rpb2, and β-tub gene sequences. These taxa are accompanied by comprehensive descriptions and illustrations. Xylariales cultures were screened for preliminary antibacterial activity against the bacterial pathogens Bacillussubtilis (Gram-positive) and Escherichiacoli (Gram-negative). Based on the screening results, two newly introduced species (Annulohypoxylonchiangraiense and Hypoxylonthailandicum) and two known species (Xylariachrysanthum and Daldiniaeschscholtzii), which exhibited antibacterial activity, were selected for secondary metabolite extraction. Crude extracts from these isolates were chemically profiled using high-performance liquid chromatography (HPLC) and Q-TOF analyses, revealing a variety of potential compounds. The present study enhances our understanding of the taxonomic diversity and bioactive potential of Xylariales by introducing five new species, reporting nine new host records—including one new geographical record—and evaluating the bioactive properties of selected Xylariales cultures.
Key words: 5 new species, Ascomycota, morphology, multi-gene phylogeny, preliminary screening test
Introduction
Xylariales was established by Nannfeldt (1932) with Xylariaceae as the type family, along with Diatrypaceae, Hypocreaceae, Hyponectriaceae, Lasiosphaeriaceae, and Polystigmataceae. Initially, Xylariales species were mainly classified based on morphological characters (Hawksworth et al. 1995). However, the incorporation of molecular data has since diversified the classification criteria (Eriksson et al. 2003). Smith et al. (2003) conducted the first multi-gene analysis and identified seven families within this order. Lumbsch and Huhndorf (2010) listed six families, while Senanayake et al. (2015) included eleven families in Xylariales. Based on morphology and multi-gene analyses, Hyde et al. (2020a) accepted fifteen families in Xylariales. Fasciatisporaceae was introduced by Hyde et al. (2020a) to accommodate Fasciatispora within Xylariales, which comprises twenty families as outlined by Wijayawardene et al. (2022). However, some taxa in this order are considered genera incertae sedis due to the uncertainty of their taxonomic positions (Hyde et al. 2020a). Currently, Xylariales is considered the second-largest order in Xylariomycetidae, with 160 genera placed at the family level and 52 in genera incertae sedis (Hyde et al. 2020b; Phukhamsakda et al. 2020; Konta et al. 2021). Recently, Samarakoon et al. (2022) revised the taxonomy of xylarialean taxa based on morphology coupled with molecular phylogeny and accepted 57 genera incertae sedis in Xylariales. Hyde et al. (2024) and Thiyagaraja et al. (2025) included 22 families, expanding the taxonomic framework of Xylariales.
Xylariales species possess both conspicuous and inconspicuous fruiting bodies and are unitunicate, perithecial ascomycetes (Smith et al. 2003). This is a diverse group of fungi with distinct stromatic characteristics, which play a major role in generic and family-level classification (Jayawardena et al. 2022; Phukhamsakda et al. 2022; Samarakoon et al. 2022; 2023). In some xylarialean taxa, a distinct morphological character called the clypeus is present. It consists of stromatic tissues or melanized hyphae that develop above partially submerged or immersed ascomata. The clypeus forms a shield-shaped structure with variable development (Læssøe and Spooner 1993). Xylariales species occur as saprobes and endophytes in temperate, subtropical, and tropical regions worldwide, associated with wood, fallen fruits or seeds, fallen leaves or petioles, and termite nests (U’Ren et al. 2016; Fournier et al. 2018).
Within Xylariales, Xylariaceae and Hypoxylaceae are the most well-known families, producing secondary metabolites significant to the pharmaceutical and chemical industries. Some endophytic Xylariales species, such as Hypoxylonrubiginosum and Xylariacf.curta, are used for biological control due to their strong antagonistic effects against fungal and other pathogens (Halecker et al. 2020; Becker and Stadler 2021; Chen et al. 2024). Xylariaceae is considered one of the largest and most diverse families in Xylariales and comprises 42 genera and nearly 852 species (Hyde et al. 2024). Xylaria, the largest genus in Xylariaceae, was introduced with X.hypoxylon as the type species (Peršoh et al. 2009). Most Xylariaceae species are endophytes or saprobes, although a few have been reported as pathogens (Rogers 2000; Okane et al. 2008; Stadler et al. 2013; Husbands et al. 2018; Pourmoghaddam et al. 2022). These species can be found in wood, leaves, fruits, seeds, dung, and soil (Karimi et al. 2024). The family is characterized by embedded, well-developed ascomata and dark-colored stromata, which may be reduced or absent. The asci are 8-spored, unitunicate, and cylindrical and may possess an amyloid apical ring. The ascospores are pigmented and exhibit germ slits or pores (Rogers 2000). The asexual morph is characterized by holoblastic conidia and sympodially or occasionally percurrently proliferating conidiogenous cells (Rogers 2000).
Hypoxylaceae comprises 19 genera and approximately 422 species (Hyde et al. 2024). Within this family, Annulohypoxylon and Hypoxylon are the largest genera, comprising 69 and 235 species, respectively (Karimi et al. 2024). Hypoxylaceae species have a cosmopolitan distribution and occur as saprobes, endophytes, and pathogens (Reyes et al. 2024). This family is characterized by erect, glomerate, pulvinate, discoid, effused-pulvinate, hemispherical, spherical, or peltate stromata, which may be solitary or confluent, brightly colored, dark or black, pruinose, or smooth. Some stromata can produce extractable pigments visible in 10% KOH (Reyes et al. 2024). The perithecia are spherical, obovoid, or tubular, with spherical, umbilicate, or papillate ostioles, with or without discs formed by dehiscence of the surrounding tissue. A nodulisporium-like asexual state, distinguishable from that in Xylariaceae, occurs in Hypoxylaceae species (Reyes et al. 2024).
In Xylariales, diatrypaceous taxa are abundant and widely distributed (Ma et al. 2023). Diatrypaceae species occur as saprobes, endophytes, and pathogens on a wide range of crops and woody plants (Vasilyeva and Ma 2014; Konta et al. 2020a; Samarakoon et al. 2022; Li et al. 2023). Members of this family can produce extracellular, ligninolytic enzymes that degrade plant cell walls, playing an important role in wood decomposition (Bucher et al. 2004; Trouillas et al. 2011; Mehrabi et al. 2015). Diatrypaceae species are characterized by eustromatic or pseudostromatic, black or dark brown, erumpent to immersed, and occasionally superficial stromata. Ostioles are present in the perithecia. The asci are 8-spored or polysporous, occasionally with one or two spores, and are unitunicate. The ascospores are ellipsoidal, globose, filiform, or allantoid, and hyaline to light brown (Senanayake et al. 2015). The asexual morph is characterized by acervulus-like subcortical, erumpent conidiomata. The conidia are hyaline, filiform, curved, or rarely straight (Senanayake et al. 2015; Wijayawardene et al. 2017; Phukhamsakda et al. 2022). Currently, Diatrypaceae comprises 31 genera (Hyde et al. 2024). Given the widespread occurrence and varied lifestyles of Xylariales species, further taxonomic and ecological studies are essential.
Within Xylariales, the families Hypoxylaceae and Xylariaceae are the most prolific producers of secondary metabolites (Becker and Stadler 2021). Numerous unique secondary metabolites have been discovered from the stromata and mycelial cultures of Xylariales fungi, many of which have potential pharmaceutical and agrochemical applications (Becker and Stadler 2021). For example, Kretzschmariazonata, a plant pathogenic fungus, produces various enzymes such as β-glucosidases, endoglucanases, hemicellulases, pectinases, and xylanases (da Luz Morales et al. 2021). In Xylariaceae, Xylaria species have yielded a wide range of bioactive compounds, including alkaloids, aromatic compounds, cytochalasins, polyketides, and terpenoids. These compounds exhibit diverse biological activities, including antibacterial, antifungal, anticancer, antimalarial, anti-inflammatory, and α-glucosidase inhibitory activities (Chen et al. 2024). Additionally, Whalley and Edwards (1995, 1998) reported that many chemical metabolites produced by Xylariaceae cultures exhibit a degree of genus specificity.
Beyond their bioactivity, chemotaxonomic data support the segregation of genera within Hypoxylaceae, such as Annulohypoxylon, Hypomontagnella, Hypoxylon, and Jackrogersella (Becker et al. 2020; Cedeño-Sanchez et al. 2024). Species in Hypoxylaceae are known to produce a diverse array of secondary metabolites, particularly azaphilones and binaphthalenes (Kuhnert et al. 2021). To date, more than 200 metabolites with various bioactivities have been identified from Hypoxylon species, including hypoxyloamide, 8-methoxynaphthalene-1,7-diol, and hypoxylonol, which exhibit antimicrobial and anticancer activities (Tan and Zou 2001; Cheng et al. 2020). The growing focus on chemotaxonomic studies of Xylariales species is likely to uncover many previously unknown secondary metabolites with significant bioactive potential.
The main objectives of this study are to introduce five novel taxa, report nine new host records including one new geographical record of species belonging to Diatrypaceae, Hypoxylaceae, and Xylariaceae in Xylariales, and to investigate bioactive compounds produced by cultures of selected Xylariales species. Morphological characteristics and multi-gene phylogenetic analyses using maximum likelihood (ML) and Bayesian inference (BI) confirmed the phylogenetic placement of the studied fungal taxa within the order Xylariales. Preliminary antibacterial screening was conducted for selected species, including newly described taxa in Annulohypoxylon and Hypoxylon. Additionally, bioactive compounds from the cultures of these species were analyzed using HPLC/Q-TOF techniques.
Materials and methods
Specimen collections, morphological studies, and isolations
Specimens (dead wood) were collected during March–July 2024 in Thailand. Samples were enclosed in zip-lock plastic bags and transported to the laboratory. Observations followed the methodology of Senanayake et al. (2020). Morphological characteristics were examined using a LEICA EZ4 stereomicroscope (Leica Microsystems, Germany) and an AXIOSKOP 2 PLUS compound microscope (Carl Zeiss Microscopy, Germany). Microscopic structures were photographed with a Canon 550D digital camera mounted on the microscope. Melzer’s reagent, Congo red, and Indian ink were used as needed. Measurements were made using ZEN2 (Blue Edition) software and calculated with the Tarosoft® Image Framework program. Figures and photo plates were assembled using Adobe Photoshop CS3 Extended version 10.0 (Adobe Systems, USA).
Single-spore isolations were carried out on potato dextrose agar (PDA) following Senanayake et al. (2020). Herbarium specimens were deposited at the Mae Fah Luang University Herbarium (MFLU), Chiang Rai, Thailand, and living cultures were preserved in the Mae Fah Luang University Culture Collection (MFLUCC). For newly introduced taxa, Faces of Fungi numbers were obtained according to Jayasiri et al. (2015), and Index Fungorum numbers were registered via Index Fungorum (2025). All data generated in this study were deposited in the Greater Mekong Subregion Fungal Database (Chaiwan et al. 2021).
DNA extraction, PCR amplification, and sequencing
Genomic DNA was extracted from 50–100 mg of fungal mycelium using the E.Z.N.A Fungal DNA Mini Kit (D3390-02, Omega Bio-Tek, USA) according to the manufacturer’s instructions. Extracted DNA was stored at 4 °C for short-term use and at -20 °C for long-term storage. Polymerase chain reactions (PCR) were performed for the large subunit rDNA (LSU), the internal transcribed spacer region (ITS), β-tubulin (β-tub), and RNA polymerase II second largest subunit (rpb2) gene regions as described in previous studies (Thiyagaraja et al. 2019; Konta et al. 2020a; Karimi et al. 2023).
The LSU, ITS, β-tub, and rpb2 gene regions were amplified using the primers LR0R/LR5 (Vilgalys and Hester 1990; Rehner and Samuels 1994), ITS4/ITS5 (White et al. 1990), T1/T22 (O’Donnell and Cigelnik 1997), Bt2a/Bt2b (Glass and Donaldson 1995), and fRPB2-5f/fRPB2-7cR (Liu et al. 1999), respectively. PCR reactions were carried out in a final volume of 50 μl, consisting of 25 μl of 2× Power Taq PCR Master Mix, 1 μl of each forward and reverse primer, 2 μl of genomic DNA, and 21 μl of deionized water. The PCR products were visualized on 1.5% agarose gels, stained with 4S Green Stain, and subsequently sequenced at SolGent Co., Ltd. (South Korea). Newly generated nucleotide sequences were deposited in GenBank (Table 1).
Table 1.
Taxa used in the phylogenetic analysis and their GenBank accession numbers. New isolates generated in this study are in bold, and type strains are indicated in superscript ‘T’.
N/A- Sequences not available; “–” - Sequences not used for analyses.
Phylogenetic analyses
The quality of the sequence chromatograms was checked using BioEdit v. 7.0.9.0 (Hall 1999). Forward and reverse sequences were assembled into consensus sequences using Lasergene SeqMan Pro v. 7. Newly generated sequences were searched using the BLASTn search engine at NCBI (https://www.ncbi.nlm.nih.gov) against the GenBank database, and related literature was referred to (Thiyagaraja et al. 2019; Karimi et al. 2023). Each locus (LSU, ITS, β-tub, and rpb2) was individually aligned using MAFFT 6.864b (Katoh et al. 2019), trimmed using trimAl v. 1.2 software (Capella-Gutiérrez et al. 2009), and manually adjusted for improvement where necessary using BioEdit v. 7.2 (Hall 1999). Single gene alignments and the concatenated aligned dataset were analyzed separately using ML and BI. Best-fit models for BI analyses were selected using MrModeltest v. 2.2 (Nylander 2004) under the AIC (Akaike Information Criterion) implemented in PAUP v. 4.0b10. The GTR+G model was selected as the best model for BI analyses for all gene regions.
The ML analyses were performed using IQ-TREE with bootstrap support obtained from 1,000 pseudoreplicates (Nguyen et al. 2015; Chernomor et al. 2016). The BI analyses were conducted with MrBayes v. 3.2.6 (Ronquist et al. 2012). The Markov Chain Monte Carlo (MCMC) algorithm of six chains was initiated for 1,000,000 generations. The trees were sampled at every 100th generation, resulting in 10,000 trees. The first 10% of trees were discarded as the burn-in phase, while the remaining 90% were used to calculate the posterior probabilities (PP) in the majority rule consensus tree. Phylograms were visualized in the FigTree v. 1.4.0 program (Rambaut 2012) and reorganized in Microsoft PowerPoint (2010).
