683 MycoKeys MycoKeys 120: 193-229 (2025) DOI: 10.3897/mycokeys.120.153906 Research Article Additions of New Endolichenic Fungi to Herpotrichiellaceae (Chaetothyriales, Ascomycota) from northern Thailand Chanokned Senwanna’'2®, Jaturong Kumla'23®, Pratthana Kodchasee2®, Nutchanan Duangkon™, Nakarin Suwannarach'23® 1 Office of Research Administration, Chiang Mai University, Chiang Mai 50200, Thailand 2 Center of Excellence in Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai 50200, Thailand 3 Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand Corresponding author: Nakarin Suwannarach (suwan.462@gmail.com) OPEN Qrceess Academic editor: Gerhard Rambold Received: 25 March 2025 Accepted: 10 July 2025 Published: 30 July 2025 Citation: Senwanna C, Kumla J, Kodchasee P, Duangkon N, Suwannarach N (2025) Additions of New Endolichenic Fungi to Herpotrichiellaceae (Chaetothyriales, Ascomycota) from northern Thailand. Mycokeys 120: 193-229. https://doi. org/10.3897/mycokeys. 120.153906 Copyright: © Chanokned Senwanna et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Endolichenic fungi associated with lichen thalli in Thailand are poorly known in terms of species diversity. During a study conducted in Chiang Mai Province, Thailand, in 2023, eight endolichenic fungal strains were isolated from healthy thalli of the foliose lichen, Parmotrema sp. These eight strains were identified as members of the family Herpo- trichiellaceae using a combination of three nuclear ribosomal regions (ITS, LSU, and SSU), tub2 sequence data, and morphological characteristics. Two strains of Atrokylindriopsis racemosospora and three strains of Veronaea endolichena were identified as new species within the Herpotrichiellaceae, while three other strains were identified as the previously known species Phialophora chinensis. This study provides the first report of P chinensis as an endolichenic fungal taxon and its first discovery in Thailand. Descriptions, illus- trations, and phylogenetic placements of these eight strains are provided. Additionally, a discussion and update on the ecology and genera within the Herpotrichiellaceae are included. The findings of this study offer valuable information that enriches the diversity of endolichenic species associated with lichens in Thailand and contributes to enhancing our understanding of the ecology and taxonomy of the Herpotrichiellaceae. Key words: Chaetothyriales, endolichenic fungi, fungal diversity, fungal taxonomy, lichen, new taxa Introduction The family Herpotrichiellaceae (Chaetothyriales, Eurotiomycetes) was es- tablished by Munk (1953) and typified by Herpotrichiella moravica, which is synonymized under Capronia pilosella based on the teleomorph-anamorph connection (Miller et al. 1987; Untereiner 1997). The sexual morph of Herpo- trichiellaceae is characterized by setose, ostiolate ascomata, bitunicate, sac- cate to ovoid asci with a thickened apex and long endotunica, and didymo- sporous, phragmosporous, or dictyosporous ascospores. The asexual morph consists of morphologically diverse dematiaceous hyphomycetes, predomi- nantly characterized by black yeast (Muller et al. 1987; Untereiner et al. 1995; Arzanlou et al. 2007; Tian et al. 2021). These fungi primarily reproduce asexu- 193 Chanokned Senwanna et al.: New endolichenic fungi from Thailand ally, and exhibit limited morphological characters (Untereiner et al. 1995; More- no et al. 2018; Tian et al. 2021). The internal transcribed spacer (ITS) and the large subunit (LSU) of ribosomal DNA sequences are commonly used to clar- ify the taxonomic placement of members in the Herpotrichiellaceae; however, many genera in this family have shown polyphyletic relationships (Quan et al. 2020; Tian et al. 2021; Torres-Garcia et al. 2023). Multi-gene sequence data are essential for resolving the morphological confusion among these genera and for establishing boundaries between genera and species. Species of Herpo- trichiellaceae are not only recognized for causing human infections but also for their extremotolerance, thriving in a wide range of environments worldwide, including extreme conditions of high or low temperature, nutrient scarcity, des- iccation, and solar irradiation (Miller et al. 1987; Untereiner 1997; Arzanlou et al. 2007; Teixeira et al. 2017; Muggia and Grube 2018; Quan et al. 2020, 2024; Chang et al. 2023; Thitla et al. 2023; Torres-Garcia et al. 2023; Diederich et al. 2024). Before this study, nineteen genera were recognized in this family, viz. Aciculomyces, Aculeata, Atrokylindriopsis, Capronia, Cladophialophora, Exophia- la, Fonsecaea, Marinophialophora, Melanoctona, Minimelanolocus, Petriomyces, Phialophora, Phaeoannellomyces, Pleomelogramma, Rhinocladiella, Thysano- rea, Uncispora, Valentiella, and Veronaea (Tian et al. 2021; Bezerra et al. 2022; Thitla et al. 2023; Torres-Garcia et al. 2023). Lichens are ubiquitous on a wide range of surfaces and can occur in various environmental conditions (Miller 2001; Molnar and Farkas 2010; Liicking et al. 2017; Muggia and Grube 2018). They serve not only as bioindicators for assess- ing air pollution and evaluating environmental health, but also serve as sources of secondary metabolites with diverse pharmaceutical properties (Casale et al. 2015; Crawford 2019; Panta 2020; Wethalawe et al. 2021; Zhao et al. 2021). Lichen thalli provide a habitat conducive to the growth of other photosynthetic and non-photosynthetic organisms, such as algae, bacteria, filamentous fungi, and yeast, while shielding the photobiont from external harm (Suryanarayanan and Thirunavukkarasu 2017; Hawksworth and Grube 2020). Many lichen-asso- ciated fungi, including both endolichenic and lichenicolous fungi, are currently being studied and described at both morphological and molecular levels (Wang et al. 2016; Cometto et al. 2023, 2024; Si et al. 2023). Lichenicolous fungi are visible inhabitants of lichen thalli, whether they are host-specific parasites, sap- rotrophs, broad-spectrum pathogens, or commensals (Lawrey and Diederich 2003; Diederich et al. 2018). In contrast, endolichenic fungi are non-obligate fungi that colonize the internal tissues of lichen thalli without causing visible symptoms or negative effects. Their occurrence is similar to that of endophytic fungi (Arnold et al. 2009; Gao et al. 2016; Kellogg and Raja 2017; Elkhateeb and Daba 2021). Since lichenicolous fungi have been shown to be present in asymptomatic lichens, differentiating between lichenicolous and endolichenic fungi can be scientifically challenging (Yang et al. 2022). Species diversity of en- dolichenic fungi influenced not only by lichen host species but also by the envi- ronment, habitat, host, and geographic distribution (U’Ren et al. 2012; Gao et al. 2016; Muggia and Grube 2018; Agrawal et al. 2020; Elkhateeb and Daba 2021). Moreover, studies on the diversity of endolichenic fungi in recent years have re- lied on culture-based and metabarcoding approaches (U’Ren et al. 2010; Zhang et al. 2016; Fernandez-Mendoza et al. 2017; Yang et al. 2021; Si et al. 2023). According to several previous studies, ascomycetes are the dominant group MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 194 Chanokned Senwanna et al.: New endolichenic fungi from Thailand of endolichenic fungi associated with lichens, followed by basidiomycetes and mucoromycetes. (Gao et al. 2016; Muggia and Grube 2018; Chakarwarti et al. 2020; Elkhateeb and Daba 2021; Cometto et al. 2024). Concurrently, various endolichenic fungal genera associated with lichen thalli worldwide have been identified, including Aspergillus, Chaetomium, Cladophialophora, Cladospori- um, Penicillium, Rhinocladiella, Trichoderma, and Xylaria (Muggia et al. 2017; Maduranga et al. 2018; Muggia and Grube 2018; Yang et al. 2021; Cometto et al. 2023, 2024). Thailand has a humid tropical climate and is recognized as one of the most biodiverse countries in the world (Tungmunnithum et al. 2022; Kaewsangsai et al. 2024). Numerous studies have been conducted on bacteria, insects, plants, fungi (endophytes, saprophytes, and pathogens), and lichens; however, the diversity of endolichenic fungi remains relatively unexplored (Ta- nasupawat 2009; Maneechan and Prommi 2015; Dathong 2016; Buaruang et al. 2017; Hyde et al. 2018; Jayawardena et al. 2020; Tayung 2022; Bamrungpanich- tavorn et al. 2023; Krongdang et al. 2023; Kaewsangsai et al. 2024; Nimnoi et al. 2024). Therefore, this research aimed to study the diversity of endolichenic fun- gi in Thailand. During a 2023 study of endolichenic fungi in northern Thailand, eight fungal strains belonging to the family Herpotrichiellaceae were isolated from foliose lichen thalli (Parmotrema sp.), including three strains of Phialopho- ra chinensis and five strains of unidentified fungal taxa. Based on morphology, growth temperature, and multi-gene phylogenetic analyses, five Herpotrichiella- ceous fungi are identified as novel taxa. Additionally, we provide an update on the ecology and genera within the Herpotrichiellaceae family. Materials and methods Lichen collection The foliose lichen thalli of Parmotrema sp. were collected from Chai Prakan and Mueang Chiang Mai Districts, Chiang Mai Province, northern Thailand, during June to July 2023. During the collection period, Chai Prakan District experienced daily rainfall of 81.2 mm, with a temperature range of 27 °C to 36 °C, whereas Mueang Chiang Mai District received daily rainfall of 153.2 mm, with tempera- tures ranging from 24 °C to 34 °C. Healthy specimens were carefully removed from tree bark