Reappraisal of Immotthia in Dictyosporiaceae, Pleosporales: Introducing Immotthia bambusae sp. nov. and Pseudocyclothyriella clematidis comb. et gen. nov. Based on Morphology and Phylogeny

Immotthia is a poorly known genus, and currently, no DNA sequence data are available to ascertain its proper phylogenetic placement and evolutionary relationships with other bitunicate fungi. To date, there are only two species accepted in the genus. During our ongoing research study of bambusicolous fungi in southwest China and Thailand, a fungus associated with stromata of Hypoxylon sp. was found on dead bamboo culms in Loei Province, Thailand. Preliminary morphological identification revealed that the fungal collection belongs to Immotthia. A novel species, Immotthia bambusae, is introduced herein based on a comparison of morphological characteristics with the type specimen of I. hypoxylon (≡ Amphisphaeria hypoxylon Ellis and Everh.), a synonym of I. atrograna (Cooke and Ellis) M. E. Barr. Phylogenetic analyses of a concatenated ITS, LSU, SSU, and TEF1-α DNA sequence matrix showed that Immotthia belongs to Dictyosporiaceae, Pleosporales. Despite I. bambusae strains constituting a supported subclade, they are nested with the genus Pseudocoleophoma. Pseudocoleophoma clematidis is morphologically different from all other Pseudocoleophoma species, while its conidial characteristics are similar to Cyclothyriella. Multigene phylogenetic analyses showed that P. clematidis formed a clade basal to Immotthia, separated from Pseudocoleophoma with strong statistical support. Therefore, we introduce a monotypic genus, Pseudocyclothyriella Phukhams. and Phookamsak, gen. nov. to accommodate the single species, Pseudocyclothyriella clematidis (Phukhams. and K. D. Hyde) Phukhams. and Phookamsak, comb. nov. Detailed descriptions, color micrographs, and phylogenetic trees to show the placement of the new taxa are provided. In addition, an updated taxonomic treatment of the genera Immotthia and Pseudocyclothyriella is also provided based on the study of the type materials and phylogeny generated from DNA sequence data.

Immotthia is a poorly known genus, and currently, no DNA sequence data are available to ascertain its proper phylogenetic placement and evolutionary relationships with other bitunicate fungi. To date, there are only two species accepted in the genus. During our ongoing research study of bambusicolous fungi in southwest China and Thailand, a fungus associated with stromata of Hypoxylon sp. was found on dead bamboo culms in Loei Province, Thailand. Preliminary morphological identification revealed that the fungal collection belongs to Immotthia. A novel species, Immotthia bambusae, is introduced herein based on a comparison of morphological characteristics with the type specimen of I. hypoxylon (≡ Amphisphaeria hypoxylon Ellis and Everh.), a synonym of I. atrograna (Cooke and Ellis) M. E. Barr. Phylogenetic analyses of a concatenated ITS, LSU, SSU, and TEF1-α DNA sequence matrix showed that Immotthia belongs to Dictyosporiaceae, Pleosporales. Despite I. bambusae strains constituting a supported subclade, they are nested with the genus Pseudocoleophoma. Pseudocoleophoma clematidis is morphologically different from all other Pseudocoleophoma species, while its conidial characteristics are similar to Cyclothyriella. Multigene phylogenetic analyses showed that P. clematidis formed a clade basal to Immotthia, separated from Pseudocoleophoma with strong statistical support. Therefore, we introduce a monotypic genus, Pseudocyclothyriella Phukhams. and Phookamsak, gen. nov. to

INTRODUCTION
Immotthia was introduced by Barr (1987a) with I. hypoxylon (Ellis and Everh.) M. E. Barr (≡ Amphisphaeria hypoxylon Ellis and Everh.) as the type species. Through examinations of the type material of I. hypoxylon and Australian collections of I. atrograna (Cooke and Ellis) M. E. Barr (≡ Sphaeria atrograna Cooke and Ellis), Jaklitsch et al. (2002) concluded that these two taxa are conspecific. To date, two species are accepted in this genus, viz. I. atrograna and I. atroseptata (Piroz.) M. E. Barr (Species Fungorum, 2020) based on morphology, but no DNA sequence data are available to confirm their phylogenetic placement (Hyde et al., 2017;Doilom et al., 2018).
Most of the Pseudocoleophoma species have been represented by their asexual morphs Tennakoon et al., 2019;Li et al., 2020;Phukhamsakda et al., 2020). Only three species have been reported for both sexual and asexual morphs, viz. P. bauhiniae, P. calamagrostidis, and P. polygonicola Jayasiri et al., 2019). Currently, species of Pseudocoleophoma are only known from Europe (Great Britain, Italy, and Netherlands) and Asia (Taiwan and Thailand), and they were found as saprobes on various hosts and substrates from both terrestrial and freshwater habitats Hyde et al., 2016;Jayasiri et al., 2019;Tennakoon et al., 2019;Li et al., 2020;Phukhamsakda et al., 2020).
In the present study, a fresh collection of Immotthia is examined and compared with other Immotthia species based on morphological characteristics. The new collection is described as a novel species in Immotthia and illustrated. Through DNA sequencing of the fresh material, we also resolved the phylogenetic placement of Immotthia for the first time, based on maximum likelihood and Bayesian inference analyses. In addition, the novel genus Pseudocyclothyriella is also introduced to accommodate Pseudocyclothyriella clematidis comb. nov. (≡ Pseudocoleophoma clematidis) based on morphological distinctiveness and multigene phylogenetic analyses.