Preliminary screening for antibacterial activity
Preliminary screening for antibacterial activity was conducted for new isolates from the present study, along with selected existing Xylariales cultures from the Mae Fah Luang University Culture Collection (MFLUCC), following the methods described by Mapook et al. (2020). Ampicillin antibacterial discs were used as a positive control for the screening tests (Alam et al. 2019). The agar plug diffusion method was employed to assess antibacterial activity against Bacillussubtilis (gram-positive bacteria) and Escherichiacoli (gram-negative bacteria) (Balouiri et al. 2016). Both bacterial strains were cultured on nutrient agar (NA) for 24 hours. Prior to inoculating the Mueller-Hinton agar medium, bacterial cell concentrations were determined using a hemocytometer (6.7 × 105 cells/mL), as described by Mapook et al. (2020). Fungal mycelial plugs from the test samples were transferred onto Mueller-Hinton agar plates and incubated at room temperature (25 °C) for 24 hours. Inhibition zones were measured and compared with both the positive and negative controls.
Chemical extraction and HPLC/Q-TOF analyses
Xylariales cultures that showed positive results in the antibacterial activity assay were selected for fermentation and crude extraction. Culture conditions for the selected fungi were optimized using potato dextrose broth (PDB; potatoes, infusion from 200 g/L; dextrose, 20 g/L; pH 5.1 ± 0.2), yeast malt broth (YMB; dextrose, 10 g/L; malt extract, 3 g/L; peptic digest of animal tissue, 5 g/L; yeast extract, 3 g/L; pH 6.2 ± 0.2), and malt extract broth (MEB; malt extract, 30 g/L; mycological peptone, 5 g/L; agar, 15 g/L; pH 5.4 ± 0.2). Five mycelial plugs were cut from freshly grown cultures on each of the three media using a sterilized cork borer. These plugs were inoculated into 250 mL of the respective liquid medium in 500 mL Erlenmeyer flasks. The flasks were incubated on an Innova rotary shaker 43/43R (Eppendorf, Germany) at 140 rpm at 23 °C for 3–5 days. During fermentation, glucose depletion was monitored daily using Bayer Harnzuckerstreifen glucose test strips (Bayer, Leverkusen, Germany), and pH levels were measured using litmus paper (Merck KGaA, Darmstadt, Germany). When glucose levels reached zero and the pH dropped below 7.0, the cultures were processed for extraction.
Fungal mycelium and supernatant were separated by vacuum filtration. Crude extracts were prepared from both fractions. For the supernatant, ethyl acetate extraction was performed three times by mixing it with an equal volume of ethyl acetate in a separatory funnel. An equal volume of acetone was added to the mycelium, which was then freeze-dried and extracted three times with methanol (MeOH) at 40 °C in an ultrasonic bath for 30 minutes (Hellwig et al. 2005). After centrifugation at 1000 × g for 10 minutes, the supernatants were evaporated, and 50–100 mL of deionized water was added. Ethyl acetate extraction was then performed three times on the mycelium using an equal volume of ethyl acetate to yield a crude extract. These extracts were weighed.
Crude extracts obtained from both supernatant and mycelium were dissolved in methanol to a final concentration of 1 mg/mL. The solutions were filtered through a 0.22 µm membrane filter to remove particulates prior to HPLC injection. High-performance liquid chromatography (HPLC) was conducted using a Waters ACQUITY Arc System. Detection was performed with a photodiode array (PDA) detector and a fluorescence (FLR) detector (Waters, USA) using a reverse-phase C18 column (Kinetex® 5 µm EVO C18 100 Å, 150 × 2.1 mm LC) maintained at 30 °C. The mobile phase consisted of water with 0.2% formic acid (A) and methanol (B). Elution was carried out at a flow rate of 0.2 mL/min using the following linear gradient: 5–30% B (0–8 min), 30% B (8–10 min), 30–95% B (10–18 min), 95% B (18–22 min), 95–5% B (22–25 min), and 5% B (25–32 min). Samples were analyzed using a photodiode array detector set at 256 and 425 nm. Data analysis was conducted using Empower 3 software.
Liquid chromatography–quadrupole time-of-flight mass spectrometry (LC-QTOF-MS) was performed using an Agilent II/G6545B QTOF/MS and 1290 Infinity system equipped with an electrospray ionization (ESI) source. The instrument was operated in both positive and negative ionization modes under UHPLC pressure (1,300) and a mass range of 100–10,000.
Results
. Astrocystis
Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 123 (1873) [1875]
E52E5287-8EFF-559F-8D79-FC67E4A3E461
Index Fungorum: IF439
Facesoffungi Number: FoF00420
Notes.
Berkeley and Broome (1875) introduced Astrocystis, with A.mirabilis as the type species. Morphologically, this genus is characterized by uni- or occasionally multi-peritheciate stromata development, often beneath the host cuticle or on the surface; relatively short stipe asci; and ascus apical apparatus that are relatively small, amyloid, and stopper-shaped (Smith et al. 2003). Currently, 42 records are available in the Index Fungorum (2025).
Phylogenetic analyses for Astrocystis
For Astrocystis, 32 taxa were included in the combined data set (ITS, β-tub, and rpb2) with Xylotumulusgibbisporus (ATCC MYA-4109), Xylariaglebulosa (GMB1053), and X.schweinitzii (HAST 92092023) as outgroup taxa. The final alignment consisted of 1953 characters, including gaps (ITS = 435 bp, β-tub = 514 bp, and rpb2 = 1004 bp). Both ML and BI analyses exhibit similar tree topology. The best-scoring RAxML tree was obtained (Fig. 1), with a final likelihood value of -12708.0675. The matrix included 851 distinct alignment patterns, with 17.88% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.242776, C = 0.267065, G = 0.262377, and T = 0.227782; substitution rates were AC = 1.393798, AG = 4.050932, AT = 1.300292, CG = 1.282338, CT = 7.225689, and GT = 1.0; and the gamma distribution shape parameter α = 0.297082. In the BI analyses, the average standard deviation of the split frequencies was 0.006 after 1,000,000 generations of runs. The phylogenetic tree topology is similar to the previous study by Li et al. (2024). According to the phylogenetic analyses, our strain MFLUCC 25-0022 clades within Astrocystis, with Astrocystisbambusae strains (HAST 8902190 and GMB0700).
Figure 1.
Phylogram generated from ML analysis based on the combined dataset of ITS, β-tub, and rpb2. The tree is rooted to Xylotumulusgibbisporus (ATCC MYA-4109), Xylariaglebulosa (GMB1053), and X.schweinitzii (HAST 92092023). Bootstrap support values for ML ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.90 are noted at the node. Strain numbers are noted after the species names. Strains isolated in this study are represented in blue, and type strains are in bold.
Taxonomy
. Astrocystis bambusae
(Henn.) Læssøe & Spooner, Kew Bull. 49(1): 13 (1994) [1993]
5B8DF55F-6917-52A6-9668-B5754B8EF78E
Index Fungorum: IF361739
Facesoffungi Number: FoF17292
Figure 2.
Astrocystisbambusae on a dead twig of Bambusavulgaris (MFLU 24-0522, a new host record). a. Substrate; b, c. Appearance of stromata on the host; d. Cross section of the stroma; e. Peridium; f. Paraphyses; g–j. Asci; k. Ascus apical apparatus (stained in Melzer’s reagent); l–q. Ascospores; r. Ascospores with sheath; s, t. Colony on the PDA (s upper, t lower). Scale bars: 5 mm (b); 1 mm (c); 500 μm (d); 20 μm (f–j); 10 μm (e, k–r).
Rosellinia bambusae Henn. 1908. Basionym.
Description.
Saprobic on dead culms of Bambusavulgaris. Sexual morph: Stromata 1.2–0.9 mm diam., 0.8–1 mm high, scattered, solitary, superficial, black, appear as black raised spots on the host surface, hexagonal prism-shaped, containing one ascoma, with a circle of black tissue at the bottom. Perithecia 0.5–0.7 mm diam., 0.4–0.5 mm high, comprising black, fragile, carbonaceous tissue. Peridium 15–45 μm wide, 5–8 layers, brown to dark brown cells of textura angularis. Hamathecium comprising 2–8 μm wide, oblong to cylindrical, septate, unbranched, cellular, paraphyses. Asci 55–90 × 5–6.5 µm (x̄ = 75 × 6.2 µm, n = 30), 8- or 6-spored, unitunicate, cylindrical, short pedicellate, persistent, apically rounded, with amyloid, cuboid, apical apparatus, staining blue in Melzer’s reagent, 2–3 µm high × 1.4–2.44 μm wide (x̄ = 2.6 × 1.9 μm). Ascospores 10–13.5 × 4–6 µm (x̄ = 12 × 5 μm, n = 30), uniseriate, unicellular, hyaline when immature, dark brown at maturity, aseptate, equilateral ellipsoid, with rounded ends, smooth, guttulate, with a straight germ slit nearly full-length, surrounded by a sheath. Asexual morph: undetermined.
Culture characteristics.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on PDA reaching 2–2.5 cm diam. after 5 days at 25 °C, circular in shape, white at first, cottony, slightly thinning towards the edge, white color in the front view, and light brown in the reverse view.
Material examined.
Thailand • Chiang Rai, Mae Chan, Mae Chan District, on dead culms of Bambusavulgaris (Poaceae), 18 March 2024, Hsan Win (MFLU 24-0522), living culture MFLUCC 25-0022.
Known distribution and hosts.
China, India, Thailand (Bambusa sp.); Ghana (Oxytenantheraabyssinica); Philippines (Bambusa sp., Schizostachyum sp.) (Læssøe and Spooner 1993; Li et al. 2024); Thailand (Bambusavulgaris) (this study).
Notes.
Morphologically, our collection (MFLUCC 25-0022) exhibits characteristics similar to the holotype of A.bambusae (basionym: Roselliniabambusae) (Merrill 5030) and other isolates of A.bambusae (GMB0700). These similarities include scattered, solitary, superficial, black stromata containing one ascomata, with a circle of black tissue at the bottom and unitunicate, cylindrical, short pedicellate asci, with an apical apparatus that stains blue in Melzer’s reagent (Li et al. 2024). The ascospores have a straight germ slit nearly full-length and are surrounded by a sheath (Ju and Rogers 1990; Li et al. 2024). However, the asci and ascospores in our collection (MFLUCC 25-0022, 55–90 μm and 10–13.5 μm, respectively) are smaller than the holotype (100–130 μm and 10.5–15(–16) μm) (Ju and Rogers 1990). Based on multi-gene phylogenetic analyses (ITS, β-tub, and rpb2), our strain (MFLUCC 25-0022) clustered with other authentic strains (HAST 89021904 and GMB0700) in a well-supported clade (89% ML, 0.95 BYPP) (Fig. 1). Astrocystisbambusae has previously been recorded on Bambusa sp. in China, India, the Philippines, and Thailand (Ju and Rogers 1990; Læssøe and Spooner 1993; Li et al. 2024). In our study, we reported a new host record for A.bambusae on Bambusavulgaris.
. Annulohypoxylon
Y.M. Ju, J.D. Rogers & H.M. Hsieh, Mycologia 97(4): 855 (2005)
806C1BA3-7232-544D-A1F9-660E6FD1ED71
Index Fungorum: IF500298
Facesoffungi Number: FoF02983
Notes.
Annulohypoxylon was introduced with A.truncatum as the type species (Hsieh et al. 2005). Kuhnert et al. (2017) conducted a concise revision of this genus based on molecular phylogeny and chemotaxonomic data, resulting in the identification of several additional species, such as A.massivum, A.violaceopigmentum, A.viridistratum, and A.yungensis. Annulohypoxylon is characterized by effused-pulvinate or pulvinate, glomerate stromata, waxy or carbonaceous tissue immediately beneath the surface and between perithecia, spherical, obovoid, with a carbonaceous stromata layer surrounding individual perithecia. Asci are light- to dark-colored, 8-spored, cylindrical, stipitate, persistent, with discoid apical ring, amyloid or infrequently inamyloid, while ascospores are light- to dark-colored, ellipsoid or short fusoid, inequilateral, narrowly rounded, or with broadly rounded ends, with a germ slit, perispore dehiscent or indehiscent in 10% KOH (Li et al. 2016). Annulohypoxylon species have mainly been recorded in tropical and subtropical regions as saprobes associated with dead dicotyledonous wood and as endophytes in seed plants (Kuhnert et al. 2017). Hyde et al. (2024) listed 60 species under this genus, while 73 species are included in the Index Fungorum (2025).
Phylogenetic analyses for Annulohypoxylon
Forty-six taxa of Annulohypoxylon were included in the combined data set (ITS, LSU, β-tub, and rpb2) with Biscogniauxiapetrensis (HKAS102388) as the outgroup taxon. After alignment, the dataset comprised 2832 characters, including gaps (ITS = 585 bp, LSU = 850 bp, β-tub = 380 bp, rpb2 = 1017 bp). The tree topology of the BI analysis (not shown) was similar to the ML tree. The best-scoring RAxML tree was obtained (Fig. 3), with a final likelihood value of -19758.666775. The matrix had 1149 distinct alignment patterns with 46.81% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.248997, C = 0.254092, G = 0.262808, T = 0.234103; substitution rates AC = 1.551284, AG = 4.096946, AT = 1.717695, CG = 1.101636, CT = 7.150044, GT = 1.0; gamma distribution shape parameter α = 0.220052. In BI analyses, the average standard deviation of split frequencies was 0.008 after 3,000,000 generations of runs. The phylogenetic tree topology is similar to that by Kuhnert et al. (2017). Our strains (MFLUCC 24-0606, MFLUCC 24-0607, MFLUCC 24-0608, MFLUCC 24-0609, MFLUCC 24-0610, and MFLUCC 25-0023) cluster within Annulohypoxylon.
Figure 3.
Phylogram generated from ML analysis based on the combined dataset of ITS, LSU, β-tub, and rpb2. The tree is rooted to Biscogniauxiapetrensis (HKAS102388). Bootstrap support values for ML ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.90 are noted at the nodes. Strain numbers are noted after the species names. Strains isolated in this study are represented in blue, and type strains are in bold.
Taxonomy
. Annulohypoxylon bahnphadengense
J. Fourn. & M. Stadler, Fungal Diversity 40: 30 (2010)
F08BA9AC-8591-5E4C-AF60-0F9CCAF2F19E
Index Fungorum: IF512545
Facesoffungi Number: FoF17293
Figure 4.
Annulohypoxylonbahnphadengense on dead wood of Berryacordifolia (MFLU 24-0526, a new host record). a. Substrate; b, c. Appearance of ascostromata on host; d, e. Horizontal section through ascomata (arrow shows the ostiolar disc); f. Peridium; g. Paraphyses; h–m. Asci; n. Apical apparatus stained blue with Melzer’s reagent; o–q. Ascospores; r. Germinated ascospores; s, t. Colony on the PDA (s upper, t lower). Scale bars: 2 mm (b); 1 mm (c); 500 μm (d); 200 μm (e); 20 μm (f, g, i–m); 10 μm (h, r); 5 μm (n–q).