using a sterile knife, transferred into individual plastic bags, the collection details were recorded (Rathnayaka et al. 2024), and the specimens were stored at 4 °C until processing within 48-96 hours after sampling. Fungal isolation The lichen thalli were carefully observed under an Olympus SZ40 stereo micro- scope to avoid damage and prevent contamination of any parts. Fungal isola- tion was performed following the protocols of Maduranga et al. (2018) and Si et al. (2023), with some modifications. The healthy lichen thalli were thoroughly washed in running water for 10 min to remove excess dirt and dried with sterile paper towels. The thalli were randomly dissected aseptically into 1 cm? frag- ments using a sterile razor blade under a stereo microscope, followed by sur- face sterilization with immersion in 70% ethanol for 1 min, 1% NaOCl for 1 min, and rinsing three times in sterile distilled water for 1 min. The upper and lower MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 195 Chanokned Senwanna et al.: New endolichenic fungi from Thailand cortex layers of surface-sterilized lichen fragments were carefully scratched under a stereo microscope using a sterile surgical blade and tweezers. The medulla tissue was carefully removed and cleaned with sterile distilled water. The fragment was then cut into a 0.5 cm? segment. Six lichen segments were evenly placed in a 9 cm Petri dish containing potato dextrose agar (PDA; Con- dalab, Laboratorios Conda S.A., Spain) and dichloran rose-bengal agar (DRBC; Difco, Becton, Dickinson and Company, USA). For each specimen, three repli- cations of the isolation plates were made. The plates were incubated at 25 °C until the growth of endolichenic fungi. The fungal hypha that grew from the segments was transferred onto freshly prepared PDA plates and incubated at 25 °C. The pure fungal strains were stored short-term on PDA slants at 4 °C and long-term in 20% glycerol at -80 °C at the Sustainable Development of Biolog- ical Resources culture collection (SDBR-CMU), Faculty of Science, Chiang Mai University, located in Chiang Mai Province, Thailand. Additionally, fungal strains were deposited and permanently maintained in a metabolically inactive state at the Chiang Mai University Biology Department’s Herbarium (CMUB), Chiang Mai University, Chiang Mai Province, Thailand. New fungal taxa were registered in the MycoBank database (MycoBank 2025). Morphological studies Pure fungal colonies were investigated on different media for induce pig- mentation and sporulation viz., corn meal agar (CMA; Difco, BBL™, USA), cornmeal dextrose agar (CMD; Difco corn meal agar + 2% dextrose), malt extract agar (MEA; Gibco, Life Technologies Corporation, USA), oatmeal agar (OA; Difco, Becton, Dickinson and Company, USA) and potato carrot agar (PCA; 200 g of each boiled and filtered carrots and potatoes, 17 g agar, 1 L distilled water). Mycelial plugs (3 mm x 3 mm) from 7-day-old PDA culture were cultured on each medium. The plates were incubated at 25 °C. In ad- dition, the cardinal growth temperatures at 4, 25, 30, and 35 °C of fungal strains were determined on MEA for two weeks in the dark. Microscopic features, including mycelia, branching patterns, and sporulating structures, were observed and photographed using a Nikon DS-Ri2 camera connected to a Nikon ECLIPSE Ni (Tokyo, Japan) compound microscope. The fungal structure measurement was performed using the Tarosoft Image Frame- work program (v. 0.9.0.7). Adobe Photoshop version 22.4.2 (Adobe Systems U.S.A.) was employed to create the photographic plates. DNA extractions, polymerase chain reaction, and sequencing DNA extractions were performed using the Fungal DNA Extraction Kit (FAVOR- GEN, Ping-Tung, Taiwan) from mycelium scraped from colonies grown on PDA using a sterile scalpel. DNA concentration and quality were determined by Nan- oDrop One® Microvolume UV-Vis Spectrophotometer (Thermo Scientific, Wilm- ington, DE, USA). The nuclear ribosomal internal transcribed spacer (ITS) region, 28S large subunit (LSU), 18S small subunit (SSU), and beta tubulin gene (tub2) were amplified using ITS4/ITS5 (White et al. 1990), LROR/LR5 (Vilgalys and Hes- ter 1990), NS1/NS4 (White et al. 1990), and Bt2a/Bt2b (Glass and Donaldson 1995) primers, respectively. Polymerase chain reaction (PCR) amplification was MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 196 Chanokned Senwanna et al.: New endolichenic fungi from Thailand performed using peqSTAR thermal cycler (PEQLAB Ltd., Fareham, UK) in a final volume of 20 uL containing 10 uL of 2 x Quick TaqTM HS DyeMix (TOYOBO, Ja- pan), 6 uL of sterile deionized water, 1 uL of 10 uM of each forward and reverse primer, and 2 uL of DNA. The PCR conditions for ITS, LSU, and SSU amplification were as follows: initial denaturing step of 95 °C for 5 min, followed by 35 cycles of denaturation at 94 °C for 30 s, annealing at 52 °C for 45 s, elongation at 72 °C for 1 min, and final extension at 72 °C for 10 min. The PCR conditions for tub2 amplification were as follows: initial denaturing step of 94 °C for 4 min, followed by 35 cycles of denaturation at 94 °C for 40 s, annealing at 52 °C for 30 s, elon- gation at 72 °C for 1 min, and final extension at 72 °C for 7 min. The PCR product was checked by 1% agarose electrophoresis gels under UV light. PCR clean-up Gel extraction NucleoSpin® Gel and PCR Clean-up Kit (Macherey-Nagel, Ger- many) was used to purify the PCR products according to the manufacturer’s protocol, which were sequenced by 1st BASE Company (Kembangan, Malaysia). New sequences generated in this study were deposited in GenBank. Sequence alignment and phylogenetic analyses The ITS, LSU, SSU, and tub2 sequence data were edited, quality-checked, and assembled using the SeqMan 5.00 software. The consensus sequences were BLAST-searched using the NCBI nucleotide database (http://blast.ncbi.nIm.nih. gov/) to assess the closely related species. Sequences generated in the analy- ses were chosen from related sequences of the genera in Herpotrichiellaceae which were derived from GenBank and recent publications (Su et al. 2023; Thit- la et al. 2023; Torres-Garcia et al. 2023). Cyphellophora laciniata (CBS 190.61) and C. suttonii (CBS 449.91) were selected as the out-group taxa (Table 1). The alignment of each locus was conducted using the MAFFT sequence alignment server (Katoh et al. 2019; http://mafft.cbrc.jp/alignment/server/) and improved manually where necessary in BioEdit v.7.0.9.1 (Hall 1999). Se- quence data for ITS and LSU were analyzed individually to examine the incon- gruence in the topology of the phylogenetic tree. Concatenated ITS, LSU, SSU, and tub2 sequence data were then analyzed and used to generate phylogenetic trees based on maximum likelihood (ML) and Bayesian inference (BI) methods. The ML tree was executed using the GTRGAMMA substitution model of nucleotide substitution and set to 1,000 bootstrapping replicates via the CIP- RES web portal. The BI tree was executed using a Markov Chain Monte Car- lo (MCMC) algorithm with MrBayes v. 3.2.6 (Ronquist et al. 2012), employing the best-fit model of sequence evolution determined by MrModeltest v. 2.3 (Nylander 2008). The GTR+I+G substitution model was the best-fitting model of sequence evolution, which was determined based on the Akaike Informa- tion Criterion (AIC) using the MrModeltest v2.3 (Nylander 2008) implemented in PAUP v. 4.0610 (Swofford 2002). Parameters for Bayesian inference: Four simultaneous Markov chains were set to run 20,000,000 generations, with the tree sampling every 100* generation, resulting in 200,000 trees. The first 25% of generated trees as part of a burn-in procedure were discarded, and the remain- ing trees were evaluated for posterior probabilities (PP) of a majority rule con- sensus tree. The resulting trees from ML and BI were displayed using FigTree v1.4.0 (Rambaut 2016) and modified using Adobe Illustrator Version 25.2.3 and Adobe Photoshop Version 22.4.2 (Adobe Systems, United States of America). MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 197 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Table 1. Taxa used in this study, along with their corresponding GenBank accession numbers. Ex-type strains are indicated with superscript “T”. The taxa obtained in this study are in bold. “—” is indicated the absence of sequence data in GenBank. GenBank Accession number Taxa name Strain number om tse) Atrokylindriopsis racemosospora PQ523731 Cladophialophora sribuabanensis OR139238 Cladophialophora subtilis KX822283 = Cladophialophora tengchongensis MG012731 MG012750 Cladophialophora thailandensis OR141865 OR139235 MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 198 Chanokned Senwanna et al Taxa name Phaeoannellomyces elegans Phaeoannellomyces elegans Phialophora americana Phialophora americana Phialophora americana Phialophora americana Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora chinensis Phialophora ellipsoidea Phialophora ellipsoidea Phialophora expanda Phialophora expanda Phialophora macrospora Phialophora macrospora Phialophora tarda Phialophora submersa Phialophora submersa Phialophora submersa Phialophora verrucosa Rhinocladiella anceps Rhinocladiella atrovirens Rhinocladiella mackenziei Rhinocladiella phaeophora Rhinocladiella quercus Thysanorea aquaticus Thysanorea clavatus Thysanorea papuana Thysanorea submersus Thysanorea thailandensis Uncispora sinensis Uncispora wuzhishanensis Valentiella maceioensis Valentiella maceioensis Veronaea aquatica Veronaea botryosa Veronaea botryosa Veronaea