Description.
Saprobic on the dead wood of Berryacordifolia. Sexual morph: Ascostromata 3–14 × 2–10 × 0.5–0.2 mm (x̄ = 7 × 6 × 0.3 mm, n = 8), effused-applanate, superficial, pulvinate to hemispherical, clustered, hard-textured, shiny, surface black, carbonaceous. Ascomata 0.5–3.5 mm high × 0.3–0.5 mm diam. (x̄ = 2 × 0.4 mm, n = 15), immersed in the stroma, subglobose to globose, black, ostiolate, papillate, encircled with a flattened truncatum-type disc 0.2–0.25 mm diam. (x̄ = 0.22 mm, n = 10). Peridium 40–60 μm wide, composed of several layers of hyaline to dark brown cells of textura angularis. Hamathecium 4–6 μm wide, comprising long, hyaline, unbranched, septate paraphyses. Asci 54–130 × 3–5 μm (x̄ = 94 × 4.5 μm, n = 20), 8-spored, unitunicate, cylindrical, short pedicellate, with an apical ring bluing in Melzer’s reagent. Ascospores 6–8 × 3–4 μm (x̄ = 7.5 × 3.5 μm, n = 40), uniseriate, one-celled, inequilaterally ellipsoidal, with narrowly rounded ends, hyaline when immature, becoming light brown to dark brown at maturity, guttulate. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. Colonies on the PDA reaching 2.0–2.5 cm diam. after six days at 25 °C, circular in shape, white at first, cottony, white color in the front view, brown in the middle, and pale brown at the margin in the reverse view.
Material examined.
Thailand • Chiang Rai, Phan District, Sai Khao, forest area near Wat Udom Waree, on decaying wood of Berryacordifolia (Malvaceae), 05 July 2024, Achala Rathnayaka, AA28 (MFLU 24-0526); living culture MFLUCC 24-0608.
Known distribution and hosts.
China (decaying wood) (Ke et al. 2024); Thailand (dead bark or wood, Berryacordifolia) (Fournier et al. 2010b; this study).
Notes.
Morphologically, our collection (MFLUCC 24-0608) shows similar characteristics to the holotype of A.bahnphadengense (MFU08-1552), including shiny, black, carbonaceous ascostromata; 8-spored, cylindrical, short-pedicellate asci with an apical ring bluing in Melzer’s reagent; and uniseriate, one-celled, inequilaterally ellipsoidal, ascospores with narrowly rounded ends (Fournier et al. 2010b). According to the multi-gene phylogenetic analyses (ITS, LSU, β-tub, and rpb2), our strain (MFLUCC 24-0608) clusters with the ex-type strain of A.bahnphadengense (STMA 13115) with 89% ML bootstrap and 0.84 PP support (Fig. 3). Based on the morpho-molecular evidence, we identified our collection as a new host record of A.bahnphadengense on Berryacordifolia in Thailand.
. Annulohypoxylon chiangraiense
Rathnayaka, K.D. Hyde & Chethana sp. nov.
187AE288-FC89-5D75-875E-1B77BCD88780
Index Fungorum number: IF903883
Facesoffungi Number: FoF17288
Figure 5.
Annulohypoxylonchiangraiense on a dead branch of Tamarindusindica (MFLU 24-0524, Holotype). a. Sustrate; b, c. Stromata on the host; d. Cross section of the stroma; e. Peridium; f. Paraphyses; g–j. Asci; k. Ascal apical apparatus (not staining in Melzer’s reagent); l–o. Ascospores; p, q. Colony on the PDA (p upper, q lower). Scale bars: 5 mm (b); 1 mm (c); 500 μm (d); 20 μm (e–k); 10 μm (l–o).
Etymology.
The epithet chiangraiense refers to Chiang Rai Province, where the fungus was collected.
Holotype.
MFLU 24-0524.
Description.
Saprobic on the dead branch of Tamarindusindica. Sexual morph: Ascostromata 0.4–0.6 × 0.8–1.5 mm (x̄ = 0.5 × 1.2 mm, n = 10), semi-immersed to superficial, with the base immersed, pulvinate to hemispherical, solitary or clustered, spherical surface black, carbonaceous. Ascomata immersed in stroma, globose to subglobose, black. Peridium 18–30 μm wide, composed of several layers of hyaline to dark brown cells of textura angularis. Paraphyses 3–6 μm wide, hyaline, abundant, persistent, unbranched, septate. Asci 90–145 × 8–5 μm (x̄ = 124 × 7 μm, n = 20), 8-spored, unitunicate, cylindrical, short pedicellate, with an apical ring not bluing in the Melzer’s reagent (without KOH pretreatment). Ascospores 10–12 × 4–7 μm (x̄ = 11 × 5 μm, n = 30), uniseriate, one-celled, inequilaterally ellipsoidal, with narrowly rounded ends, hyaline at immature stages, dark brown when mature, guttulate. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on the PDA reaching 1.0–2.0 cm diam. after five days at 25 °C, circular in shape, white at first, cottony, slightly thinning towards the edge, with white color in the middle and pale yellow color in the margin of the front view, and pale yellow in the reverse view.
Material examined.
Thailand • near Nang Lae waterfall, Chiang Rai, on decaying wood of Tamarindusindica (Fabaceae), 18 March 2024, Achala Rathnayaka, AA11 (MFLU 24-0524, holotype); ex-type living culture, MFLUCC 24-0606.
Notes.
Annulohypoxylon is a speciose genus with more than 60 species; however, the present study shows the genus to be more diverse as predicted by Bhunjun et al. (2024). Based on the multi-gene phylogeny (ITS, LSU, β-tub, and rpb2), Annulohypoxylonchiangraiense (MFLUCC 24-0606) formed a distant lineage sister to A.archeri (SGNLB 5) and A.microdiscum (HMAS 285320) with 100% ML bootstrap and 1.00 PP support (Fig. 3). Annulohypoxylonchiangraiense fits within the generic concept of Annulohypoxylon by having spherical, carbonaceous ascostromata; 8-spored, cylindrical asci; and ellipsoid, light- to dark-brown ascospores (Li et al. 2016). Annulohypoxylonchiangraiense differs from both A.archeri and A.microdiscum by having smaller ascostromata (0.4–0.6 × 0.8–1.5 mm vs. 8–20 × 5–10 mm and 0.5–4 × 0.3–2 cm) (Raei et al. 2012; Cruz et al. 2020). The asci of A.chiangraiense are shorter and wider (90–145 × 8–5 μm) than A.microdiscum (130–187 × 5–6.5 μm). However, asci were not observed in A.archeri (Cruz et al. 2020). In A.chiangraiense, the apical ring does not turn blue in Melzer’s iodine reagent, whereas in A.microdiscum, the apical ring turns blue in Melzer’s iodine reagent (Raei et al. 2012). While ascospores of both A.archeri and A.microdiscum have a straight germ slit, such a character was not observed in the ascospores of A.chiangraiense (Raei et al. 2012; Cruz et al. 2020). When comparing the ITS base pair differences of A.chiangraiense with A.archeri and A.microdiscum, it shows 1.6% (8/564) and 1.9% (10/533) differences (without gaps), respectively. Based on the distinct morphology and phylogenetic evidence, we established Annulohypoxylonchiangraiense as a new species.
. Annulohypoxylon crowfoothodgkiniae
Y.P. Tan, Bishop-Hurley, Bransgr. & R.G. Shivas, Index of Australian Fungi 1: 1 (2022)
3BF3D690-588C-50BE-8D58-39E4C3204566
Index Fungorum: IF900010
Facesoffungi Number: FoF17294
Figure 6.
Annulohypoxyloncrowfoothodgkiniae on decaying wood of Swieteniamacrophylla (MFLU 24-0522, a new host and geographical record). a. Substrate; b, c. Stromata on the host; d. Cross section of the stroma; e. Paraphyses; f–j. Asci; k. Ascus apical apparatus (not stained in Melzer’s reagent); l–q. Ascospores; r, s. Colony on PDA (r upper, s lower). Scale bars: 5 mm (b); 1 mm (c); 200 μm (d); 20 μm (e–k); 5 μm (l–q).
Description.
Saprobic on dead wood of Swieteniamacrophylla. Sexual morph: Ascostromata 0.5–0.7 × 0.3–0.6 mm (x̄ = 0.6 × 0.5 mm, n = 10), semi-immersed to superficial, with the base immersed, pulvinate to hemispherical, clustered, shiny, surface black, carbonaceous. Ascomata immersed in the stroma, subglobose to hemispherical, black, ostiolate, papillate. Hamathecium 3–7 μm wide, comprising long, hyaline, unbranched, aseptate paraphyses. Asci 55–65 × 4–6 μm (x̄ = 60 × 4.5 μm, n = 20), 8-spored, unitunicate, cylindrical, short pedicellate, with an apical ring not bluing in the Melzer’s reagent. Ascospores 5–6 × 2–3 μm (x̄ = 5.4 × 2.8 μm, n = 30), uniseriate, one-celled, ellipsoidal inequilaterally, with narrowly rounded ends, hyaline when immature, becoming brown at maturity. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on the PDA reaching 2.0–2.5 cm diam. after five days at 25 °C, circular in shape, white at first, cottony, slightly thinning towards the edge, white color in the front view, and light brown in the reverse view.
Material examined.
Thailand • Nang Lae Village, Chiang Rai, on decaying wood of Swieteniamacrophylla (Meliaceae), 21 May 2024, Zaw Lin Tun, AZ2 (MFLU 24-0523); living culture MFLUCC 25-0023.
Known distribution and hosts.
Australia (Pandanustectorius) (Tan and Shivas 2022a); Thailand (Swieteniamacrophylla) (this study).
Notes.
In the multi-gene phylogeny (ITS, LSU, β-tub, and rpb2), our strain (MFLUCC 25-0023) and the ex-type strain of A.crowfoothodgkiniae (BRIP 72527 h) clustered with 91% ML bootstrap and 1.00 PP support (Fig. 3). When comparing the base pair differences between our strain (MFLUCC 25-0023) and the ex-type strain of A.crowfoothodgkiniae (BRIP 72527 h), the ITS shows a 0.3% (3/894) difference, and there are no differences in LSU. The morphology of the holotype is not recorded. Therefore, we could not compare the morphology between the holotype and our strain. In here, we provide complete morphology with an illustration for A.crowfoothodgkiniae. Based on molecular evidence, we introduce our collection as a new host record of A.crowfoothodgkiniae from Swieteniamacrophylla and also as a new geographical record from Thailand.
. Annulohypoxylon spougei
Suwannasai, M.P. Martín, Phosri & Whalley, Persoonia 44: 353 (2020)
5CC6F3EF-0165-5012-B3A7-96BFD7DB8A69
Index Fungorum: IF811164
Facesoffungi Number: FoF17295
Figure 7.
Annulohypoxylonspougei on dead wood of Antidesmamadagascariense (MFLU 24-0526, a new host record). a. Substrate; b, c. Appearance of ascostromata on the host; d. Ostiolar discs in ascomata (indicated by arrows); e. Horizontal section through the ascomata; f. Paraphyses; g-i. Asci; j. Apical apparatus stained blue with Melzer’s reagent; k-p. Ascospores; q. Germinated ascospores; r, s. Colony on the PDA (r upper, s lower). Scale bars: 2 mm (b); 500 μm (c, d); 200 μm (e); 20 μm (f); 10 μm (g–i, q); 5 μm (j–p).
Description.
Saprobic on Antidesmamadagascariense dead wood. Sexual morph: Ascostromata 1–3 cm long × 0.3–2 cm broad and 0.8–1.2 mm thick (x̄ = 2 × 1.4 × 1 mm, n = 10), hemispherical, effused-pulvinate, shiny, surface black, carbonaceous. Ascomata 0.25–0.6 mm high × 0.25–0.5 mm diam. (x̄ = 0.4 × 0.3 mm, n = 10), immersed in the stroma, subglobose to globose, black, ostioles papillate, encircled with a flattened, truncatum-type disc, 0.2–0.25 mm diam. (x̄ = 0.23 mm, n = 8). Hamathecium 3–5 μm wide, comprising long, hyaline, unbranched, septate paraphyses. Asci 27–42 × 2–3 μm (x̄ = 36 × 2 μm, n = 20), the spore-bearing parts 17–25 µm long with stipes 9–15 µm long, 8-spored, unitunicate, cylindrical, with an apical ring bluing in Melzer’s iodine reagent. Ascospores 7–9 × 3–4 μm (x̄ = 7.6 × 3.6 μm, n = 40), uniseriate, one-celled, inequilaterally ellipsoidal, with narrowly rounded ends, hyaline when immature, becoming brown at maturity, guttulate. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. Colonies on the PDA reaching 1.5–2.0 cm diam. after five days at 25 °C, circular in shape, white at first, cottony, white color in the front view, brown in the middle, and pale brown at the margin of the reverse view.
Material examined.
Thailand • Chiang Rai, near Ang Kep Nam Huai Luang Than Thong Reservoir, on decaying wood of Antidesmamadagascariense (Phyllanthaceae), 05 July 2024, Achala Rathnayaka, AA24 (MFLU 24-0525); living culture MFLUCC 24-0607.
Known distribution and hosts.
China (rotten wood) (Ke et al. 2024); Thailand (on corticated wood, Antidesmamadagascariense) (Crous et al. 2020; this study).
Notes.
According to the multi-gene phylogenetic analyses (ITS, LSU, β-tub, and rpb2), our strain (MFLUCC 24-0607) clustered with the ex-type strain of A.spougei (SWUF09-032) with 100% ML bootstrap and 1.00 PP support (Fig. 2). Our fungal collection (MFLUCC 24-0607) exhibits morphological characteristics similar to the holotype of A.spougei (SWUFH099), including black, shiny carbonaceous ascostromata; 8-spored, unitunicate, cylindrical asci, with an apical ring that bluing in Melzer’s iodine reagent; and unicellular, inequilaterally ellipsoidal, brown ascospores (Crous et al. 2020). However, the ascospores of the A.spougei holotype show a straight germ slit along the full length of the spore, which is not observed in our isolate (MFLUCC 24-0607). Based on the morpho-molecular evidence, we identified our collection as a new host record of A.spougei on Antidesmamadagascariense in Thailand.
. Annulohypoxylon purpureonitens
(Y.M. Ju & J.D. Rogers) Y.M. Ju, J.D. Rogers & H.M. Hsieh, Mycologia 97(4): 861 (2005)
2147B750-CED2-549B-A177-6B263B9FD47B
Index Fungorum: IF500323
Facesoffungi Number: FoF17296
Figure 8.
Annulohypoxylonpurpureonitens on the dead wood of Sterculiatragacantha (MFLU 24-0527, a new host record). a. Substrate; b, c. Appearance of ascostromata on host; d. Ostiolar discs in ascomata (indicated by arrows); e. Horizontal section through ascomata; f. Paraphyses; g–l. Asci; m. Apical apparatus (not staining in Melzer’s reagent); n–q. Ascospores (q: arrows indicate the germ slit); r Germinated ascospores; s, t Colony on the PDA (s upper, t lower). Scale bars: 5 mm (b); 500 μm (c, d); 200 μm (e); 20 μm (f–l); 5 μm (m–r).
Hypoxylon purpureonitens Y.M. Ju & J.D. Rogers 1996. Basionym.
Description.
Saprobic on the dead wood of Sterculiatragacantha. Sexual morph: Ascostromata 2–10 mm long × 3–8 mm broad and 0.25–0.35 mm thick (x̄ = 7.5 × 5.5 × 0.3 mm, n = 10), hemispherical, effused-pulvinate, solitary or clustered, shiny, surface black, carbonaceous. Ascomata 0.4–0.5 mm high × 0.3–0.4 mm diam. (x̄ = 0.45 × 0.35 mm, n = 15), immersed in the stroma, subglobose to globose, black, ostioles papillate, encircled with a flattened truncatum-type disc 0.18–0.25 mm diam. (x̄ = 0.2 mm, n = 10). Hamathecium 2.5–4.5 μm wide, comprising long, hyaline, unbranched, aseptate paraphyses. Asci 90–110 × 5–7 μm (x̄ = 100 × 6.2 μm, n = 20), the spore-bearing parts 60–70 µm long with stipes 20–40 µm long, 6-spored, unitunicate, cylindrical, with an apical ring not bluing in Melzer’s iodine reagent. Ascospores 7–9 × 3–6 μm (x̄ = 8.5 × 4.4 μm, n = 40), uniseriate, one-celled, inequilaterally ellipsoidal, with narrowly rounded ends, hyaline when immature, becoming dark brown at maturity, guttules present at the immature stage, with straight, spore length germ slit. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. Colonies on the PDA reaching 2–2.5 cm diam. after seven days at 25 °C, circular in shape, white at first, cottony, white color in the front view, dark brown or black in the middle, and brown at the margin in the reverse view.
Material examined.
Thailand • Chiang Rai, Phan District, Sai Khao, forest area near Wat Udom Waree, on decaying wood of Sterculiatragacantha (Malvaceae), 05 July 2024, Achala Rathnayaka, AA30 (MFLU 24-0527); living culture MFLUCC 24-0609.
Known distribution.
Brazil (on an unidentified branch of a dicotyledonous tree) (Pereira et al. 2009); Thailand (on rotten wood, Sterculiatragacantha) (Suwannasai et al. 2013; this study).
Notes.
According to the multi-gene phylogenetic analyses (ITS, LSU, β-tub, and rpb2), our strain (MFLUCC 24-0609) clustered with the ex-type strain of A.purpureonitens (MFLUCC 14-1225) with 97% ML bootstrap and 0.84 PP support (Fig. 3). Our fungal collection (MFLUCC 24-0609) shows morphological characteristics similar to the holotype of A.purpureonitens (WSP 71615), including effused-pulvinate ascostromata and unicellular, ellipsoid-inequilateral, brown ascospores with a straight, spore-length germ slit (Pereira et al. 2009). In this study, we introduced our fungal collection as a new host record of A.purpureonitens on Sterculiatragacantha in Thailand.
. Annulohypoxylon violaceopigmentum
Sir & Kuhnert, Fungal Diversity: 10.1007/s13225-016-0377-6, [9] (2016)
D28D608A-AFA8-5EC4-834A-57EDF81E1D20
Index Fungorum: IF552341
Facesoffungi Number: FoF02507
Figure 9.
Annulohypoxylonviolaceopigmentum on the dead wood of Syzygiumpolyanthum (MFLU 24-0528, a new host record). a. Substrate; b, c. Appearance of ascomata on the host (ostiolar discs indicated by the arrow); d. Horizontal section through ascomata; e. Paraphyses; f–k. Asci; l. Ascus apical apparatus (not stained in Melzer’s reagent); m–q. Ascospores (q: arrow indicates the germ slit); r A germinated ascospore; s, t Colony on the PDA (s upper, t lower). Scale bars: 2 mm (b); 500 μm (c); 200 μm (d); 20 μm (e–k, r); 5 μm (l–q).
Description.
Saprobic on the dead wood of Syzygiumpolyanthum. Sexual morph: Ascostromata 8–15 mm long × 5–10 mm broad and 0.5–1.2 mm thick (x̄ = 12 × 5.5 × 0.8 mm, n = 10), hemispherical, effused-pulvinate, clustered, developing within cuticle, surface black, carbonaceous. Ascomata 0.4–0.5 mm high × 0.3–0.4 mm diam. (x̄ = 0.45 × 0.35 mm, n = 10), immersed in the stroma, subglobose to globose, black, ostioles papillate, encircled with a flattened truncatum-type disc 0.2–0.25 mm diam. (x̄ = 0.24 mm, n = 5). Hamathecium 1–2 μm wide, comprising long, hyaline, unbranched, aseptate paraphyses. Asci 74–110 × 5–7 μm (x̄ = 97 × 6.7 μm, n = 20), the spore-bearing parts 70–77 µm long with stipes 29–32 µm long, 8-spored, unitunicate, cylindrical, with an apical ring not bluing in Melzer’s iodine reagent. Ascospores 7–10 × 4–6 μm (x̄ = 8.7 × 4.6 μm, n = 40), uniseriate, one-celled, inequilaterally ellipsoidal with narrowly rounded ends, hyaline when immature, becoming dark brown at maturity, guttules at immature stage, with a spore length straight germ slit. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. Colonies on the PDA reaching 2–2.5 cm diam. after five days at 25 °C, circular in shape, white at first, cottony, white color in the front view, brown in the middle, and pale brown at the margin in the reverse view.
Material examined.
Thailand • Chiang Rai, Phan District, Sai Khao, forest area near Wat Udom Waree, on decaying wood of Syzygiumpolyanthum (Myrtaceae), 05 July 2024, Achala Rathnayaka, AA31 (MFLU 24-0528); living culture MFLUCC 24-0610.
Known distribution and hosts.
Thailand (on dead wood, Syzygiumpolyanthum) (Kuhnert et al. 2017; this study).
Notes.
The morphological description of our collection (MFLUCC 24-0610) aligns with the holotype of A.violaceopigmentum (MFLU 14-0314), including effused-pulvinate ascostromata; black, ostioles, papillate ascomata encircled by a flattened truncatum-type disc; 8-spored, cylindrical asci; and brown, unicellular, inequilaterally ellipsoidal ascospores with broadly rounded ends and a straight germ slit along the spore length (Kuhnert et al. 2017). Based on the multi-gene phylogenetic analyses (ITS, LSU, β-tub, and rpb2), our strain (MFLUCC 24-0610) clusters with the ex-type strain of A.violaceopigmentum (MFLUCC 14-1225) with 97% ML bootstrap and 0.85 PP support (Fig. 3). Considering the morpho-molecular evidence, we conclude that our collection is a new host record of A.violaceopigmentum on Syzygiumpolyanthum in Thailand.
. Halorosellinia
Whalley, E.B.G. Jones, K.D. Hyde & Læssøe, Mycol. Res. 104(3): 368 (2000)
63FA1191-BB72-5099-A27F-5311F6CD8849
Index Fungorum: IF28368
Facesoffungi Number: FoF03045
Notes.
Halorosellinia was introduced by Whalley et al. (1999) as a monotypic genus to accommodate H.oceanica (previously referred to as Hypoxylonoceanicum). This genus is characterized by uniperitheciate ascomata immersed in a pseudostroma (Hyde et al. 2016). Halorosellinia currently comprises five species (Index Fungorum 2025). Only three Halorosellinia species are included in Wijayawardene et al. (2022), while five species are listed in the Index Fungorum (2025) and Hyde et al. (2024).
Phylogenetic analyses for Xylariaceae
For Xylariaceae, the ITS, rpb2, and β-tub gene regions were used in the combined data set. Seventy-two isolates of Xylariaceae species were included in the analysis, with Hypoxylonfragiforme (HAST 383 and MUCL 51264) as the outgroup taxa. After alignment, the dataset comprises 2747 characters, including gaps (ITS = 580 bp, rpb2 = 1122 bp, β-tub = 1045 bp). The topology of the BI tree was similar to that of the ML tree. The best-scoring RAxML tree, with a final likelihood value of -49702.0557, is shown in Fig. 10. The matrix comprises 1637 distinct alignment patterns, with 16.93% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.243971, C = 0.271335, G = 0.241282, and T = 0.243412; substitution rates were AC = 1.350468, AG = 5.371719, AT = 1.169242, CG = 1.195494, CT = 7.102768, and GT = 1.0; and the gamma distribution shape parameter α = 0.335659. In the BI analysis, the average standard deviation of split frequencies was 0.01 after 3,000,000 generations of runs. The phylogenetic tree topology is similar to the study by Konta et al. (2020a). According to the phylogenetic analyses, our strains (MFLU 24-0536 and MFLUCC 25-0025) cluster with Haloroselliniaxylocarpi (MFLU 18-0545) with 100% ML bootstrap and 1.00 PP support, while MFLUCC 24-0611 clusters sister to Stilbohypoxylonquisquiliarum (YMJ 172) with 90% ML bootstrap and 0.98 PP support (Fig. 10).
Figure 10.
Phylogram generated from ML analysis based on the combined dataset of ITS, rpb2, and β-tub. The tree is rooted to Hypoxylonfragiforme (HAST 383 and MUCL 51264). Bootstrap support values for ML ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.90 are noted at the nodes. Strain numbers are noted after the species names. Strains isolated in this study are represented in blue, and type strains are in bold.
Taxonomy
. Halorosellinia xylocarpi
Dayar & K.D. Hyde, Mycosphere 11(1): 158 (2020)
D67773CC-30AE-5878-9C0F-DB4DD040DBEC
Index Fungorum number: IF556600
Facesoffungi Number: FoF06192
Figure 11.
Haloroselliniaxylocarpi on decaying submerged wood of Arecaceae sp. (MFLU 24-0536, a new host record). a. Substrate; b. Appearance of an ascoma on the host; c. A horizontal section through an ascoma; d. Peridium; e. Paraphyses; f–k. Asci; l, m. Apical apparatus stained blue with Melzer’s reagent; n–s. Ascospores (r: arrow shows the germ slit on the ventral side); t A germinated ascospore; u, v Colony on the PDA (u upper, v lower). Scale bars: 100 μm (b, c); 20 μm (d–k); 10 μm (l–t).
Description.
Saprobic on decaying submerged wood of Arecaceae sp. Sexual morph: Pseudostromata 0.6–1.0 × 0.5–0.8 mm (x̄ = 0.8 × 0.6 mm, n = 5), superficial, pulvinate to hemispherical, in clusters of uni-peritheciate pseudostromata, surface black, carbonaceous, lacking ascomatal projections. Ascomata 0.3–0.34 × 0.34–0.36 mm (x̄ = 0.33 × 0.35 mm, n = 5), superficial, globose or subglobose to hemispherical, black, ostioles papillate. Peridium 25–38 μm wide, consists of 6–7 layers of brown to dark brown textura angularis cells. Paraphyses 4–8 μm wide, hyaline, abundant, persistent, unbranched, septate. Asci 85–115 × 9–17 μm (x̄ = 97 × 12 μm, n = 20), 8-spored, unitunicate, cylindrical, long pedicellate, with J+, cylindrical apical ring. Ascospores 13–17 × 7–9 μm (x̄ = 15 × 8 μm, n = 30), overlapping 1–2-seriate, hyaline, becoming opaque green and dark brown when mature, more or less equilaterally ellipsoid, straight, both ends often pointed, 1-celled, guttulate, without appendages, a spore length germ slit on the ventral side, straight. Asexual morph: Undetermined.
Culture characteristics.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on the PDA at 25–28 °C, reaching 6 cm in seven days, circular in shape, zonate with diffused margins, white color in front view, and pale yellow in reverse view.
Material examined.
Thailand • Chang Wat Prachuap Khiri Khan Province, Pran Buri District, Pran Buri riverbank, 26 February 2023, Tharindu Bhagya, on decaying submerged wood of Arecaceae sp., TB (MFLU 24-0536), living culture, MFLUCC 25-0025.
Known distribution and hosts.
Thailand (submerged wood of Xylocarpus sp., Rhizophora sp., and submerged wood of Arecaceae sp.) (Dayarathne et al. 2020b; this study)
Notes.
Morphologically, our collection (MFLU 24-0536/MFLUCC 25-0025) resembles the holotype of H.xylocarpi (MFLU 18-0545) in having superficial, carbonaceous, uni-perithecial pseudostromata; 8-spored, cylindrical, unitunicate, long-pedicellate asci with a J+, cylindrical apical ring; and dark brown, unicellular, ellipsoid, straight ascospores with both ends often pointed and a straight germ slit on the ventral side along the spore length (Dayarathne et al. 2020b). However, asci (85–115 × 9–17 μm vs. 126–135 × 20–28 μm) and ascospores (13–17 × 7–9 μm vs. 20–26 × 10–14 μm) of our collection (MFLUCC 25-0025) are smaller than the holotype (Dayarathne et al. 2020b). According to multi-gene phylogeny (ITS, rpb2, and β-tub), our strains (MFLU 24-0536 and MFLUCC 25-0025) cluster with the ex-type strain of H.xylocarpi (MFLU 18-0545) with 100% ML bootstrap and 1.00 PP support (Fig. 10). Considering the morpho-molecular evidence, we conclude that our collection is a new host record on decaying submerged wood of Arecaceae sp. in Thailand.
. Stilbohypoxylon
Henn., Hedwigia 41: 16 (1902)
4941AF0D-E617-5406-95A8-4A4CCC6B16E9
Index Fungorum: IF5264
Facesoffungi Number: FoF03071
Notes.