botryosa Veronaea botryosa Veronaea botryosa Veronaea compacta Veronaea endolichena Veronaea endolichena Veronaea endolichena Veronaea japonica Veronaea polyconidia Veronaea polyconidia Veronaea polyconidia .. New endolichenic fungi from Thailand Strain number GenBank Accession number yc easiaasy ws SSC SC~*S ewstanss7 | rozeues | keoaaso7 ewszeras | eustasse | eusiasoa easaoosy | eustasss | eusiasos wat t0e7s* | estasse | eusiasss am t0ers | estassy | eusiass7 em oasss | reersa0 | koous rewsigse | ss0779 | aassorre eww oor | reerss7 | ksooe emu ooiso | va700507 | kor? ewvoiosr | va700858 | ksoo8 emu ores: | varonons | kaos emu orss | va7005s8 | kos emu oresa | varono0s | ksois6 eases” | arosoze | arosoz wszmas7 [sta PCS” ewworeas | roetssa [Kuma ews van2587 | reese? | Ksss005 ewsarea” | arosozey [arson —ewstons7 =| ross = CCS ees irrse57 | xsossee [CS rwrrsoy | ownosasa | onoossae -wr8995 | ono0sa67 | onoosse7 | rwr9997 | ona0s068 | onoossae (ewsavrar | —asosrns [e665 ews soar | avasrsto | arosoze ees erase? | wrsnere | wxaosree ewsarzg6r | wosasr2 | waren rence isn067 | woresere | roms esse | _roanaoe | znanies ews teres? | vans | xan ~~ sauceases" | oases | anos ewstazssa | rozeazo | krone eassr290 | ween | wnerae20 | a eas 7i600" | evoenere | euoners MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 tub2 KF928571 EU514707 EU514708 EU514712 EU514713 KF971731 KF971765 KF971757 KM658123 KM658129 KM658101 KM658107 KM658111 PQ523732 PQ523733 PQ523734 EU514715 KU306354 KF971734 KF971737 EU514714 KU306355 KU306347 ON667800 ON667798 ON667799 KF971761 GU079661 MW248394 JN112505 KF928557 PQ523735 PQ523736 PQ523737 OR817660 OR817661 OR817659 199 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Results Phylogenetic analyses The concatenated ITS, LSU, SSU, and tub2 sequence dataset consists of 114 representative strains of species in the families Herpotrichiellaceae and Cyphellophoraceae (outgroup). The total alignment length comprises 3,413 characters (ITS: 1-779; LSU: 780-1,644; SSU: 1,645-2,872; tub2: 2,873- 3,413), including gaps. The topologies of the trees generated from ML and BI analyses were congruent. The resultant ML tree is shown in Fig. 1. The best RAxML tree with a final likelihood optimization value of -32004.987525 is presented. The matrix contained 1,459 distinct alignment patterns, with 38.84% of the characters being undetermined or gaps. Estimated base frequencies were as follows: A = 0.251383, C = 0.234823, G = 0.265952, T =0.247842; substitution rates AC = 1.510868, AG = 3.198001, AT = 1.566806, CG = 0.906099, CT = 6.013302, GT = 1.000000; gamma distribution shape parameter a = 0.548282. The average standard deviation of split frequen- cies was 0.009767 at the end of total MCMC generations. The phylogenetic tree, based on the analysis of a combined ITS, LSU, SSU, and tub2 sequence data, shows the relationships of taxa within the family Her- potrichiellaceae (Fig. 1). The results indicated that all eight endolichenic fungal strains obtained in this study belong to the family Herpotrichiellaceae. Three strains (SDBR-CMU504, SDBR-CMU505, and SDBR-CMU506) clustered within the Phialophora species and were found to be phylogenetically related to P. chin- ensis, with 86% ML and 1 PP statistical support. Interestingly, the other five strains formed distinct lineages from previously known taxa. Two fungal strains (SDBR-CMU502 and SDBR-CMU503) of a novel taxon formed a monophyletic clade clustering with Atrokylindriopsis setulosa, with 100% ML and 1 PP support values. In addition, three remaining strains (SDBR-CMU507, SDBR-CMU508, and SDBR-CMU509) are also introduced as a new species, which forms a distinct lineage in Veronaea with 92% ML and 1 PP and is in a sister clade to V. botryosa. Taxonomic descriptions Atrokylindriopsis racemosospora Senwanna, J. Kumla & N. Suwannar., sp. nov. MycoBank No: 858499 Fig. 2 Etymology. In reference to the spore arrangement resembling a raceme form. Type. THAILAND * Chiang Mai Province: Chai Prakan District, Nong Bua Sub- district, endolichenic from the medulla of foliose lichen (Parmotrema sp.) on Prunus domestica, 19°42'23"N, 99°1'32"E, elevation 1160 m, 2 June 2023, C. Senwanna and N. Suwannarach, CMUB40067 (Holotype, preserved in a meta- bolically inactive state. Ex-type living culture LCO5-1 = SDBR-CMU502). Cultural characteristics. Colonies on different agar media were incubat- ed in the dark at 25 °C for 2 months; colonies flat, irregular, with edge undu- late, velvety; on PDA (18 to 27 mm in diameter) surface grayish brown, light brown at the margin, reverse olivaceous black, light brown at the margin; on MEA (18 to 23 mm in diameter) surface grayish brown, dark brown to black at the margin, reverse olivaceous black, producing brown pigment in agar; MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 200 Chanokned Senwanna et al.: New endolichenic fungi from Thailand rancor Gladophialophora carrionll GBS 160.54 covo.os| | | Cl@dophialophora yegresil CBs 114405 Cla yaphara Cladophialophora rupestricola SDBR-CMU446 Cladophialophora tengchongensis CGMCC 3.15201 Cladophialophora nyingchiensis CGMCC 3.17330 Cladophialophora denticulata FMR 18992 Cladophialophora arxii CBS 306.94 Rhinocladiella anceps CBS 181.65 Rhinocladiella phaeophora CBS 496.78 Rhinocladiella ocladiella mackenziel CBS 650.93 97/0.99) Thysanorea aquaticus MFLUCC 15-0414 96/1|_ Thysanorea submersus KUMCC 15-0206 Thysanorea thailandensis MFLUCC 15-0974 3 Cyphellophora laciniata CBS 190.61 Cyphellophoraceae, OUTGROUP 0.07 Figure 1. Phylogram generated from maximum likelihood analysis of members in Herpotrichiellaceae based on a com- bined ITS, LSU, SSU, and tub2 sequence dataset. Bootstrap values = 50% MLBS and Bayesian posterior probabilities (PP) = 0.90 are shown above nodes and defined as ML/PP. The tree is rooted to Cyphellophora laciniata (CBS 190.61) and C. suttonii (CBS 449.91). The fungal strains obtained in this study are blue. Ex-type strains are in bold. The scale bar represents the expected number of nucleotide substitutions per site. MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 201 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Figure 2. Atrokylindriopsis racemosospora (SDBR-CMU502, ex-type). A-F. Colonies on PDA, MEA, OA, PCA, CMD and CMA, re- spectively, after 48 d at 25 °C; G, H. Conidiophores sporulating on PCA; I. Conidiophores sporulating on OA; J. Conidiophores sporulating on CMA; K, L. Conidiophores and conidia; M-N. Hyphal coil; 0, P Hyphal anastomosis; Q, R. Terminal chlamydo- spores; S-W. Conidiophores with conidiogenous loci; X-Y. conidia. Scale bars: 10 um (G-J); 20 pm (K-L); 10 um (M-Y). MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 502 Chanokned Senwanna et al.: New endolichenic fungi from Thailand on OA (17 to 23 mm in diameter) surface grayish brown, reverse olivaceous black, sporulation absent; on PCA (17 to 20 mm in diameter) surface and re- verse brownish gray, reverse olivaceous black, sporulation absent. On CMD (15 to 20 mm in diameter) surface and reverse grayish brown; and on CMA (20 to 27 mm in diameter) surface and reverse brownish grey, sporulation absent. Asexual morph in vitro dematiaceous hyphomycetes. Hyphae 1-2.5 um wide, pale brown, simple to branched, septate, smooth, thin-walled, coiling, anasto- mosis observed. Conidiophores (10—)20-111(-—153) x (1.4-)1.8-2.6(-2.9) um (x = 53.17 x 2.15, n = 40), macronematous, monomematous, straight or slightly flexuous, unbranched, continuous or 1-3 septate, dark brown, paler terminally, smooth-, thick-walled. Conidiogenous cells (6.6—)15.5-36(-45.5) x (1.4-)1.8- 2.6(—2.9) um (X = 25.5 x 2.16, n = 40), integrated, polyblastic, terminal to mostly intercalary, proliferating sympodial and producing conidia from short denticles, subcylindrical, pale brown to brown, fertile parts subhyaline; denticles scat- tered, slightly darkened, 0.4—1 um wide. Conidia (2.5-)2.8—4.4(—5) x (1.8-)2- 3.3 um (X = 3.72 x 2.68, n = 40), abundant, obovoid or subglobose, with a round apex, and slightly truncate base, aseptate, subhyaline to pale brown, smooth- walled, with inconspicuous conidial scars, 0.5-1 um wide. Chlamydospores rare, solitary or in chains, terminal, globose to pyriform, without or one-septate, pigmented, dark brown, smooth-, thick- walled. Sexual morph unknown. Cardinal temperatures for growth on MEA after two weeks (mm). Optimum at the range of 25 °C to 30 °C (10 to 14). No growth 4 °C and 35 °C. Additional materials examined. THAILAND * Chiang Mai Province, Chai Pra- kan District, Nong Bua Subdistrict, endolichenic from the medulla of foliose lichen (Parmotrema sp.) on Prunus domestica, 19°42'23"N, 99°1'32"E, eleva- tion 1160 m, 2 June 2023, C. Senwanna and N. Suwannarach, living culture (LCO5-3 = SDBR-CMU503). Additional GenBank numbers. act and tef7 for SDBR-CMU502: PQ523738 and PQ523739; tef1 for SDBR-CMU503: PQ523740. Ecology and distribution. Endolichenic fungi from the medulla of foliose li- chen (Parmotrema sp.) in Thailand. Notes. Based on a blast search of the NCBI’s GenBank nucleotide database of the ITS sequence, Atrokylindriopsis racemosospora has the closest match with At. setulosa (strain HMAS245592; KP337330, ex-type) with 99.33% simi- larity and is similar to Aciculomyces restrictus (strain FMR 18994; ON009870, ex-type) with 93.79% similarity (identities = 468/499, 10 gaps) and Exophiala siamensis (strain SDBR-CMU417; NR_184988, ex-type) with 92.39% similarity (identities = 583/631, 16 gaps). The closest matches using the LSU sequence are At. setulosa (strain HMAS245592; KP337329) with 