Stilbohypoxylon was established by Hennings (1902) to accommodate S.moelleri as the type species. Morphologically, this genus is characterized by black, globose to pulvinate stromata, cylindrical asci with a J+, apical ring; and brown, ellipsoidal ascospores surrounded by a thin mucilaginous sheath and a straight or spiral germ slit. Stilbohypoxylon species have geniculosporium-like asexual morphs (Hennings 1902; Rogers and Ju 1997; Petrini 2004; Daranagama et al. 2018). Based on morphology and phylogenetic studies by Daranagama et al. (2018) and Wendt et al. (2018), Stilbohypoxylon was accepted in Xylariaceae. Stilbohypoxylon species cluster with Xylaria species in two subclades, providing evidence that this genus is polyphyletic (Hsieh et al. 2010; Li et al. 2017; Daranagama et al. 2018; Wendt et al. 2018). Hyde et al. (2024) listed 20 species under this genus, while 18 are included in the Index Fungorum (2025).
. Stilbohypoxylon chiangraiense
Rathnayaka, K.D. Hyde & Chethana sp. nov.
15643B0D-C30C-577A-B93F-47A23D7CBEBF
Index Fungorum number: IF903884
Facesoffungi Number: FoF17289
Figure 12.
Stilbohypoxylonchiangraiense on the dead branch of Saraca sp. (MFLU 24-0529, Holotype). a. Substrate; b, c. Appearance of stromata on the host, showing yellow scales (arrows indicating yellow scales); d, e. A horizontal section through a stroma; f. Peridium; g. Paraphyses; h, i. Immature asci; j. Apical apparatus stained in blue with Melzer’s reagent; k–m. Ascospores; n, o. Ascospores with a germ slit (arrows indicating a spiral germ slit); p, q. Colony on the PDA (p upper, q lower). Scale bars: 1 mm (b); 500 μm (c); 200 μm (d, e); 20 μm (g–j); 10 μm (f, k–o).
Etymology.
The epithet chiangraiense refers to Chiang Rai Province, where the fungus was collected.
Holotype.
MFLU 24-0529.
Description.
Saprobic on a dead branch of Saraca sp. Sexual morph: Stromata superficial, visible as a black conical or globose structure on the host surface, solitary, showing yellow scales on mature stromata, carbonaceous, brittle, fragile. Ascomata 420–440 × 450–510 μm (x̄ = 427 × 471 μm, n = 10), black, carbonaceous, globose to mammiform, 1 per stroma, covered with remnants of the host tissue, ostioles papillate. Peridium 10–20 μm wide, thick-walled, composed of several layers of cells of textura angularis, dark brown to black. Paraphyses 2–3 μm wide, filamentous, cylindrical, aseptate, unbranched, longer than asci. Asci 48–58 × 7–9 μm (x̄ = 53 × 8 μm, n = 10), unitunicate, cylindrical, long pedicellate, apically rounded, with a J+, apical ring (rarely seen). Ascospores 21–27 × 10–14 μm (x̄ = 24 × 12 μm, n = 30), uniseriate, hyaline when immature, dark brown at maturity, equilateral ellipsoidal to broadly fusoid, unicellular, guttulate, with a spiral germ slit over the whole spore length. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. The slow-growing colonies on the PDA reached 1–1.5 cm diam. After five days at 25 °C, circular in shape, cottony, slightly less dense towards the edge, white color in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Nang Lae village, on decaying branch of Saraca sp. (Fabaceae), 18 March 2024, Achala Rathnayaka, AA13 (MFLU 24-0529, holotype); ex-type living culture, MFLUCC 24-0611.
Notes.
Based on the multi-gene phylogeny (ITS, rpb2, and β-tub), our collections (MFLU 24-0529 and MFLUCC 24-0611) formed a distinct lineage sister to S.quisquiliarum (YMJ 172) with 90% ML bootstrap and 0.98 PP support (Fig. 10). Stilbohypoxylonchiangraiense shares morphologies similar to the Stilbohypoxylon genus by having superficial, solitary, globose stromata, cylindrical asci with a J+, apical ring; and brown, ellipsoidal ascospores with a spiral germ slit. However, asci are very rarely observed in S.chiangraiense. Stilbohypoxylonquisquiliarum differs from our fungal collection in that its yellow scales turn brown when mature, which are present on the stromata. In addition, our fungal collection has shorter ascospores (21–27 μm) than S.quisquiliarum (27.5–28.5 μm) (Petrini 2004). The base pair differences between S.chiangraiense (MFLUCC 24-0611) and S.quisquiliarum (YMJ 172) are as follows: ITS = 1.9% (11/578), β-tub = 11.9% (123/1033). Considering the morpho-molecular data analysis, we established S.chiangraiense as a new species in Stilbohypoxylon.
. Hypoxylon
Bull., Hist. Champ. Fr. (Paris) 1: 168 (1791)
07775400-3971-5843-993C-A1103C1CD36C
Index Fungorum: IF2456
Facesoffungi Number: FoF02980
Notes.
Bulliard (1791) introduced Hypoxylon to accommodate H.fragiforme (basionym: H.coccineum) as the type species. The sexual morph of this genus is characterized by ascomata embedded in a colorful, effused, or pulvinate stroma containing secondary metabolites (Ju and Rogers 1996). The ascomata are perithecioid, monostichous, and open separately through umbilicate, rarely slightly papillate ostioles. The asci are 8-spored, unitunicate, cylindrical, stipitate, and provided with a typically amyloid apical apparatus. The ascospores are unicellular, ellipsoid, and brown and have a germ slit on the most convex side of the inequilateral ascospores (Ju and Rogers 1996). The asexual morph is characterized by a nodulisporium-like morph, but other types of conidial states have also been observed, such as sporothrix-like, virgariella-like, and periconiella-like (Ju and Rogers 1996). The evolutionary relationships of hypoxylaceous fungi have been studied using phylogenetic, chemotaxonomic, and morphological data (Kuhnert et al. 2021). Most of the Hypoxylon species have been able to produce highly bioactive secondary metabolites, which are released from the stromata (Ju and Rogers 1996; Kuhnert et al. 2014b; Fournier et al. 2016). Hypoxylon species have a cosmopolitan distribution and are recorded as saprotrophs that grow on dead wood, endophytes in seed plants, and facultative parasites on diseased hosts (Ju and Rogers 1996; Stadler 2011; Kuhnert et al. 2014a; Daranagama et al. 2018; Helaly et al. 2018; Rogers 2018). Hyde et al. (2024) listed 200 species under this genus, while 466 are included in the Index Fungorum (2025).
Phylogenetic analyses for Hypoxylaceae
For Hypoxylon, 150 taxa were included in the combined data set (ITS, LSU, rpb2, and β-tub). Graphostromaplatystomum (CBS 270.87), Natonodosaspeciosa (CLM RV86), Xylariaarbuscula (CBS 126415), and X.hypoxylon (CBS 122620) were used as the outgroup taxa. After alignment, the dataset comprised 3003 characters, including gaps (ITS = 614 bp, LSU = 822 bp, rpb2 = 1017 bp, β-tub = 550 bp). Both the ML and BI analyses exhibit a similar tree topology. The best-scoring RAxML tree was obtained (Fig. 13), with a final likelihood value of -75469.623433. The matrix included 1852 distinct alignment patterns, with 29.09% undetermined characters or gaps. The estimated base frequencies were as follows: A = 0.247189, C = 0.251478, G = 0.262907, and T = 0.238426; substitution rates were AC = 1.284950, AG = 4.479967, AT = 1.344479, CG = 1.074839, CT = 6.471802, and GT = 1.0; and the gamma distribution shape parameter α = 0.313445. In the BI analysis, the average standard deviation of the split frequencies was 0.01 after 5,000,000 generations of runs. The phylogenetic tree topology is similar to the study by Karimi et al. (2023). According to the phylogenetic analyses, our strains (MFLU 24-0530, MFLUCC 25-0024, MFLUCC 24-0613, MFLU 24-0532, and MFLUCC 24-0612) cluster within Hypoxylon and Hypomontagnella.
Figure 13.
Phylogram generated from ML analysis based on the combined dataset of ITS, LSU, rpb2, and β-tub. The tree is rooted to Graphostromaplatystomum (CBS 270.87), Natonodosaspeciosa (CLM RV86), Xylariaarbuscula (CBS 126415), and X.hypoxylon (CBS 122620). Bootstrap support values for ML ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.90 are noted at the nodes. Strain numbers are noted after the species names. Strains isolated in this study are represented in blue, and type strains are in bold.
Taxonomy
. Hypoxylon thailandicum
Rathnayaka, K.D. Hyde & Chethana sp. nov.
8CECEFFF-0DB3-539F-ADEE-0E56590A994E
Index Fungorum number: IF903885
Facesoffungi Number: FoF00373
Figure 14.
Hypoxylonthailandicum on a dead branch of Bambusavulgaris (MFLU 24-0530, Holotype). a. Substrate; b–d. Appearance of ascostromata on the host; e, f. A horizontal section through an ascoma; g. Peridium; h–j. Asci; k. Inamyloid apical ascal apparatus stained with Melzer’s reagent; l–o. Ascospores; p. A germinated ascospore; q, r. Colony on the PDA (q upper, r lower). Scale bars: 1 mm (b); 200 μm (c, d); 100 μm (e, f); 20 μm (h–k); 10 μm (g, l–p).
Etymology.
The epithet thailandicum refers to Thailand, where the fungus was collected.
Holotype.
MFLU 24-0530.
Description.
Saprobic on a dead branch of Bambusavulgaris. Sexual morph: Stromata 0.3–1 cm long × 0.1–0.5 cm wide, pulvinate, with conspicuous perithecial mounds, gregarious, surface bright orange; orange-red granules immediately beneath the surface and between ascomata, the tissue below the perithecial layer inconspicuous. Ascomata 200–220 × 195–203 × 180–210 µm (x̄ = 210 × 200 × 190 µm, n = 5), globose, ostiolate. Peridium 21–27 μm wide, two-layered, outer layer composed of dark brown to brown cells of textura angularis, inner layer composed of hyaline cells of textura angularis. Asci 60–105 × 8–11 µm (x̄ = 80 × 9.4 µm, n = 15), 8-spored, unitunicate, cylindrical, pedicellate, with an inamyloid, apical ascal apparatus. Ascospores 13–18 × 7–10 µm (x̄ = 16 × 8.4 µm, n = 30), uniseriate, slightly overlapping, one-celled, ellipsoid, with narrowly rounded ends, straight, initially light brown, becoming brown to dark brown at maturity, rough surface, guttulate. Asexual morph: Undetermined.
Culture characters.
Ascospores germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. The slow-growing colonies on the PDA reach 1–2 cm diam. after seven days at 25 °C, circular in shape, cottony, slightly less dense towards the edge, white color in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Mae Chan District, Mae Chan village, on a dead branch of Bambusavulgaris (Poaceae), 18 March 2024, Achala Rathnayaka, AA06 (MFLU 24-0530, holotype); ex-type living culture, MFLUCC 25-0024.
Notes.
In multi-gene phylogeny (ITS, LSU, rpb2, and β-tub), our novel isolates (MFLU 24-0530 and MFLUCC 25-0024) formed a separate lineage sister to H.begae (S99 and YMJ 215) and H.blackburniae (BRIP 72467b) with 97% ML bootstrap and 0.95 PP support (Fig. 13). Morphologically, our new fungal collection (MFLUCC 25-0024) is similar to Hypoxylon by having ascomata embedded in colorful effused or pulvinate stromata (Ju and Rogers 1996). Due to the lack of morphological data for H.begae and H.blackburniae, we could not compare the morphological characters between these three species. When comparing the ITS base pair differences (without gaps) between H.thailandicum (MFLUCC 25-0024) with H.begae (YMJ 215) and H.blackburniae (BRIP 72467b), there are 9.5% (46/482) and 9.12% (44/482) differences, respectively. For β-tub, there is a 10.21% (43/423) base pair difference (without gaps) between H.thailandicum (MFLUCC 25-0024) and H.begae (YMJ 215). However, due to the lack of sequence availability, we were unable to compare the base pair differences between H.thailandicum (MFLUCC 25-0024) and H.blackburniae (BRIP 72467b). Additionally, the absence of rpb2 sequences prevented a comparison among H.begae, H.blackburniae, and our collection. Based on the available morphological and phylogenetic evidence, we propose H.thailandicum as a new species.
. Hypomontagnella
Sir, L. Wendt & C. Lamb., in Lambert, Wendt, Hladki, Stadler & Sir, Mycol. Progr. 18(1–2): 190 (2019)
7C5C3D09-44C5-57EC-874C-F912589A8731
Index Fungorum: IF827251
Facesoffungi Number: FoF06136
Notes.
Lambert et al. (2019) introduced Hypomontagnella to accommodate H.monticulosa as the type species and included several species previously described under Hypoxylon. Hypomontagnella differs from Annulohypoxylon and Jackrogersella by smooth perispores or transversally striate ornamentations. Additionally, Hypomontagnella species are distinguished from Hypoxylon species by woody to carbonaceous stromata that lack colored granules (Lambert et al. 2019). They have papillate ostioles, usually with black annulate discs, without apparent KOH-extractable pigments in mature stromata (Lambert et al. 2019). The cultures of Hypomontagnella species produce sporothrolide-type strong antifungal polyketides. Species in this genus have been reported as saprobic or endophytic on plants (Lambert et al. 2019). Six species are listed under Hypomontagnella in Hyde et al. (2024) and Index Fungorum (2025).
. Hypomontagnella hibisci
Rathnayaka, K.D. Hyde & Chethana sp. nov.
B0C3A28D-4C43-5037-9064-D8FB59AF9633
Index Fungorum number: IF903886
Facesoffungi Number: FoF17290
Figure 15.
Hypomontagnellahibisci on a decaying branch of Hibiscus sp. (MFLU 24-0532, Holotype). a. Substrate; b. Mature stroma on the bark; c. Stromatal surface showing papillate and ostiolar discs (indicated by white arrows); d–f. Stromata in vertical sections; g. Peridium; h. Paraphyses; i–m. Asci; n. Apical apparatus with Melzer’s reagent; o–r. Ascospores; s, t. Colony on the PDA (s upper, t lower). Scale bars: 2 mm (b); 1 mm (c); 200 μm (d); 100 μm (e); 50 μm (f); 10 μm (g, n); 20 μm (h–m); 5 μm (o–r).
Etymology.
In reference to the host genus from which the fungus was collected
Holotype.
MFLU 24-0532
Description.
Saprobic on a dead branch of Hibiscus sp. Sexual morph: Stromata effused-pulvinate, with conspicuous to inconspicuous perithecial mounds, surface blackish, carbonaceous tissue immediately beneath the surface and between the perithecial surface and perithecia. Ascomata 157–300 × 133–218 μm (x̄ = 223 × 178 μm, n = 5), globose to spherical, ostioles higher than the stromatal surface. Peridium 13–26 μm diam., composed of thin-walled, brown to dark brown cells of textura angularis. Paraphyses 2–3 μm wide, hyaline, filamentous, long, branched, aseptate, arising from the base of ascomata. Asci 78–100 × 5–6 μm (x̄ = 90 × 5.4 μm, n = 25), the spore-bearing parts 46–53 µm long, 8-spored, cylindrical, with J-, apical ring and a stipe of 28–36 µm long. Ascospores 7–9 × 3–5 μm (x̄ = 8 × 4 μm, n = 30), unicellular, uni-seriate, slightly overlapping, ellipsoid, with narrowly rounded ends, slightly curved, hyaline to dark brown, smooth to finely roughened, guttulate. Asexual morph: Undetermined.
Culture characteristics.
Ascospores are germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on the PDA reach 1.5–2.5 cm diam. after seven days at 25 °C, circular in shape, flat, cottony, slightly less dense towards the edge, white color in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Mae Fah Luang University premises, on a decaying branch of Hibiscus sp. (Malvaceae), 08 March 2024, Zaw Lin Tun, AZ01 (MFLU 24-0532, holotype); ex-type culture MFLUCC 24-0613.
Notes.
Hypomontagnellahibisci (MFLUCC 24-0613) is similar to Hypomontagnella in having stromata with conical, papillate ostioles and cylindrical asci with a short pedicel (Lambert et al. 2019). According to the multi-gene phylogenetic analyses, our strains (MFLU 24-0532 and MFLUCC 24-0613) formed a separate lineage sister to H.monticulosa (MUCL 54604 and MFLUCC 24-0613) with 100% ML bootstrap and 1.00 PP support (Fig. 13). However, H.hibisci has globose to spherical perithecia and asci with J-, apical rings, whereas H.monticulosa has spherical to obovoid perithecia and asci with J+ discoid apical rings (Chethana et al. 2021a). Additionally, H.monticulosa differs from H.hibisci by having ascospores with a straight, spore-length germ slit, which is not observed in H.hibisci (Chethana et al. 2021a). With the evidence of unique morphology and distinct phylogeny, we introduce H.hibisci as a new species.
. Hypomontagnella monticulosa
(Mont.) Sir, L. Wendt & C. Lamb., in Lambert, Wendt, Hladki, Stadler & Sir, Mycol. Progr. 18(1–2): 190 (2019)
6CFB1552-E384-563C-A0BF-D92B6618326B
Index Fungorum: IF827252
Facesoffungi Number: FoF06781
Figure 16.
Hypomontagnellamonticulosa on a dead branch of Macarangapeltata (MFLU 24-0531, a new host record). a. Substrate; b, c. Appearance of mature stroma on the host; d. A horizontal section through ascomata; e. Paraphyses; f–i. Asci; j. Apical apparatus stained blue with Melzer’s reagent; k–n. Ascospores; o. A germinated ascospore; p, q. Colony on the PDA (p upper, q lower). Scale bars: 2 mm (b); 500 μm (c); 200 μm (d); 20 μm (e–i); 10 μm (j, o); 5 μm (k–n).
Description.
Saprobic on a dead branch of Macarangapeltata. Sexual morph: Stromata effused-pulvinate, with conspicuous to inconspicuous perithecial mounds, surface blackish, woody to carbonaceous tissue immediately beneath the surface and between the perithecial surface and the perithecia. Perithecia globose to subglobose, ostioles higher than the stromatal surface. Paraphyses 4–7 μm wide, hyaline, abundant, persistent, unbranched, septate. Asci 70–95 × 4–6.5 μm (x̄ = 82 × 5.4 μm, n = 20), the spore-bearing parts 40–50 µm long with stipes 36–44 µm long, 8-spored, unitunicate, cylindrical, with J+, discoid apical ring. Ascospores 6–8 × 3–4 μm (x̄ = 7.2 × 3.4 μm, n = 30), uniseriate, unicellular, ellipsoid-inequilateral, with broadly to less frequently narrowly rounded ends, light brown to brown, smooth. Asexual morph: Undetermined.
Culture characteristics.
Ascospores are germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. The slow-growing colonies on the PDA reach 1.0–1.5 cm diam. after seven days at 25 °C, circular in shape, flat, cottony, slightly less dense towards the edge, white color in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Nang Lae village, on decaying branch of Macarangapeltata (Euphorbiaceae), 18 March 2024, Achala Rathnayaka, AA10 (MFLU 24-0531); living culture, MFLUCC 24-0612.
Known distribution and hosts.
Argentina (Ficusmaroma) (Lambert et al. 2019), French Polynesia (dead wood) (Lambert et al. 2019), Indonesia, Malaysia (lichen, Sargassum seaweed) (Zainee et al. 2018), Paraguay (dead wood) (Lambert et al. 2019), Thailand (Leucaenaleucocephala, Macarangapeltata) (Chethana et al. 2021a, this study), USA (Cladonialeporina) (U’Ren et al. 2016).
Notes.
Morphologically, our collection (MFLUCC 24-0612) is similar to the ex-type strain of H.monticulosa (MUCL 54604), which was collected from a dead branch of Leucaenaleucocephala in Thailand (Chethana et al. 2021a). However, asci (70–95 μm vs. 85–110 μm) and ascospores (6–8 μm vs. 7.5–9.3 μm) of our collection (MFLUCC 24-0612) are shorter than the ex-type strain (MUCL 54604) (Chethana et al. 2021a). According to multi-gene phylogeny (ITS, LSU, rpb2, and β-tub), our strain (MFLUCC 24-0612) clusters with the ex-type strain of H.monticulosa (MUCL 54604) with 100% ML bootstrap and 1.00 PP support (Fig. 13). Based on the morpho-molecular evidence, we established the first host record of H.monticulosa on Macarangapeltata in Thailand.
. Diatrypella
(Ces. & De Not.) De Not., Sfer. Ital.: 29 (1863)
A090A059-70E2-5033-8C01-436EEC16FB85
Index Fungorum: IF1505
Facesoffungi Number: FoF11777
Notes.
Cesati and De Notaris (1863) introduced Diatrypella with D.verruciformis as the type species. This genus is characterized by stromata, which are conical to truncate, cushion-like or discoid, and usually delimited by a black zone within host tissues, umbilicate or sulcate ostiolar necks. Asci are cylindrical, polysporous, and long-stalked, and ascospores are hyaline to yellowish. Diatrypella species have a libertella-like coelomycete asexual morph (Kirk et al. 2008; Hyde et al. 2020a). There are 74 Diatrypella species in Species Fungorum (2024), and only 23 of them have sequence data.
Phylogenetic analysis for Diatrypaceae
The ITS and β-tub combined data set consists of 144 taxa representing strains of Diatrypaceae, including Kretzschmariadeusta (CBS 826.72) and Xylariahypoxylon (CBS 122620) as the outgroup taxa. The aligned data set comprises 1404 characters, including gaps (ITS = 504 bp, β-tub = 897 bp). The topology of the BI tree was similar to that of the ML tree. The best-scoring RAxML tree, with a final likelihood value of -20102.616796, is shown in Fig. 17. The matrix comprises 944 distinct alignment patterns, with 36.45% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.222804, C = 0.272806, G = 0.233404, T = 0.270985; substitution rates AC = 1.101521, AG = 3.499467, AT = 1.354291, CG = 0.845871, CT = 4.738241, GT = 1.0; gamma distribution shape parameter α = 0.360572. In the BI analysis, the average standard deviation of split frequencies was 0.009 after 9,000,000 generations of runs. The phylogenetic tree topology is similar to the study by Dissanayake et al. (2024). According to the phylogenetic analyses, our strains, MFLU 24-0533 and MFLU 24-0534, formed a separate clade with Diatrypellaoregonensis (CA117 and DPL200), D.pseudooregonensis (GMB0041 and GMB0040), and D.verruciformis (UCROK1467 and UCROK754) with 93% ML bootstrap and 0.94 pp, while MFLUCC 24-0614 clusters with Paraeutrypellacitricola (HVGRF01 and HKAS 133111) with 99% ML and 0.97 pp bootstrap support.
Figure 17.
Phylogram generated from ML analysis based on the combined dataset of ITS and β-tub. The tree is rooted to Kretzschmariadeusta (CBS 826.72) and Xylariahypoxylon (CBS 122620). Bootstrap support values for ML ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.90 are noted at the nodes. Strain numbers are noted after the species names. Strains isolated in this study are presented in blue, and type strains are in bold.
Taxonomy
. Diatrypella thailandica
Rathnayaka, K.D. Hyde & Chethana sp. nov.
91BF7F2A-2103-54FF-983D-E06CE865E42E
Index Fungorum number: IF903887
Facesoffungi Number: FoF17291
Figure 18.
Diatrypellathailandica on a dead branch of Fabaceae sp. (MFLU 24-0533, holotype) a. Substrate; b, c. Stromata on the substrate; d. Cross-section of a stroma; e. Vertical section through stroma showing ostiole and perithecia; f. Ostiole; g. Peridium; h. Paraphyses; i–l. Asci; m. Apical apparatus in Melzer’s reagent; n–q. Ascospores. Scale bars: 2 mm (b); 500 μm (c); 200 μm (d); 100 μm (e, f); 10 μm (g); 20 μm (h–m); 5 μm (n–q).
Etymology.
The epithet thailandica refers to Thailand, from where the fungus was collected.
Holotype.
MFLU 24-0533.
Description.
Saprobic on a dead branch of Fabaceae sp. Sexual morph: Stromata 0.5–1 mm in diam., well-developed, with groups of 10–15 perithecia, solitary to gregarious, erumpent, black, immersed, globose to subglobose or conical shape. Endostroma white to light yellow. Ascomata 410–450 μm high × 275–370 μm diam. (x̄ = 434 × 326 μm, n = 10), perithecial, immersed in stromata, 2–4 perithecial arrangement, subglobose, with an individual ostiole. Ostiolar canal 200–253 μm high, 110–132 μm diam., cylindrical, periphysate, with yellowish pigment around ostioles. Peridium 10–25 μm wide, composed of 3–7 layers, hyaline to brown, thick-walled cells of textura angularis. Hamathecium 2.4–6 μm wide, comprising dense, hyaline, aseptate, unbranched paraphyses, tapering towards the apex, embedded in a hyaline gelatinous matrix. Asci 80–150 × 11–23 μm (x̄ = 107 × 16 μm, n = 25), polysporous, unitunicate, clavate to cylindric-clavate, with a J-apical ring and a long pedicel. Ascospores 6–8 × 1–3 μm (x̄ = 7.5 × 2.3 μm, n = 30), multi-seriate, crowded, initially hyaline, becoming pale yellowish at maturity, oblong to allantoid, aseptate, slightly curved, smooth-walled, mostly with small guttules. Asexual morph: Undetermined.
Culture characteristics.
Ascospores are germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from both sides of the ascospore. Colonies on the PDA at 25–28 °C reach 2 cm in 10 days, medium dense, circular to slightly irregular, cottony, white at first, becoming light brownish yellow in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Nang Lae village, on a decaying branch of Morus sp. (Moraceae), 18 March 2024, Achala Rathnayaka, AA14 (MFLU 24-0533, holotype); ibid., on a dead branch of Fabaceae sp. (Fabaceae), 20 March 2024, Achala Rathnayaka, AA15 (MFLU 24-0534, topotype).
Notes.
Our new fungal collection (MFLU 24-0533 and MFLU 24-0534) fits within Diatrypella by having conical-shaped stromata, white to light yellow, well-developed endostroma, cylindrical, polysporous, and long-stalked asci, and hyaline to yellowish ascospores (Hyde et al. 2020a). According to the multi-gene phylogenetic analyses (ITS and β-tub), our strains (MFLU 24-0533 and MFLU 24-0534) formed a separate clade sister to D.oregonensis (CA117, DPL200), D.pseudooregonensis (GMB0039, GMB000), and D.verruciformis (UCROK1467, UCROK754) with 93% ML bootstrap and 0.94 PP support (Fig. 17). Morphologically, D.thailandica differs from D.oregonensis, D.pseudooregonensis, and D.verruciformis, as mentioned in Table 2. Diatrypellathailandica has polysporous asci, while D.oregonensis and D.pseudooregonensis have 8-spored asci. Diatrypellaverruciformis differs from our fungal collection by having diamond- or star-shaped ascomata (Table 2). When comparing the ITS and β-tub base pairs (without gaps) between D.thailandica (MFLU 24-0533) with D.oregonensis (CA117), D.pseudooregonensis (GMB0039), and D.verruciformis (UCROK1467), there are 2.15% (11/512), 3.3% (16/485), and 2.73% (14/512) base pair differences in the ITS and 2.17% (18/828), 2.53% (21/828), and 1.7% (14/828) for β-tub, respectively. Based on both morphological and molecular evidence, we introduce Diatrypellathailandica (MFLU 24-0533) as a new species in Diatrypella.
Table 2.
Synopsis of morphological characters of sexual morphs between D.thailandica and species in the sister clade.
| Species | Stromata | Perithecial neck (μm) | Asci | Ascospores | References |
|---|---|---|---|---|---|
| D.thailandica | groups of 10–15 perithecia, globose to subglobose or conical shape, 0.5–1 mm in diam | 200–253 high, 110–132 diam. | Polysporous, 80–150 × 11–23 (x̄ = 107 × 16) μm | 6–8 × 1–3 (x̄ = 7.5 × 2.3) μm, L/W = 3.26 | This study |
| D.oregonensis | pustules of 1–30 perithecia pulvinate, hemispherical or forming linear stripes, 0.3–0.6 mm diam | – | 8-spored, 50–65 (–80) × 6–9.5 μm | (7–)10–12 (14) × 2–2.5 | Trouillas et al. (2010) |
| D.pseudooregonensis | groups of 3–16 perithecia, 2 × 1.5 mm | 218.5–465 high, 112–257 diam. | 8-spored, 95–149 × 6.5–11.5 (av. = 120 × 10.5) μm | 11–16 × 1.5–3.5 (x̄ = 14 × 2.5) μm, L/W = 5.6 | Long et al. (2021) |
| D.verruciformis | Diamond or star shape, 5–6 × 2–3 mm | Difficulty to recognize | Multispored, 120–140 (170) × 10–14 (16) μm | 6–7 (8) × 1.5–2 µm | http://www.taunuspilz.de/coppermine/displayimage.php?pid=9376 |
. Paraeutypella
L.S. Dissan., J.C. Kang, Wijayaw. & K.D. Hyde, Biodiversity Data Journal 9: e63864, 11 (2021)
D95C3B42-DBE2-50A3-BE83-694941A3E59B
Index Fungorum: IF557954
Facesoffungi number: FoF09231
Notes.