100% similarity (identi- ties = 548/548, 0 gap), Ex. ramosa (strain FMR 18632; ON009933, ex-type) with 98.97% similarity (identities = 865/874, 1 gap), and Melanoctona tectonae (strain MFLUCC 12-0389; NG_059687) with 98.43% similarity (identities = 879/893, 2 gaps). The closest matches using SSU sequence are Ex. siamensis (strain SD- BR-CMU417; ON555826) with 99.53% similarity (identities = 1055/1060, 1 gap), Capronia dactylotricha (strain CBS 604.96; NG_062636) with 98.45% similarity (identities = 1082/1099, 1 gap), and E. yunnanensis (strain YMF 1.06739, ex-type) with 98.31% similarity (identities = 1049/1067, 1 gap). The closest matches MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 503 Chanokned Senwanna et al.: New endolichenic fungi from Thailand using the tub2 sequence are Biscogniauxia arima (strain YMJ 122; AY951672) with 93.08% similarity (identities = 121/130, 1 gap) and B. mediterranea (strain ISN9LDC31; 0Q942633) with 92.37% similarity (identities = 121/131, 1 gap). A combined multilocus-based phylogenetic analysis showed that both strains (SDBR-CMU502 and SDBR-CMU503) of Atrokylindriopsis racemosos- pora formed sister taxon to At. setulosa (HMAS245592, ex-type strain) with 100% ML and 1 PP statistical support, and also clustered with Aciculomyces restrictus (FMR 18994, ex-type strain) with 95% ML and 1 PP statistical support (Fig. 1). The ITS and LSU nucleotide sequence comparisons reveal 3/580 and 0/548 base pair differences with At. setulosa, respectively. In contrast, a com- parison of ITS, LSU, tef7, and tub2 base pairs shows that At. racemosospora differs from Ac. restrictus by 34/494 bp of ITS, 8/770 bp of LSU, 38/158 bp of tef1, and 108/430 bp of tub2. The morphology of Atrokylindriopsis differs from that of Aciculomyces in having unbranched, macronematous conidiophores, monophialidic conidiogenous cells, pigmented, septate, setulate conidia (Ma et al. 2015; Torres-Garcia et al. 2023). Atrokylindriopsis racemosospora shares similar features, including sympodial proliferation, denticulate conidiogenous cells, and aseptate, obovoid or subglobose conidia, with species Ac. restrictus (Torres-Garcia et al. 2023), but Ac. restrictus has longer conidiophores (10- 153 x 1.4-2.9 um vs 19-105.5 x 1.5-2.5 um) and smaller conidia (2.5—5 x 1.8- 3.3 um vs 2-4 x 1.5-2.5 um). Moreover, the conidiogenous cells and conidia formation of At. racemosospora is more abundant than that of Ac. restrictus, and the conidial scars are not conspicuous. In addition, At. setulosa differs from At. racemosospora by its larger, cylindrical to clavate or rounded-cuboid conidia, which have 4-5 longitudinal eusepta and are attached to the conidiog- enous locus at the midpoint of their long side, appearing to form a ‘T’ (Ma et al. 2015). These morphological characteristics clearly distinguish At. racemosos- pora from Ac. restrictus and At. setulosus. Phialophora chinensis Ya L. Li, de Hoog & R.Y. Li, in Li, Xiao, de Hoog Wang, Wan, Yu, Liu & Li, Persoonia 38: 11 (2016) MycoBank No: 815345 Fig. 3 Cultural characteristics. Colonies on different agar media were incubated in the dark at 25 °C for 1 months; colonies flat, irregular, with edge entire, velvety; on PDA (39 to 44 mm in diameter) surface grayish brown, dark brown to black at the margin, reverse olivaceous black; on MEA surface Blackish Grey, black at the margin, reverse olivaceous black; on MEA (49 to 53 mm in diameter) surface Blackish Grey, black at the margin, reverse olivaceous black; on OA (69 to 74 mm in diameter) surface grayish brown, reverse olivaceous black; on PCA (46 to 55 mm in diameter) surface and reverse grayish brown, dark brown at the margin, reverse olivaceous black, sporulation absent; on CMD (42 to 50 mm in diameter) surface and reverse brownish gray; and on CMA (33 to 41 mm in diameter) surface and reverse brownish grey, sporulation ab- sent. Asexual morph in vitro dematiaceous hyphomycetes. Hyphae 1-3.4 um wide, subhyaline to light brown, simple to branched, septate, smooth-, thin- walled, coiling, anastomosis observed. Conidiophores (4—)5.5-31(—45.3) MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 504 Chanokned Senwanna et al.: New endolichenic fungi from Thailand AE ® AE. G@ AHy Ab 0 7A JAK. Figure 3. Phialophora chinensis (SDBR-CMU506, new host record). A-F. Colonies on PDA, MEA, OA, PCA, CMD and CMA, respectively, after 30 d at 25 °C; G. Conidiophores sporulating on CMA; H. Conidiophores sporulating on PCA; I-M. Hyphal anastomosis; N-P. Hyphal coil; Q-AA. Phialides at different stages of development; AB—AD. Conidia; AE-AK. Chlam- ydospores (intercalary and chains formation). Scale bars: 100 um (G, H); 10 pm (I-AA); 5 um (AB-AD); 20 pm (AE-AK). x (2-)3-4.3(-4.8) um (x = 17 x 3.6, n = 55), micro- or semi-macronema- tous, straight, simples or poorly branched, septate, slightly constricted at septa, smooth; micronematous conidiophores consisting in conidiogenous cells growing directly from vegetative hyphae, lateral or terminal. Phialides 4-11(-15.5) x 2.3-4(-5) um (Xx = 8.3 x 3.3, n = 75), regularly flask-shaped to elongate-ampulliform or subulate, with an apical conspicuous collarette; collarettes (1.6-)2-3.9(-4.8) x (1.6-)2-3.9(-4.2) um (x = 3 x 3, n = 65), MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 505 Chanokned Senwanna et al.: New endolichenic fungi from Thailand usually funnel-shaped, darker than the rest of the phialide. Conidia 2—4(—5) x 1.6-2.7(-3.9) um (X = 3.3 x 2.2, n = 75), hyaline to subhyaline, mostly broad- ly ellipsoidal, more rarely obovoidal, smooth-walled. Chlamydospores 9.5- 20(-—23) um wide, abundantly produced in aerial hyphae, mostly intercalary, solitary or in branched chains, sub globose to ellipsoidal or barrel-shaped, pigmented, light to dark brown, smooth-, thick-, dark-walled, without or one-septate, constricted near the septa. Sexual morph unknown. Cardinal temperatures for growth on MEA after two weeks (mm). Opti- mum at the range of 25 °C to 30 °C (19 to 24), maximum 35 °C (17 to 19). No growth 4 °C. Materials examined. THAILAND * Chiang Mai Province: Mueang Chiang Mai District, Suthep Subdistrict, endolichenic from the medulla of foliose lichen (Parmotrema sp.) on unidentified tree trunk, 18°48'27'"N, 98°56'37.7'E, elevation 343 m, 26 June 2023, C. Senwanna, living culture: LC10-10 = SDBR-CMU504, LC10-11 = SDBR-CMU505, and LC10-17 = SDBR-CMU506. Ecology and distribution. Endolichenic fungi from the medulla of foliose li- chen (Parmotrema sp.) in Thailand (this study); pathogenic as chromoblasto- mycosis in human in China and Mexico (Li et al. 2017; Ahmed et al. 2021), as fungal keratitis in India and USA (Ply et al. 2023; Mitra et al. 2024), as phaeo- hyphomycosis in France (Pruvot et al. 2023); saprobic from bamboo in China, from the environment in Japan, from plant materials in Brazil and China, and soil in China and from wheat straw in Brazil (Li et al. 2017). Notes. A BLASTn search using ITS and tub2 sequence data in NCBI has revealed sequence similarities of 98.98-99.83% and 98.46-99.78% be- tween our strains (SDBR-CMU504, SDBR-CMU505, and SDBR-CMU506) and Phialophora chinensis strains. While the closest matches using the LSU se- quence are P ellipsoidea (strain MUCL 9768; AF050282) with 99.89% sim- ilarity (identities = 885/886, no gap), PR macrospora (strain MUCL 15541; EU514701), and P. americana (strain UAMH 10872; EU514691) with 99.77% similarity (identities = 864/866, no gap). The closest matches using the SSU sequence are Capronia semiimmersa (strain UAMH 10872; JN941209) with 99.71% similarity (identities = 1018/1021, 1 gap), P verrucosa (strain AF- TOL-ID 670; EF413614) with 99.60% similarity (identities = 992/996, 2 gaps), and P americana (strain CBS 840.69; AY554291) with 99.33% similarity (identities = 1033/1040, 4 gaps). In our multigene phylogenetic study, strains SDBR-CMU504, SDBR-CMU505, and SDBR-CMU506 form a clade with close affinity to P chinensis (Fig. 1). How- ever, the species segregation within the taxa is not discrete in a multigene phy- logeny. Comparing the ITS, LSU, SSU, and tub2 regions between our strains and CBS 140326 (type strain), only 1 bp difference was found in the ITS, 3 bp differ- ence in LSU and SSU, and no base pair differences in the tub2. The morphology of our strains resembles the species description of P chinensis provided by Li et al. (2017), except for the appearance of the conidiophores and chlam- ydospores and the lack of budding conidia. Although our strains have shorter conidia compared to P. chinensis CBS 140326, the holotype isolated from skin lesions of a human chromoblastomycosis patient (2-5 x 1.6-3.9 um vs 3-6 x 2.0-5.5 um), the detail of other structures (i.e., collarette, conidiophore, conid- iogenous cells and chlamydospores) was not mentioned. Hence, an updated morphology for the species P. chinensis is provided. MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 206 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Veronaea endolichena Senwanna, J. Kumla & N. Suwannar., sp. nov. MycoBank No: 856324 Fig. 4 Etymology. Refers to the host substrate. Type. THAILAND * Chiang Mai Province: Mueang Chiang Mai District, Suthep Subdistrict, endolichenic from the medulla of foliose lichen (Parmotrema sp.) on unidentified tree trunk, 18°48'27"N, 98°56'37.7"E, elevation 343 m, 26 June 2023, C. Senwanna, CMUB40068 (Holotype, preserved in a metabolically inac- tive state. Ex-type lliving culture LC10-9-2 = SDBR-CMU507). Cultural characteristics. Colonies on different agar media were incubated in the dark at 25 °C for 2 weeks; colonies flat, irregular, with edge entire, velvety; on PDA (63 to 67 mm in diameter) surface grayish brown, dark at the middle and margin, reverse olivaceous black; on MEA (42 to 59 mm in diameter) surface brownish gray, reverse olivaceous black, producing dark brown pigment in agar; on OA (57 to 60 mm in diameter) surface brownish gray, reverse olivaceous black; on PCA (61 to 63 mm in diameter) surface and reverse brownish gray, reverse olivaceous black, sporulation absent; on CMD (54 to 55 mm in diam- eter) surface and reverse grayish brown; and on CMA (51 to 58 mm in diame- ter) surface and reverse brownish gray, sporulation absent. Asexual morph in vitro dematiaceous hyphomycetes. Hyphae 1-—3.8 um wide, subhyaline to light brown, simple to branched, septate, smooth, thin-walled, coiling observed. Co- nidiophores (23-)38-200(—264) x 2.3-3.7(—4.9) um (xX = 118.6 x 3.1, n = 25), macronematous, mononematous, erect, straight or slightly flexuous, branched, septate, cylindrical, rough-, thick-walled, light to dark brown. Conidiogenous cells (7.5-)12—102(-136.5) x 2.1-3.5(-3.9) um (X = 53.5 x 2.9, n = 30), in- tegrated, polyblastic, terminal to mostly intercalary, cylindrical, pale brown to brown, fertile parts subhyaline, rachis with crowded, flat to slightly prominent, unthickened scars. Conidia (5-)6.5-11(-—13.5) x 2.5-4(—4.6) um (X = 8.5 x 3.3, n= 90), solitary, cylindrical to pyriform, rounded at apex, truncate at base, lower cell longer and wider than upper one, with a prominent scar, 0.8-1.9 um wide, pale brown, without or one median septate, constricted at the septa, smooth- walled. Chlamydospores absent. Sexual morph unknown. Cardinal temperatures for growth on MEA after two weeks (mm). Optimum 25 °C (35 to 40), maximum 30 °C (20 to 21). No growth 4 °C and 35 °C. Additional materials examined. THAILAND * Chiang Mai Province, Mueang Chiang Mai District, Suthep Subdistrict, endolichenic from the medulla of foliose lichen (Parmotrema sp.) on unidentified tree trunk, 26 June 2023, 18°48'27"N, 98°56'37.7"E, elevation 343 m, C. Senwanna, living culture LC10-2 = SDBR-CMU508 and LC10-12 = SDBR-CMU509. Ecology and distribution. Endolichenic fungi from the medulla of foliose li- chen (Parmotrema sp.) in Thailand. Notes. The closet match in a BLASTn search in GenBank with the ITS, LSU, and tub2 sequence of V. endolichena had highest similarity to V. bo- tryosa strain GZCC:19-0557 (OP377853, with 99.65%, identities = 577/579, 2 gaps), strain CBS 127264 (MH875936, with 100%, identities = 908/908, no gap) and strain CBS 121506 (JN112502, with 93.61%, identities = 381/407, no gap), respectively. While the match using SSU sequence are Ex. yunnanensis (strain YMFT 1.06739; MZ781222, holotype) with 99.72% (identities = 1053/1056, MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 507 Chanokned Senwanna et al.: New endolichenic fungi from Thailand tively, after 14 d at 25 °C G. Conidiophores sporulating on PCA; H. Conidiophores sporulating on CMA; I-O. Conidiophores at different stages of development; P—Q. Hyphal coil; R-Y. Conidiophores with conidiogenous loci; W-A.C Conidia. Scale bars: 100 um (G-1I); 50 um (J-O); 10 um (P-V); 5 um (W-AC). no gap), Thysanorea amniculi (strain SGT69-1; OP378033, holotype) with 99.71% (identities = 1019/1022, no gap) and Ex. aquamarine (strain IMP- BG-H0001; MH813287) with 99.63% (identities = 1072/1076, no gap). Multigene MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 208 Chanokned Senwanna et al.: New endolichenic fungi from Thailand phylogenetic analysis of the combined dataset revealed that three strains (SD- BR-CMU507, SDBR-CMU508, and SDBR-CMU509) of V. endolichena clustered as a sister taxon to V. botryosa; however, the ITS and tub2 sequences of V. en- dolichena differ from V. botryose in 6 bp/610 and 25/393 bp, respectively. Mor- phologically, V. endolichena differs from V. botryosa in longer conidiophores, the number of septa (0-1 vs 0-3 septa) and size of the conidial scar (0.8-1.6 vs 0.5 um) (Table 2) (Arzanlou et al. 2007, Yang et al. 2023). Discussion Several studies have revealed that lichen thalli host diverse microbial com- munities, including fungi (Oh et al. 2020; Yang et al. 2021, 2022; Diederich et al. 2022, 2024; Miral et al. 2022; Cometto et al. 2023; Si et al. 2023). Endol- ichenic fungi that reside inside lichens without causing any symptoms not only thrive in harsh environments but also generate unique pharmacological activities and bioactive compounds similar to those of lichens (Kellogg and Raja 2017; Singh et al. 2017; Suryanarayanan and Thirunavukkarasu 2017; Moreno et al. 2018; Agrawal et al. 2020; Yang et al. 2021; Zhang et al. 2021, 2024; Perera et al. 2022; Cometto et al. 2023). The presence of endolichenic fungi within lichen thalli suggests a symbiotic or commensal relationship, whereby these fungi enhance lichen health and resilience by producing pro- tective secondary metabolites and potentially participating in organic mat- ter decomposition and nutrient recycling within the thallus or its substrate (Tsurykau and Etayo 2017; Muggia and Grube 2018; Diederich et al. 2024). These functional roles may help explain the ecological success of lichens in diverse and often extreme environments, highlighting the significance of endolichenic fungi within lichen ecosystems and underscoring the need for further studies to elucidate their functions. Most endolichenic fungi have been documented from tropical areas, primarily China and India, and it is anticipated that research in other areas will yield a wealth of new species, biological compounds, and ecological data (Kellogg and Raja 2017; Sury- anarayanan et al. 2017; Agrawal et al. 2020; Si et al. 2023). Recent publica- tions have identified eight new Chaetothyriales species (Cladophialophora endolichena, C. guttulate, C. haematommatis, C. heterodermiae, C. holoseri- cea, C. olivacea, C. yunnanensis, and Paracladophialophora lichenicola) and a new genus, Intumescentia in Teratosphaeriaceae, based on phylogeny and morphology and other known species in Eurotiomycetes, Dothideomycetes, and Sordariomycetes from lichen thalli (Chang et al. 2023; Cometto et al. 2023, 2024; Si et al. 2023; Diederich et al. 2024). Likewise, studies on sec- ondary metabolites and their activities, isolated from endolichenic fungi growing in various environments, are increasing (Zhang et al. 2021; Toure et al. 2022; Varli et al. 2022, 2023; Zhang et al. 2024). Therefore, the investiga- tion of new ecosystems, particularly in tropical countries, is likely to reveal novel endolichenic fungi with unique potential, thereby enhancing scientific knowledge. This research employed a culture-based approach, focusing on endolichenic fungi, and is part of an ongoing study examining the diversity of endolichenic fungi in northern Thailand. Three endolichenic fungal species in Herpotrichiellaceae were identified based on morphology and phylogenet- ic evidence from combined datasets of ITS, LSU, SSU, and tub2 genes. MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 909 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Table 2. Veronaea species with a synopsis of the characteristics and relevant references. Veronaea species V. aquatica V. botryosa V. caricis V. carlinae V. compacta V. coprophila V. endolichena V. ficina V. filicina V. gobica V. grewiicola Conidiophore Up to 280 x 2.5-4 um. Erect, the lower part is usually straight, and the upper half is usually flexuous, usually loosely branched, sometimes geniculate, pale brown to dark brown, smooth-walled. Up to 250 x 2-3 um. Erect, straight or flexuose, unbranched or occasionally loosely branched, sometimes geniculate, pale brown to olivaceous brown, smooth-walled. Up to 550 x 4-6 um. Erect, straight or flexuose, sometimes swollen at the base to 10-14 um, with numerous scars towards the apex, unbranched or occasionally loosely branched, sometimes geniculate, mid to dark brown, smooth-walled. Up to 130 x 2-4 um. Simple or loosely branched, straight or flexuous, septate, pale brown, smooth, with numerous scars. Up to 60 um long. Slightly differentiated from vegetative hyphae, lateral or occasionally terminal, often wider than the supporting hypha, up to 4 um wide, unbranched or branched at acute angles, with 1-3 additional septa, cells often inflated and flask-shaped, pale brown. Up to 350 x 3—4.5 um. Straight or flexuous, septate, mid to dark brown, paler towards the apex where there are a number of small, flat scars. (23-)38-200(-264) x 2.3-3.7(- 4.9) um. Erect, straight or slightly flexuous, branched, septate, cylindrical, light to dark brown, rough-, thick-walled. 30-120 x 2.5-4.75 um. Superficial, arising singly as lateral or terminal branches from external hyphae ranging from macronematous to mononematous, erect, straight to flexuous or slightly curved, smooth- walled, unbranched, septate with brown base and paler apex. 400-800 x 2-2.5 um. Base rarely swollen, rarely branched, sparsely septate, at first thin-walled, roughened walls, olivaceous then to brown. Up to 350 x 1.5-2.5 um. Straight or flexuous, septate, smooth, with numerous minute scars towards the apex, brown. 