Paraeutypella was introduced by Dissanayake et al. (2021) to accommodate P.guizhouensis as the type species, together with P.citricola and P.vitis, which were previously classified under Eutypella sensu lato. The genus is characterized by erumpent, clustered, irregularly shaped, dark brown to black, poorly developed stromata, 8-spored asci; and ascospores that are allantoid, overlapping, and subhyaline (Trouillas et al. 2011; de Almeida et al. 2016; Dissanayake et al. 2021). A coelomycetous asexual morph has been recorded in this genus, which was characterized by black, subconic, multi-loculate, largely prosenchymatous conidiomata with yellowish conidial masses. Conidia are hyaline, single-celled, slightly curved, and guttulate (Glawe and Jacobs 1987). There are six species listed in the Index Fungorum (2025).
. Paraeutypella citricola
(Speg.) L.S. Dissan., Wijayaw., J.C. Kang & K.D. Hyde, Biodiversity Data Journal 9: e63864, 14 (2021)
2D28D00C-1D4A-5D13-8A8E-923EDBB3BB27
Index Fungorum: IF557954
Facesoffungi Number: FoF09150
Figure 19.
Paraeutypellacitricola on a dead branch of Swieteniamacrophylla (MFLU 24-0535, a new host record). a. Substrate; b. Stromata on the substrate; c. A cross-section of a stroma; d. A vertical section through the stroma shows ostioles and perithecia; e. Peridium; f. Paraphyses; g–k. Asci; l–o. Ascospores; p, q. Colony on the PDA (p upper, q lower). Scale bars: 5 mm (b); 1 mm (c); 200 μm (d); 20 μm (e–k); 5 μm (l–o).
Eutypella citricola Syd. & P. Syd., Hedwigia 49: 80 (1909), nom. illegit., Art. 53.1. Basionym.
Description.
Saprobic on a dead branch of Swieteniamacrophylla. Sexual morph: Stromata immersed in the bark of dead branches, erumpent, aggregated, circular to irregular in shape, superficial, carbonaceous. Endostroma white to light yellow. Ostiole opening separately, papillate or apapillate, central. Ascomata 840–880 μm high × 430–455 μm diam. (x̄ = 867 × 446 µm, n = 10), perithecial, with groups of 5–10 perithecia arranged in a valsoid configuration, black, subglobose, clustered, immersed in ascostroma with an ostiolar neck. Necks 220–265 μm long (x̄ = 248 µm, n = 10), papillate, central ostiolar canal filled with paraphyses. Peridium 25–48 μm wide, composed of two layers of textura angularis to textura prismatica; inner layer cells hyaline, outer layer cells brown to dark brown. Hamathecium 3–5 μm wide (x̄ = 4 µm, n = 15) comprises hyaline, long, narrow, unbranched, aseptate, guttulate cells, paraphyses arising from the base of perithecia. Asci 56–94 × 5–7 μm (x̄ = 67 × 6.4 μm, n = 20), 8-spored, unitunicate, thin-walled, clavate to cylindrical clavate, long pedicellate (35–55 μm), J- apical ring. Ascospores 7–9 × 2–3 μm (x̄ = 8 × 2.4 μm, n = 40), overlapping 2–3 seriate, allantoid, hyaline to light brown, smooth, aseptate, usually with small guttules. Asexual morph: Undetermined.
Culture characteristics.
Ascospores are germinated on the PDA within 24 hours at 25 °C. Germ tubes are produced from one side of the ascospore. Colonies on the PDA at 25–28 °C reaching 3–5 cm in five days, medium dense, circular to slightly irregular, cottony, white color in the front view, and pale yellow in the reverse view.
Material examined.
Thailand • Chiang Rai, Nang Lae village, on a decaying branch of Swieteniamacrophylla (Meliaceae), 08 April 2024, Achala Rathnayaka, AA16 (MFLU 24-0535); living culture, MFLUCC 24-0614.
Known distribution.
Wide host range and widely distributed in temperate, tropical, and subtropical regions (Senwanna et al. 2021).
Notes.
Based on the phylogenetic analyses, our collection (MFLUCC 24-0614) clustered with other strains of P.citricola (HKAS 13311 and HVGRF01) with 100% ML bootstrap and 1.00 PP support (Fig. 17). Morphologically, our collection is similar to the holotype of P.citricola (HMAS 290660), which was collected from the dead twigs of Acerpalmatum in China (Dissanayake et al. 2021). Both specimens share similar morphological characteristics, including immersed, erumpent, aggregated, superficial, carbonaceous stromata; black, subglobose, clustered ascomata immersed in the ascostroma with an ostiolar neck; 8-spored, unitunicate, clavate to cylindrical-clavate, long pedicellate asci with a J- apical ring; and allantoid, hyaline to light brown, aseptate ascospores, usually with small guttules (Dissanayake et al. 2021). However, our collection has a shorter neck (220–265 µm vs. 360–390 µm) and longer asci (56–94 µm vs. 70–75 µm) than the holotype (Dissanayake et al. 2021). Based on the morpho-molecular evidence, Paraeutypellacitricola has been recorded from Thailand on various woody plants, including Heveabrasiliensis (Senwanna et al. 2021), Magnolia sp. (de Silva et al. 2022), and Microcospaniculata (Afshari et al. 2023). We identified our collection as a new host record of Paraeutypellacitricola from Swieteniamacrophylla from Thailand.
Preliminary screening for antibacterial activity
In the present study, we conducted a preliminary screening to assess the antibacterial activity of selected fungal species against Bacillussubtilis (TISTR 1248) and Escherichiacoli (TISTR 527). Two newly introduced species from Hypoxylaceae and Xylariaceae, Annulohypoxylonchiangraiense (MFLUCC 24-0606) and Hypoxylonthailandicum (MFLUCC 25-0024), showed antibacterial activity against Bacillussubtilis, each producing a 2 mm zone of inhibition, indicating partial inhibition compared to the positive control. Additionally, other Xylariales species, including Hypoxylon sp. (MFLUCC 18-1207), Daldiniaeschscholtzii (MFLUCC 18-1207), and Xylariachrysanthum (MFLUCC 21-0014), demonstrated antibacterial activity against E.coli, each producing a 3 mm zone of inhibition, respectively, compared to the positive control (Fig. 20).
Figure 20.
Preliminary screening of antimicrobial activity using the agar plug diffusion method against Bacillussubtilis (a, b) and E.coli (c, d). a.Annulohypoxylonchiangraiense (MFLUCC 24-0606); b.Hypoxylonthailandicum (MFLUCC 25-0024); c.Daldiniaeschscholtzii (MFLUCC 18-1207), and Xylariachrysanthum (MFLUCC 21-0014). Positive control (ampicillin discs): Left side of the plate in a and b; middle of the petri plate in c.
Metabolite profiling of selected fungal extracts by HPLC coupled to LC-QTOF-MS analyses
The crude extracts of Annulohypoxylonchiangraiense, Hypoxylonthailandicum, Daldiniaeschscholtzii, and Xylariachrysanthum were weighed at 0.01 g, 0.02 g, 0.08 g, and 0.02 g, respectively. An untargeted screening approach initially detected secondary metabolites at wavelengths of 256 and 425 nm. Secondary metabolites have been spotted by the information from spectra and molecular weight, matched with reference compounds from the online databases. Some characteristic HPLC chromatograms are shown in Figs 21, 22.
Figure 21.
HPLC chromatograms of the culture extracts of Annulohypoxylonchiangraiense (MFLUCC 24-0606) a. Mycelium in MEB media; b. Supernatant in MEB media. Hypoxylonthailandicum (MFLUCC 25-0024); c. Mycelium in YMB media; d. Supernatant in YMB media.
Figure 22.
HPLC chromatograms of the culture extracts of Xylariachrysanthum (MFLUCC 21-0014) a. in MEB media; b. Phosphoenol-4-deoxy-3-tetrulosonate; c. in YMB media; d. Benzbromarone; e.Daldiniaeschscholtzii (MFLUCC 18-1207) in MEB media; f. Chlorfenson. Peaks corresponding to metabolites are indicated in the HPLC profiles.
As Fig. 21 depicts, the HPLC analysis revealed distinct differences in the metabolite profiles of Annulohypoxylonchiangraiense and Hypoxylonthailandicum when cultivated in different media. For Annulohypoxylonchiangraiense, the metabolic output was highly dependent on the growth medium. The mycological extract derived from malt extract broth (MEB) exhibited a complex metabolite profile with numerous peaks, indicating the production of a diverse array of secondary metabolites. This suggests that MEB provides favorable conditions for the production of a broad range of secondary metabolites. Similar findings have been reported in previous studies, where MEB was shown to enhance the production of secondary metabolites in various fungi, including Hypoxylon and Xylaria species, due to its rich composition of carbohydrates, amino acids, and vitamins (Kuhnert et al. 2015). In contrast, potato dextrose broth (PDB) resulted in significantly fewer peaks, suggesting a reduced metabolic diversity under these conditions. This could be attributed to the simpler composition of PDB, which may not provide the necessary nutrients or environmental conditions to induce the production of a broad spectrum of secondary metabolites.
Similarly, yeast malt broth (YMB) also showed fewer peaks compared to MEB, suggesting that PDB and YMB may not be as conducive for secondary metabolite production in Annulohypoxylonchiangraiense. However, it is noteworthy that Hypoxylonthailandicum exhibited the highest number of well-defined peaks in YMB, indicating that this medium was optimal for the secondary metabolite production of this species. Similar results have been shown in other studies where they report unique metabolites in YMB, such as hypoxylonols, which are not as prominently produced in other media (Kuhnert et al. 2017). These findings highlight the influence of nutritional composition on secondary metabolite production in different fungal taxa. The variation in peak intensities and numbers across different media indicates that metabolite expression is medium-dependent and species-specific, and hence, must be carefully considered in natural product discovery and metabolic studies.
The mass spectrum obtained from the analysis suggested an identification of many compounds. Based on the HPLC profile of the Annulohypoxylonchiangraiense (MFLUCC 24-0606) isolate, the dominant compounds in the mycelium grown in MEB medium are 5-amino-2-(p-toluidino)benzenesulphonic acid (C13H14N2O3S), sulfamethoxazole sodium (C10H10N3NaO3S), N,N-dibutyl-3-methyl-5-(4,4,5,5-tetramethyl-1,3,2-dioxaborolan-2-yl)aniline (C21H36BNO2), and 2-(benzyloxy)-5-(4,4,5,5-tetramethyl-1,3,2-dioxaborolan-2-yl)pyridine (C18H22BNO3), detected at retention times of 6.0, 11.1, 19.0, and 20.0 minutes, respectively (Fig. 21a). Among these compounds, 5-amino-2-(p-toluidino)benzenesulphonic acid is an aromatic sulfonic acid primarily used in the dye and pigment industries (National Center for Biotechnology Information 2025a). Sulfamethoxazole sodium is the sodium salt form of sulfamethoxazole, which is used as a broad-spectrum sulfonamide antibiotic. Sulfamethoxazole sodium is mainly used in treating bacterial infections, including urinary tract infections (UTIs), respiratory infections, gastrointestinal infections, and skin infections (Kemnic and Coleman 2022).
The dominant compounds of the Annulohypoxylonchiangraiense (MFLUCC 24-0606) in the supernatant in MEB medium are Pramipexole (C10H17N3S), 5-Ethyl-1-ethoxymethyl-6-(3,5-dimethylphenylthio)uracil, 10-Deoxymethymycin (C25H43NO6), and Spisulosine (C18H39NO), detected at retention times of 5.0, 11.0, 12.5, and 19.75, respectively (Fig. 21b). Pramipexole is an orally active aminothiazole dopamine and is used to treat Parkinson’s disease (Bennett and Piercey 1999). 5-Ethyl-1-ethoxymethyl-6-(3,5-dimethylphenylthio)-2-thiouracil has antiviral, anticancer, and antimicrobial properties and is active against the majority of viruses (Balzarini et al. 1995). 10-Deoxymethymycin is a glycoside, a macrolide antibiotic that is active against gram-positive bacteria (National Center for Biotechnology Information 2025b). Spisulosine is a bioactive sphingoid and has been used as an antineoplastic agent (National Center for Biotechnology Information 2025c).
According to the HPLC profile in the Hypoxylonthailandicum (MFLUCC 25-0024) isolate, the dominant compounds in the mycelium in YMB medium are 2-Naphthalenepropanol, 6-methoxy-a-methyl-, hydrogen sulfate (C15H18O5S), N-({[Dimethoxy(methyl)silyl]oxy}methyl) aniline (C10H17NO3Si), Phenethyl rutinoside (C20H30O10), and Ethyl vanillin (C9H10O3), detected at retention times of 9.54, 13.55, 20.25, and 20.75 minutes, respectively (Fig. 21c). 2-Naphthalenepropanol, 6-methoxy-α-methyl-, hydrogen sulfate is a sulfate ester of a naphthalene-derived compound, potentially related to antibacterial naphthol derivatives (Roman et al. 2016). N-({[Dimethoxy(methyl)silyl]oxy}methyl)aniline is utilized in various industries due to its unique properties. This compound can be used as a versatile silane coupling agent, enhancing the adhesion, wetting, and durability of different materials. Therefore, N-({[Dimethoxy(methyl)silyl]oxy}methyl)aniline plays a crucial role in the production of adhesives, sealants, coatings, and composites (Inno Specialty Chemicals 2017). Phenethyl rutinoside is a glycoside that exhibits antioxidant activity (Wang et al. 2004). Ethyl vanillin belongs to the benzaldehydes and is used as an antioxidant and flavoring agent. This compound is widely used in the food industry as a food additive and spice in foods, beverages, cosmetics, and drugs (National Center for Biotechnology Information 2025d).
The dominant compounds of the Hypoxylonthailandicum (MFLUCC 25-0024) in the supernatant in YMB medium are 1,4-dimethyl-7-ethylazulene (C14H16), 3-{[(Benzyloxy)carbonyl]amino}-N-(tertbutoxycarbonyl)-L-alanine—N-cyclohexylcyclo hexanamine (1/1) (C28H45N3O6) and ethyl 2-phenyl-3-furancarboxylate (C13H12O3), detected at retention times of 9.54, 13.55, and 15.59 minutes, respectively (Fig. 21d). Ethyl 2-phenyl-3-furancarboxylate is an ester derivative of furan and can be used in pharmaceuticals, fragrances, and organic synthesis (Yannai 2004).