42-104 x 2.5-6.3 um. Sometimes curved, smoothwalled, branched, 2-5 septa, pale brown to brown base and paler apex. Conidiogenous cells (3-)10-30 x 2-3.5 um. Terminally integrated, polyblastic, occasionally intercalary, cylindrical, variable in length, pale brown, later often becoming septate, fertile part subhyaline, wide at the basal part, rachis with crowded, flat to slightly prominent, faintly pigmented; scars flat, slightly pigmented, not thickened, about 0.65 um diam. 10-100 um long. Terminal, occasionally intercalary, cylindrical, pale brown, later often becoming septate, fertile part subhyaline, often as wide as the basal part, rachis with crowded, flat to slightly prominent, faintly pigmented, unthickened scars. No information available No information available Up to 10 um long. Terminal, occasionally intercalary, variable in length, cylindrical to doliiform or flask-shaped, with hardly prominent denticles, pale brown; scars flat, slightly pigmented, not thickened, about 0.5 um diam. No information available (7.5-)12-102(-136.5) x 2.1-3.5(-3.9) pm. Integrated, polyblastic, terminal to mostly intercalary, cylindrical, pale brown to brown, fertile parts subhyaline, rachis with crowded, flat to slightly prominent, unthickened scars. Integrated, polyblastic, terminal to predominantly intercalary, sympodial, cylindrical, cicatrized with thickened scars. No information available No information available Integrated, polyblastic, terminal to mostly intercalary, sympodial and cicatrized with thickened scars. MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 Conidia 6-11(-12) x 2.5-3.5(-4.0) um. Solitary, smooth, cylindrical to subpyriform and some subclavate, pale brown, most medially 1-spetate, rarely 0 or 2-septate, often constricted at the septum and the colour septum middle brown and the conidia with a round apex and truncate base; with a faintly darkened, unthickened hilum, about 0.5-0.9 um diam. (3-)6.5-8.5(-12) x (1.5-)2-2.5(- 3) um Solitary, smooth, cylindrical to pyriform, rounded at the apex and truncate at the base, pale brown, 1(-2)-septate, with a faintly darkened, unthickened hilum, about 0.5 um diam. 20-25 x 6-7 um. Straight, fusiform or obclavate, 1 septate, hyaline or subhyaline, verruculose 20-25 x 6-7 um. Cylindrical, rounded at the apex and conico- truncate at the base, or fusiform, 1-3 septate, pale brown, smooth (4-)6-7(-9) x 2-3 um. Solitary, ellipsoidal to ovoid, 0-1(-2)-septate, often constricted at the septa, with a round apex and truncate base, pale brown, smooth, thin-walled; hilum prominent, slightly darkened, unthickened, about 0.5 um diam. 6-12 x 3-5 um. Straight, cylindrical, rounded at the apex, conico-truncate at the base or ellipsoidal, 1-2 septate, smooth, pale brown. (5-)6.5-11(-13.5) x 2.5-4(-4.6) um. Solitary, cylindrical to pyriform, rounded at apex, truncate at base, lower cell longer and wider than upper one, with a prominent scar, 0.8-1.9 um wide, pale brown, without or one median septate, constricted at the septa, smooth-walled. 4.5-18.5 x 5.5 um. Holoblastic, dry, simple, not catenate, finely verruculose, obovoidal, base obconicotruncate, apex rounded, 0-3-septate, pale brown. 6-13.5 x 3-4 um. Elliptical or pyriform, not constricted at septa, 1-3- septate, olivaceous or fuscous. 4-6.5 x 2-3.5 um. Ellipsoid, smooth, Subhyaline or pale brown 7.3-19.5 x 2.6-7.5 um. Holoblastic, dry, simple, non-catenate, finely verruculose, sometimes constricted at the septa, hilum thickened with truncate base and rounded apex, 0-2(-3) septate subhyaline or pale brown Sequence data Present Present Absent Absent Present Absent Present Absent Absent Absent Absent Reference Chandrasiri et al. (2021) Arzanlou et al. (2007) Ellis (1976) Ellis (1976) Arzanlou et al. (2007) Ellis (1976) This study Kharwar and Singh (2004) Dingley (1972) Pan and Zhang (2009) Kharwar and Singh (2004) 210 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Veronaea species V. hedychii V. hippocrateae V. japonica V. latispora V. oblongispora V. polyconidia V. queenslandica V. smilacis V. tectonae V. thylacospermi Conidiophore Conidiogenous cells Conidia Sequence data Reference 19.5-86.5 x 2.5-3.5 um. 10-40 x 1.5-2.5 um. Terminal, 5-10 x 2-4 um. Ellipsoid, Absent Soares and Sometimes with an inflated base | holoblastic, sometimes cylindrical or obovoid, base Barreto (2008) up to 7 um diam., straight to intercalary, densely and minute obconic truncate, apex rounded, slightly curved, unbranched, 1-6 | cicatrized, scars thickened, 1 um__| 0-1 septate, sometimes slightly septate, golden brown to brown, wide. constricted at the septum, paler towards the apices, smooth. subhyaline to pale brown, smooth, hila thickened, somewhat darkened, 1 um diam. 27-123 x 2-4 um. Straight to No information available 4-18.5 x 1.5-3.5 ym, Solitary, Absent Kharwar and slightly curved, unbranched, paler non-catenate, smooth-walled, Singh (2004) towards the apices, 1—6 septate, obconico base, apex rounded, smooth, brown to dark brown. 0-1-septate, cylindric to obclavate, slightly curved, subhyaline to pale brown Up to 65 x 2-3 um. Slightly Up to 15 um long. Terminal, (6-)7—8(-10) x 2-2.5(-4) um. Present Arzanlou et al. differentiated from aerial occasionally intercalary, variable | Solitary, pale brown, smooth, (2007) vegetative hyphae, lateral, or in length, pale brown, cylindrical thin-walled, ellipsoidal to ovoid, terminal, often wider than the to clavate, with hardly prominent | (0—)1-septate, often constricted at supporting hypha, unbranched denticles; scars flat, slightly the septum, with a round apex and or occasionally branched, pale pigmented, not thickened, about | truncate base; hilum unthickened brown, thin-walled, smooth, with 0.5 um diam. but slightly darkened, about 1 um 1-3 additional septa. diam. Up to 90 x 1.5-2.5 um. Straight or | No information available 7.5-9.5 x 3-5 Broadly obovoid, Absent Pan and Zhang slightly curved, septate, smooth, smooth, aseptate, subhyaline to (2009) with numerous minute scars at pale brown. the upper parts, brown. Up to 320 x 3-5 um. Solitary or No information available 7-8 x 4-5 um. Oblong, smooth, Absent Morgan-Jones sometimes in fascicles, smooth, aseptate, rather thick-walled, (1982) bulbous towards the base, pale obtuse at the apex, subhyaline to brown to brown. pale brown. 1125-1515 x 45-70 um. Straight | Polyblastic, terminal, cylindrical, 11-16 x 3-5 um. Solitary and Present Su et al. (2023) or slightly flexuous, unbranched, rachis with crowded, flat to smooth, cylindrical to pyriform, solitary, cylindrical, rough-walled, | slightly prominent, pale brown, rounded at apex and truncate at thick-walled, brown to dark brown. | fertile parts subhyaline. base, pale brown, 1—3-septate (mostly 3), often constricted and medium brown at septa. No information available No information available No information available Absent Matsushima (1989); Index Fungorum (2025) 70-165 (-90-125) x 3.5-5.5 um. | No information available 21.5-32.5 x 2-5 um. Solitary, Absent Singh et al. Solitary, thick-walled, smooth, up cylindrical to slightly obclavate, (1981) to 12-septa, brown. 1-12 septate, smooth-walled, subhyaline to pale olivaceous. Up to 200 x 3.6 um. Solitary No information available 11 x 3.75 um. Solitary, simple, Absent Kamal (1980) smooth, up to 10-septa, pale smooth, usually 1-septa, brown. prominent scars at the base. 35-100 x 3-4 um. Simple, Conidiogenous loci 0.3 um high, 12-14 x 3-4 um. Two-celled, Absent Chlebicki smooth, brown, 1-3(-8) septate, | 0.5 um wide. lower cell longer and wider (2009) basal cell inflated 6-10 um wide, than upper one, wall slightly fertile part taller than basal part, verruculose, hyaline to pale brown. forming slightly flexuous rachis with scattered, hyaline and small, apically pointed denticle-like. Atrokylindriopsis was established by Ma et al. (2015) to accommodate the single species At. setulosa. Atrokylindriopsis setulosa was reported as saprobic on dead branches of an unidentified broadleaf tree and was placed in Cha- etothyriales without a family assigned, based on a BLASTn search using ITS and LSU sequence data from NCBI and morphological comparisons (Ma et al. 2015). Wijayawardene et al. (2020) considered this genus to be in the Chae- tothyriales, but later Quan et al. (2020) assigned it to the Herpotrichiellaceae. Morphologically, the pigmented, septate, setulate conidia are a unique feature of Atrokylindriopsis, distinguishing it from other members of Herpotrichiellace- ae. In the present phylogenetic tree (Fig. 1), the new species, At. racemosos- pora, is closely related to At. setulosa. Atrokylindriopsis racemosospora exhib- its morphological differences from At. setulosa, as mentioned above. Based on morphology, At. racemosospora might represent a different genus from Atrokylindriopsis. The holotype (HMAS245592) and dried culture specimens MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 511 Chanokned Senwanna et al.: New endolichenic fungi from Thailand (HSAUP H4560) of At. setulosa were not reexamined, as they had been lost. Consequently, it was not possible to reassess the morphology. Due to the sim- ilarity in the ITS and LSU sequences between At. racemosospora and At. set- ulosa, along with the unavailability of protein-coding gene sequence data of At. setulosa, it was deemed insufficient evidence to justify separating them into different genera. Since the molecular data were unclear in resolving the rela- tionships between At. setulosa and our specimens, we tentatively classified our strains as a new species within Atrokylindriopsis. Additionally, the examina- tion of newly collected specimens, morphology, and molecular data regarding protein-coding genes of At. setulosa requires further study. It is worth noting that At. racemosospora is similar to Aciculomyces and Petriomyces in having sympodial conidiogenous cells that produce conidia from short denticles and obovoid, aseptate conidia (Thitla et al. 2023; Torres-Garcia et al. 2023). Based on morphological characters and multi-gene phylogenetic analysis, Aciculomy- ces was established by Torres-Garcia et al. (2023) to accommodate the single species Ac. restrictus, collected from fluvial sediments, while Petriomyces, typi- fied by P. obovoidisporus, was isolated from sandstone in a natural forest (Thit- la et al. 2023). Moreover, At. racemosospora can be distinguished from these genera by the abundance of sporulation, inconspicuous conidial scars, and the phylogenetic analysis, which also segregates these genera. Phialophora is known as a group of black yeasts and their relatives, which have been reported as pathogens in humans and animals, as well as saprobic and endophytic in plant materials and soil (Schol-Schwarz 1970; Untereiner et al. 2008; Liu et al. 2013; Li et al. 2017). The genus is typified by P verrucosa and characterized by pigmented hyphae, solitary or aggregated conidiophores, cylindrical to flask-shaped phialides bearing conspicuous, darkly pigmented collarettes, and ovoid to cylindrical, aseptate conidia (Medlar 1915; Schol- Schwarz 1970; Gams 2000; Untereiner et al. 2008). Currently, 38 epithets are listed for Phialophora in Index Fungorum (2025). Morphology alone is insuffi- cient for species identification within Phialophora; moreover, molecular data have demonstrated significant genetic variation within the species complex, now referred to as the P verrucosa species complex (Li et al. 2017). Several species have been transferred to other genera, such as Chloridium, Cyphello- phora, Entimomentora, Hyphodiscus, Lasiosphaeris, and Rhopalophora, based on morpho-molecular analyses (Bogale et al. 2010; Réblova et al. 2011, 2013, 2017; Johnston et al. 2014; Untereiner et al. 2019). Although many sequences of Phialophora are available in GenBank, in our pre-analyses, P asteris, P asteris f. sp. helianthi, P. atrovirens, P. avicenniae, P. cinerescens, P. cyclaminis, P. dan- coi, P foetens, P. intermedia, P japonica, and P mustea were grouped outside of the clade comprising members of Herpotrichiellaceae. Therefore, we excluded these sequences from our analysis. Our molecular phylogenetic tree indicated that our three strains formed a separate clade within the P. chinensis clade, with representative strains of P chinensis separated into two subclades, supported by 86% ML and 1 PP statistical support (Fig. 1). Sequence comparisons of ITS, LSU, SSU, and tub2 genes between our strains and representative strains from two sub-clades indicate that our strains are congeneric. The conidial size of our strains aligns well with P. chinensis; however, the dematiaceous hyphomycetes of our strains cannot be directly compared with the type, as Li et al. (2017) did not provide a detailed description of conidiophores, conidiogenous cells, MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 912 Chanokned Senwanna et al.: New endolichenic fungi from Thailand collarettes, and chlamydospores. According to Li et al. (2017), P chinensis has been isolated from human patients, leaves, soil, wheat straw, and wood, which may account for morphological variation within the species. Based on the available data, we identified our strains as P chinensis, which were collected as endolichenic fungi associated with foliose lichens in Thailand. Phialophora identification was initially based on morphology and molecular phylogeny (Yan et al. 1995; de Hoog et al. 1999; Untereiner et al. 2008). Although similar co- nidia can be observed among Phialophora species, genetic variation in the ITS, LSU, and tub2 gene regions can be used to distinguish species. However, since most available sequences of P. chinensis contain only three gene regions and excluded species contain only the ITS gene region, species delineation based on phylogeny remains uncertain. Therefore, more informative genes (i.e., act, tef1, and rpb2) and additional collections may help clarify species placement and reveal the possibility of a species complex. Veronaea was introduced by Ciferri and Montemartini (1957) and is typified by V. botryose. There are 20 epithets recorded in Index Fungorum (2025), but only five species have been identified with DNA sequence data. Based on multi- gene analyses and morphological comparisons, V. constricta (CBS 572.90, type strain) has been considered a synonym of V. botryosa (Su et al. 2023; Yang et al. 2023). In our phylogenetic tree, the Veronaea species clusters as a monophylet- ic clade (Fig. 1), maintaining a stable position within Herpotrichiellaceae, which concurs with the findings of Su et al. (2023) and Yang et al. (2023). Veronaea endolichena is introduced based on phylogenetic analysis and morphological data. A morphological comparison between V. endolichena and other Veronaea species is shown in Table 2. Veronaea endolichena is similar to V. caricis, V. com- pacta, V. hedychii, V. hippocrateae, V. japonica, V. tectonae, and V. thylacosper- mi in having 0-1-septate conidia. However, the conidia of V. compacta (4-9 x 2.5-4 um), V. hedychii (5-10 x 2-4 um), V. japonica (6-10 x 2-4 ym) and V. tec- tonae (11 x 3.75 um) are shorter than those of V. endolichena (5-13.5 x 2.5—4.6 um) (Ellis 1976; Kamal 1980; Kharwar and Singh 2004; Arzanlou et al. 2007; Soares and Barreto 2008; Chlebicki 2009). Veronaea caricis (20-25 x 6-7 um) and V. hippocrateae (4—18.5 x 1.5-3.5 um) have longer conidia than V. endoli- chena (Ellis 1976; Kharwar and Singh 2004). Veronaea endolichena (5-13.5 x 2.5-4.6 um) is closely related to V. thylacospermi (12-14 x 3-4 um); however, V. endolichena has longer conidiophores (23-264 x 2.3-4.9 um vs 35-100 x 3-4 um) and cylindrical to pyriform, smooth-walled conidia (Chlebicki 2009). Based on both morphological characteristics and multigene analyses, a novel species is introduced. Phylogenetically, FE. brunnea (CBS 587.66) also clustered within the Veronaea clade (Fig. 1), which concurred with the results conducted by Costa et al. (2023) and Torres-Garcia et al. (2023). Exophiala brunnea differs from Veronaea in its conidiophores and phialides (Papendorf 1969; de Hoog et al. 2011); therefore, it was maintained as a separate species pending further investigation. Most species of Veronaea are known to be saprobes on various hosts worldwide (Arzanlou et al. 2007; Soto et al. 2017; Chandrasiri et al. 2021; Su et al. 2023; Yang et al. 2023). Since Veronaea exhibits overlapping charac- teristics, species delimitation based solely on morphology is challenging (Su et al. 2023); therefore, additional sampling is necessary for further investigation. Our phylogenetic analyses indicated that Capronia, Cladophialophora, Ex- ophiala, Fonsecaea, and Rhinocladiella are polyphyletic within the family MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 913 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Herpotrichiellaceae, which is concurred with Teixeira et al. (2017), Thitla et al. (2023), and Torres-Garcia et al. (2023). Whereas Aciculomyces, Aculeata, Atroky- lindriopsis, Marinophialophora, Melanoctona, Petriomyces, Phaeoannellomyces, Phialophora, Thysanorea, Uncispora, Valentiella, and Veronaea represent a mono- phyletic group. Considering the asexual morph resemblance together with mo- lecular support, Hernandez-Restrepo et al. (2020) treat most Minimelanolocus species under Thysanorea. Although several species of Minimelanolocus have been recently reported based on morphological characteristics, these species lack sequence data, including the type species (Costa et al. 2018; Barreto et al. 2024). Pleomelogramma is also considered a doubtful genus in Herpotrichiel- laceae, pending further studies due to the lack of sequence data for the type species. (Quan et al. 2020). Based on molecular analyses with distinctive eco- logical trends, de Hoog et al. (2011) defined the members of this family into six clades: bantiana-, carrionii-, salmonis-, europaea-, dermatitidis-, and jeanselmei- clades. With morphology and multi-gene phylogeny analyses, the species in the europaea clade were assigned as members of Cyphellophoraceae (Réblovd et al. 2013). According to Wijayawardene et al. (2022), Brycekendrickomyces, Metulo- cladosporiella, Neosorocybe, and Sorocybe had previously been classified within Herpotrichiellaceae. However, the phylogenetic position of Brycekendrickomyces and Metulocladosporiella was presented in Trichomeriaceae (Quan et al. 2020, 2024). The ITS and LSU phylogenetic analyses by Quan et al. (2024) revealed that Neosorocybe and Sorocybe are separate from the Herpotrichiellaceae clade, forming a sister clade to the Epibryaceae clade in Chaetothyriales. Later, Thitla et al. 2023 treated both genera in Chaetothyriales incertae sedis. Munk (1953) defined the characteristics of anamorphic Herpotrichiellaceae as dematiaceous hyphomycetes (black yeast), typified by proliferating, percurrently conidiogenous cells, conidiogenesis with holoblastic conidia produced from min- ute pegs or denticles, and unicellular conidia often held in chains. It is worth noting that several obligate lichenicolous Herpotrichiellaceae genera, such as Cladophi- alophora, have been found on lichen thalli or apothecia. These species differ from others in forming minute conidiophores in sporodochia, and in some species, two distinct types of conidia are observed (Diederich et al. 2024). A few species, such as Aculeata aquatica, Atrokylindriopsis setulosa, and Melanoctona tectonae, can also be distinguished from other Herpotrichiellaceae species by their conid- iogenesis and conidial morphology (Ma et al. 2015; Tian et al. 