For Xylariachrysanthum (MFLUCC 21-0014), most of the bioactive compounds have been reported in MEB, followed by YMB. In MEB media, Xylariachrysanthum (MFLUCC 21-0014) showed phosphoenol-4-deoxy-3-tetrulosonate (C4H7O7P) at 1.5 min retention time. Phosphoenol-4-deoxy-3-tetrulosonate plays a key role as an intermediate compound in the biosynthesis of 3-deoxy-D-manno-octulosonic acid (KDO), an important molecule in bacterial biochemistry (Fig. 22a, b). Additionally, benzbromarone (C17H12Br2O3) has also been reported to be extracted from Xylariachrysanthum (MFLUCC 21-0014) in MEB medium, exhibiting antibacterial properties against Gram-positive pathogens, such as Enterococcusfaecalis, Staphylococcusaureus, S.epidermidis, and Streptococcusagalactiae (Meng et al. 2024).
According to the HPLC profile in the Xylariachrysanthum (MFLUCC 21-0014) isolate, the dominant compounds in the YMB medium are benzbromarone (C17H12Br2O3) (Fig. 22c, d). Benzbromarone, a benzofuran derivative, exhibits potential antibacterial activity against Gram-positive pathogens. Benzbromarone is used as a uricosuric drug that has been used in the treatment of gout and hyperuricemia (high levels of uric acid in the blood) (Heel et al. 1977; Meng et al. 2024). Additionally, monensin (C32H58O13), a polyether antibiotic widely used in veterinary medicine for its efficacy against certain Gram-positive bacteria and protozoa, was also reported in the Xylariachrysanthum (MFLUCC 21-0014) isolate in YMB medium (Łowicki and Huczyński 2013).
The HPLC analysis of Daldiniaeschscholtzii (MFLUCC 18-1207) cultured in MEB media revealed a peak corresponding to chlorfenson (C12H8Cl2O3S), with a retention time of 1.5 minutes. Chlorfenson is an organophosphorus compound, mainly used as a pesticide and acaricide (National Center for Biotechnology Information 2025e). (Fig. 22e, f). Additionally, in the Daldiniaeschscholtzii (MFLUCC 23–0263) isolate, the most abundant compound in the MEB medium is 13-methoxy-heneicosanoic acid (C22H44O3), followed by linoleic acid (C18H32O2). Both 13-methoxy-heneicosanoic acid and linoleic acid exhibited antimicrobial properties and have potential applications in research, cosmetics, pharmaceuticals, nutraceuticals, and industrial products (Kusumah et al. 2020). Furthermore, 3-methoxymandelic acid-4-O-sulfate (C9H10O8S) and formoterol (C19H24N2O4) have been identified in Daldiniaeschscholtzii on MEA, demonstrating medicinal properties.
Discussion
Thailand is part of the Indo-Malayan hub of biodiversity and is geographically located in the core of the Greater Mekong Subregion (Chaiwan et al. 2021). Thailand is well known to have tropical seasonal forests with rich and diverse plant communities. Therefore, a huge diversity of fungi can be found in Thailand (Tanaka et al. 2008; Vasilyeva et al. 2012; Hyde et al. 2018).
The present study includes the taxonomy of fungi in the families of Xylariales. Based on morphological aspects and phylogenetic analyses, we provided the taxonomic details of five novel species and ten new host/geographical records within Diatrypaceae, Hypoxylaceae, and Xylariaceae. Taxa were collected from February 2023 to July 2024 from forest areas with a high variety of trees and well-grown understory vegetation. These saprobic specimens were collected from different host families. We introduced two novel species in Hypoxylaceae: Annulohypoxylonchiangraiense from Tamarindusindica and Hypomontagnellahibisci from Hibiscus sp.; two novel species in Xylariaceae: Hypoxylonthailandicum from Bambusavulgaris and Stilbohypoxylonchiangraiense from Saraca sp.; and one novel species in Diatrypaceae: Diatrypellathailandica from Fabaceae sp. from Thailand. These five taxonomic novelties fulfilled the basic criteria for establishing new species, including distinct morphologies and multiple loci for the phylogenetic analyses, as described by Chethana et al. (2021b). Six new host records from Hypoxylaceae were recorded from Thailand: A.bahnphadengense from Berryacordifolia, A.crowfoothodgkiniae from Swieteniamacrophylla, A.purpureonitens from Sterculiatragacantha, A.spougei from Antidesmamadagascariense, A.violaceopigmentum from Syzygiumpolyanthum, and Hypomontagnellamonticulosa from Macarangapeltata. Our study also records A.crowfoothodgkiniae from Thailand for the first time. Astrocystisbambusae from Bambusavulgaris and Haloroselliniaxylocarpi from Arecaceae sp. were recorded as new host records from Xylariaceae in Thailand. In addition, we include Paraeutypellacitricola from Swieteniamacrophylla in Diatrypaceae as a new host record from Thailand.
In this study, we examined the antibacterial activity of the newly introduced species, Annulohypoxylonchiangraiense (MFLUCC 24-0606) and Hypoxylonthailandicum (MFLUCC 25-0024), using a preliminary screening test. Both species showed partial inhibition of the growth of Bacillussubtilis. Additionally, some existing Xylariales species, Daldiniaeschscholtzii (MFLUCC 18-1207) and Xylariachrysanthum (MFLUCC 21-0014), also exhibited slight inhibition zones against the bacterial pathogens. Many studies have been conducted on the antimicrobial properties of Annulohypoxylon, Daldinia, Hypoxylon, and Xylaria species (Quang et al. 2005; Yuyama et al. 2017; Pourmoghaddam et al. 2020; Gauchan et al. 2021; Segundo et al. 2022; Brooks et al. 2024; Cedeño-Sanchez et al. 2024; Chen et al. 2024; Yu et al. 2024).
The current study identified some of the secondary metabolites extracted from Xylariales isolates exhibiting biological properties, including antimicrobial, antibiotic, antiviral, and anticancer properties. All four tested fungal species exhibit antibacterial metabolites, including sulfamethoxazole sodium from Annulohypoxylonchiangraiense, 2-naphthalenepropanol, 6-methoxy-α-methyl-, hydrogen sulfate from Hypoxylonthailandicum, benzbromarone from Xylariachrysanthum, and linoleic acid from Daldiniaeschscholtzii. Therefore, this study will generate the initial data necessary for large-scale metabolite extractions for future applications.
The genus Annulohypoxylon can produce secondary metabolites with cytotoxic, antibacterial, and antioxidant properties (Maciel et al. 2018). Here, we recorded sulfamethoxazole sodium as an antibacterial compound from Annulohypoxylonchiangraiense. The genus Hypoxylon has been identified as the main producer of potential bioactive metabolites in Hypoxylaceae (Stadler et al. 2006). Hypoxyloninvadens produces flaviolin, which exhibits antibacterial activity against S.aureus (Becker et al. 2020). Additionally, 2-(4-(dimethylamino)phenyl-4H-chromen-4-one, 1-naphthalenol, 4-methoxy-, and hexadecyl methanesulfonate, produced by Hypoxylon species, showed antibacterial activity against human pathogenic bacteria, including B.subtilis, E.coli, S.aureus, and Pseudomonasaeruginosa (Mishra et al. 2020). In this study, the new antibacterial compound 2-Naphthalenepropanol, 6-methoxy-α-methyl-, hydrogen sulfate was derived from Hypoxylonthailandicum.
Additionally, a few compounds with antibacterial properties were reported in the Annulohypoxylonchiangraiense (MFLUCC 24-0606) isolate in MEB medium, including phytosphingosine (C18H39NO3), pipemidic acid (C14H17N5O3), tetranactin (C44H72O12), and neomycin palmitate (C39H78N6O15). Phytosphingosine plays an important role in innate immune defense against epidermal and mucosal bacterial infections (Başpınar et al. 2018). Pipemidic acid is a derivative of piromidic acid and is active against Gram-negative and Gram-positive bacteria (Shimizu et al. 1975). Tetranactin exhibits antibacterial, insecticidal, and mitogenic properties and inhibits the growth of Gram-positive bacteria (Ando et al. 1971). Neomycin palmitate is active against both Gram-positive and Gram-negative organisms and mediates its pharmacological action by binding to bacterial ribosomes and inhibiting protein synthesis (National Center for Biotechnology Information 2025f).
Based on LC–QTOF–MS analyses, the Hypoxylonthailandicum (MFLUCC 25-0024) isolate in YMB recorded several compounds with antibacterial properties, including pentamidine (C19H24N4O2), phytosphingosine (C18H39NO3), tiamulin (C28H47NO4S), tobramycin (C18H37N5O9), and dibucaine (C20H29N3O2). Pentamidine exhibits antibacterial activity against Gram-negative bacteria (Stokes et al. 2017). Tiamulin is used as an antibacterial drug in veterinary medicine for the treatment of swine dysentery caused by Serpulinahyodysenteriae (National Center for Biotechnology Information 2025g), while dibucaine shows antibacterial activity against Staphylococcusaureus (Chakraborty et al. 2024).
The genus Xylaria is an important source of a variety of bioactive secondary metabolites, including terpenoids, nitrogen-containing compounds, polyketides, and lactones. These metabolites exhibit a range of biological activities, such as antimicrobial, anti-inflammatory, antifungal, cytotoxic, immunosuppressive, and enzyme-inhibitory activities (Chen et al. 2024). Among terpenoids, xylareremophil exhibits weak antibacterial activity against Micrococcusluteus and Proteusvulgaris. In triterpenoids, xylarioxides E and F show antibacterial activity against Alternariaalternata, Curvularialunata, and Colletotrichumgloeosporioides. Additionally, kolokosides A and xylarchalasins A display antibacterial activity against B.subtilis, S.aureus, and E.coli, respectively (Chen et al. 2024). This is the first time benzbromarone has been recorded as an antibacterial compound from Xylariachrysanthum.
In the genus Daldinia, most chemical investigations have focused on D.concentrica and D.eschscholzii, resulting in different kinds of chemical compounds. These include alkaloids, terpenoids, polyketides, polyphenols, and steroids, which exhibit antimicrobial, anti-inflammatory, antifungal, antiviral, cytotoxic, and enzyme-inhibitory activities (Yu et al. 2024). In alkaloids, dalesindoloids A and in chromones, 8-O-methylnodulisporin F and nodulisporin H showed antibacterial activity against S.aureus. In polyketides, 5-hydroxy-2-methoxy-6,7-dimethyl-1,4-naphthoquinone and fusaraisochromenone showed antibacterial activity against Bacilluscereus and Enterococcusfaecalis, S.aureus, Escherichiacoli, and Pseudomonasaeruginosa, respectively. Additionally, Daldisones B showed moderate antibacterial activities against B.cereus, S.aureus, and Enterococcusfaecalis (Yu et al. 2024). However, this is the first time linoleic acid has been recorded from Daldiniaeschscholtzii.
Investigation of new host and geographical records of fungi is important for understanding fungal-host interactions, disease management, monitoring biodiversity, identifying fungal distribution patterns, and revealing hidden fungal diversity (Rathnayaka et al. 2024). The documentation of five new species and nine new host records, including one geographical record within Xylariales, emphasizes the fungal diversity in these provinces in Thailand. This study has led to an expansion of the taxonomic framework of Xylariales as well as to the exploration of fungal diversity in Thailand within various types of dead twigs/branches in the forest ecosystem.
Most undiscovered sexual forms of xylarialean taxa are presumably inconspicuous forms and may be isolated as endophytes. The endophytic life cycle may account for the large numbers of species found in some of these genera (Bhunjun et al. 2022, 2024). However, there are few studies that have been conducted on inconspicuous forms compared to conspicuous forms (Daranagama et al. 2016). Due to the insufficient fresh fungal collection, the taxonomic studies on inconspicuous xylarialean taxa have been limited. Additionally, the genera transferred to new families were previously accepted, although they have uncertain morphologies and phylogenies (Samarakoon et al. 2023). Therefore, it is important to collect more fresh fungal samples to resolve the taxonomic placement of inconspicuous xylarialean taxa.
Supplementary Material
Acknowledgments
Achala Rathnayaka expresses her profound gratitude to the Basic Research Fund (Fundamental Fund) of the National Science, Research, and Innovation Fund (NSRF) for their generous support and funding for the project entitled “Taxonomy, phylogeny, and chemo-profiling of selected families in Xylariales” (Grant Nos. 652A01006, 662A01003, and 672A01003). She also gratefully acknowledges the postdoctoral fellowship from Mae Fah Luang University under the Reinventing University grant. The authors extend their sincere thanks to W.A.S. Nuwanthika, A.J. Gajanayake, and D. Thakshila for their valuable suggestions and kind support. Special thanks are due to Shaun Pennycook for his assistance in selecting species epithets for the newly described taxa. Ausana Mapook and K.D. Hyde thank the National Research Council of Thailand (NRCT) for funding the project entitled “Total fungal diversity in a given forest area with implications towards species numbers, chemical diversity, and biotechnology” (Grant No. N42A650547).
Citation
Rathnayaka AR, Chethana KWT, Manowong A, Bhagya AT, Win H, Tun ZL, Mapook A, Hyde KD (2025) Taxonomy, phylogeny, and bioactive potential of Xylariales (Sordariomycetes, Ascomycota) from Thailand: novel species discovery, new host and geographical records, and antibacterial properties. MycoKeys 120: 35–117. https://doi.org/10.3897/mycokeys.120.155915
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Use of AI
No use of AI was reported.
Funding
This study was funded by the Basic Research Fund (Fundamental Fund) of the National Science, Research, and Innovation Fund (NSRF) under the project “Taxonomy, phylogeny, and chemo-profiling of selected families in Xylariales” (Grant Nos. 652A01006, 662A01003, and 672A01003).
Author contributions
Conceptualization: ARR, KWTC. Data curation: ARR. Formal analysis: ARR, KWTC. Funding acquisition: KWTC. Investigation: ARR, AM, KWTC. Methodology: ARR. Project administration: KWTC, AM. Resources: KWTC, AM, ATB, HW, ZLT. Software: ARR. Validation: ARR, KWTC. Visualization: ARR. Writing—original draft: ARR. Writing—review and editing: ARR, KWTC, KDH.
Author ORCIDs
Achala R. Rathnayaka https://orcid.org/0000-0001-8498-2715
K. W. Thilini Chethana https://orcid.org/0000-0002-5816-9269
Amuhenage T. Bhagya https://orcid.org/0009-0002-3988-298X
Hsan Win https://orcid.org/0009-0001-3830-7222
Zaw L. Tun https://orcid.org/0009-0001-3108-4425
Ausana Mapook https://orcid.org/0000-0001-7929-2429
Kevin D. Hyde https://orcid.org/0000-0002-2191-0762
Data availability
All of the data that support the findings of this study are available in the main text.
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