2016; Dong et al. 2018). Although ITS and LSU sequence analyses support the placement of these three species in Herpotrichiellaceae, reliable genes (e.g., rpb2, tef7, and tub2) may provide better resolution for distinguishing species within this family. Due to the recent increase in the number of Herpotrichiellaceae members and the apparent polyphyletic nature of their characteristics, the delineation of the family remains inconclusive (Tian et al. 2021; Crous et al. 2023). Currently, most sequence data of Herpotrichiellaceae strains in GenBank are available only for ribosomal genes, including ITS, LSU, and SSU, while data for protein-coding genes remain limited. Although the ITS and LSU can be used for linking sexual and asexual morphs and delimitating species, their classification is often unclear or placed in unsup- ported branches (Untereiner and Naveau 1999; Quan et al. 2020, 2022; Costa et al. 2023; Thitla et al. 2023; Torres-Garcia et al. 2023). Polyphyletic genera (i.e., Capronia, Cladophialophora, Exophiala, Fonsecaea, and Rhinocladiella) and ambig- uous species within the Herpotrichiellaceae family are still waiting for resolution, MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 714 Chanokned Senwanna et al.: New endolichenic fungi from Thailand particularly with the inclusion of newly collected specimens. Likewise, sequences of protein-coding genes (e.g., act, rpb2, tef7, and tub2) from additional collections, as well as the re-sequencing of generic types within this family and polyphyletic strains, are required to clarify their phylogenetic affinity and species delineation. Members of the family Herpotrichiellaceae play various ecological roles, in- cluding endophytic, epiphytic, fungicolous, hypersaprobic, lichenicolous, patho- genic, rock-inhibiting, and saprobic taxa, on different substrates, as listed in Table 3. These fungi not only interact with their hosts in various ways but are also considered opportunistic organisms that inhabit natural environments (Crous et al. 2007; Quan et al. 2024). Members of the family Herpotrichiellaceae are known Table 3. Updated genera in Herpotrichiellaceae with their life mode, habitat, and isolation sources. Genera Aciculomyces Aculeata Atrokylindriopsis Capronia Cladophialophora Exophiala Fonsecaea Marinophialophora Melanoctona Minimelanolocus* Petriomyces Phialophora Phaeoannellomyces Pleomelogramma* Rhinocladiella Thysanorea Uncispora Valentiella Veronaea Life mode Saprobic Saprobic Saprobic and endolichenic Epiphytic, fungicolous and hypersaprobic, lichenicolous, and saprobic Endophytic, fungicolous lichenicolous, pathogenic, rock- inhabiting, and saprobic Endophytic, fungicolous, pathogenic, rock- inhabiting, and saprobic Epiphytic, pathogenic, and saprobic Saprobic Saprobic Saprobic Rock-inhabiting Endolichenic, pathogenic, and saprobic Pathogenic Saprobic Epilithic, epiphytic, lichenicolous, pathogenic, and saprobic Saprobic Saprobic Saprobic Endophytic, epiphytic, pathogenic, and saprobic * Sequence data are not available. Habitat Freshwater Freshwater Terrestrial Terrestrial Terrestrial and freshwater Aquatic, marine, and terrestrial Terrestrial Marine Terrestrial Aquatic and terrestrial Terrestrial Freshwater and terrestrial Terrestrial Terrestrial Terrestrial Aquatic and terrestrial Freshwater and terrestrial Terrestrial Freshwater and terrestrial Isolation source Fluvial sediments Submerged wood Dead branches of an unidentified broadleaf tree and lichen (Parmotrema sp.) Fungi, lichen, litter, plants, soil, water, and wood Fluvial sediments, lichen, plants, rock, sawdust, soil, wood, warm-blooded animal, and water Cold blooded animals, fungi, litter, plants, rock, sediments, soil, various substrates (i.e., contact lens, plastic foil, soap container, etc.), warm- blooded animals, and water Cold blooded animal, insect, lichen, litter, plants, soil, warm-blooded animal, and wood Associated with Halocyphina species on decaying mangrove wood Decaying wood Litter, plants, and submerged wood Rock Cold blooded animal, fluvial sediments, food, lichen, plants, soil, warm-blooded animal, water, and wood Warm-blooded animal Decaying woody plants Honey, insects, plants, soil, warm-blooded animal, and wood Plants and wood Plants and submerged wood Carton runway galleries built by Azteca brevis ants and plants Animal dung, cold-blooded animal, plants, soil, warm blooded animal, and wood MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 References Torres-Garcia et al. (2023) Dong et al. (2018) Ma et al. (2015); this study Miller et al. (1987); Etayo (2002); Untereiner (2000); Untereiner and Naveau (1999); Untereiner et al. (2011); Marincowitz et al. (2008); Teixeira et al. (2017); Phookamsak et al. (2019); Sanchez et al. (2019); Roux et al. (2020); Hollinger and Lendemer (2021); Tian et al. (2021) Teixeira et al. (2017); Quan et al. (2020); Thitla et al. (2023, 2024); Chang et al. (2023); Cometto et al. (2023) de Hoog et al. (2011); Teixeira et al. (2017); Quan et al. (2020); Tian et al. (2021); Thitla et al. (2022); Ide-Pérez et al. (2024) Vicente et al. (2012, 2014); de Azevedo et al. (2015); Teixeira et al. (2017); de Souza Lima et al. (2020); Sousa et al. (2024) Li et al. (2018) Tian et al. (2016) Castafieda Ruiz et al. (2001, 2003); Zhang et al. (2009); Ma et al. (2011); Heredia et al. (2014); Hernandez-Restrepo et al. (2013); Tian et al. (2016) Thitla et al. (2023) Li et al. (2017); Jiang et al. (2017); Song et al. (2021); Tian et al. (2021); Ply et al. (2023); Torres-Garcia et al. (2023); this study McGinnis et al. (1985) Spegazzini (1909) Arzanlou et al. (2007); Hernandez-Restrepo et al. (2016); Teixeira et al. (2017); de Souza Lima et al. (2020); Quan et al. (2020); Roeun et al. (2022); Sousa et al. (2024) Arzanlou et al. (2007); Tian et al. (2016); Dong et al. (2018); Hernandez-Restrepo et al. (2020); Yang et al. (2023) Sinclair and Morgan-Jones (1979); Li et al. (2014); Liu et al. (2018) Nepel et al. (2014); Bezerra et al. (2022) Ellis (1976); Papendorf (1976); Moustafa and Abdul-Wahid (1990); Arzanlou et al. (2007); Soares and Barreto (2008); Chlebicki (2009); Sang et al. (2011) Soto et al. (2017) Coleman et al. (2018); Chandrasiri et al. (2021); Su et al. (2023); Yang et al. (2023); this study 215 Chanokned Senwanna et al.: New endolichenic fungi from Thailand for their biochemical and ecological adaptations that enable survival in extreme environments, often linked to the production of protective compounds such as melanin (de Hoog et al. 2011; Zhan et al. 2011; Coleine and Selbmann 2021; Coelho et al. 2022; Quan et al. 2024; de Leon et al. 2025). For example, Cladophi- alophora exuberans, Exophiala dermatitidis, E. pisciphila, and E. phaeomuriformis produce melanin, which enables them to thrive under high-radiation, high-tem- perature or oligotrophic conditions (Badali et al. 2008; de Hoog et al. 2011; Zhan et al. 2011; Cordero and Casadevall 2017; Benmessaoud et al. 2023; da Silva et al. 2023; de Leon et al. 2025). Although this study employed culture-based tech- niques to isolate and identify endolichenic fungi within Herpotrichiellaceae, many taxa from this family and others likely remain undetected due to their uncultur- able nature, low abundance, or specific growth requirements. Future research incorporating metaomics approaches, including metagenomics and metatran- scriptomics, could provide a more comprehensive view of fungal diversity, func- tional gene expression, and ecological interactions within the lichen holobiont. These tools offer promising avenues to uncover hidden fungal lineages and un- derstand their metabolic roles in lichen symbiosis and environmental adaptation. Acknowledgements The authors are grateful to Hatthakom Mingrujidith and Nopparat Wannathes for their kind advice in naming the new fungal taxa. Liwei Zhou, Nalin N. Wijay- awardene, Rungtiwa Phookamsak, and Ausana Mapook are gratefully acknowl- edged for their valuable suggestions and help. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of Al No use of Al was reported. Funding Nakarin Suwannarach and Chanokned Senwanna expresses appreciation to the CMU Proactive Researcher Program, Chiang Mai University. Author contributions Conceptualization: CS, NS. Data curation: CS, JK, PK, ND, NS. Formal analysis: CS, JK, NS. Funding acquisition: CS,NS. Investigation: CS, JK, NS. Methodology: CS, NS. Project admin- istration: NS. Software: CS, JK. Supervision: JK, NS. Validation: CS, JK, NS. Visualization: CS, JK, NS. Writing — original draft: CS, JK, NS. Writing — review & editing: CS, JK, PK, ND, NS. Author ORCIDs Chanokned Senwanna ® https://orcid.org/0000-0002-1008-4514 Jaturong Kumla ® https://orcid.org/0000-0002-3673-6541 MycoKeys 120: 193-229 (2025), DOI: 10.3897/mycokeys.120.153906 716 Chanokned Senwanna et al.: New endolichenic fungi from Thailand Pratthana Kodchasee ® https://orcid.org/0000-0003-0432-4910 Nutchanan Duangkon © https://orcid.org/0009-0008-5769-0117 Nakarin Suwannarach © https://orcid.org/0000-0002-2653-1913 Data availability All of the data that support the findings of this study are available in the main text. References Agrawal S, Deshmukh SK, Reddy MS, Prasad R, Goel M (2020) Endolichenic fungi: A hid- den source of bioactive metabolites. South African Journal of Botany 134: 163-186. https://doi.org/10.1016/j.sajb.2019.12.008 Ahmed SA, Bonifaz A, Gonzalez GM, Moreno LF, Menezes da Silva N, Vicente VA, Li R, de Hoog S (2021) Chromoblastomycosis Caused by Phialophora—Proven Cases from Mexico. 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