Sample Collection, Specimen Examination, and Preservation
Dead bamboo culms were collected from Loei Province, Thailand, in 2020. The specimens were kept in a paper bag and returned to the laboratory for observation and examination. Fungal fruiting bodies on the host substrate were observed with a Motic SMZ 140 series dissecting stereoscope, and a centrum was mounted in sterilized distilled water for morphological examination and captured using a Nikon ECLIPSE Ni compound microscope connected with a Canon EOS 600D digital camera. Tarosoft (R) Image Frame Work version 0.9.7 was used to measure the size of ascomata, peridium, pseudoparaphyses, asci, and ascospores. In addition, holotypic specimens of Immotthia atroseptata [United States, North Carolina, behind N. C. Department of Agriculture, Nursery Inspection Station, 1 mile west of Linville, Avery Co., on apothecia of Pestalopezia rhododendri on the leaves of Rhododendron maximum L.   were also re-examined and illustrated. Adobe Photoshop CS6 software (Adobe Systems Inc., United States) was used to edit and provide the photographic plates based on captured pictures of the fungal structures. Good practices for morphological examinations as outlined by Senanayake et al. (2020b) were followed for the morphological study, while phylogenetic methods as outlined by Dissanayake et al. (2020) were followed for phylogenetic analyses. The holotype is deposited in the herbarium of Cryptogams Kunming Institute of Botany Academia Sinica (KUN-HKAS), Yunnan, China. The isotype was stored in Herbarium Mycologicum Academiae Sinicae (HMAS), Beijing, China. The Facesoffungi and Index Fungorum numbers are registered for the newly described taxa Index Fungorum, 2020). New species are established based on the guidelines provided by Jeewon and Hyde (2016).

Alignment and Phylogenetic Analyses
The newly generated ITS, LSU, SSU and TEF1-α sequences of the new taxon were subjected to the BLASTn search tool 1 for initial verification and search of reference taxa for further analyses. Similarity indices based on BLASTn search showed that five new strains are closely related to Pseudocoleophoma Kaz. Tanaka and K. Hiray (Dictyosporiaceae, Pleosporales). In order to investigate the phylogenetic status of the new taxa, a combined dataset of taxa including members of the Dictyosporiaceae was analyzed based on DNA sequence data available in recent publications (Iturrieta-González et al., 2018;Yang et al., 2018;Hyde et al., 2020a,b;Phukhamsakda et al., 2020). DNA sequences of representative taxa used are shown in Table 1. Individual DNA sequence alignments were initially performed via the online platform MAFFT v. 7.474 2 (Katoh et al., 2019) and were improved manually using BioEdit v. 5.0.6 (Hall, 2001). Preliminary phylogenetic analyses of a concatenated LSU-SSU-TEF1α-RPB2-ITS sequence matrix represented the relationships of Immotthia in Dictyosporiaceae with other families in Pleosporales (Supplementary Figure 3), and a concatenated ITS-LSU-TEF1-α sequence dataset (Supplementary Figure 1) was also analyzed by maximum-likelihood (ML) analysis via the web portal CIPRES Science Gateway v. 3.3 (Miller et al., 2010), with the help of the tool RAxML-HPC v.8 on XSEDE (8.2.12).
ML analysis using the GAMMA model of nucleotide substitution was performed via the web portal CIPRES Science Gateway v.3.3 (Miller et al., 2010), with the help of the tool RAxML-HPC v.8 on XSEDE (8.2.12). The evolutionary model of nucleotide substitution for Bayesian inference (BI) analysis was selected independently for each locus using MrModeltest 2.3 (Nylander, 2008). GTR + I + G is the best fit for ITS and LSU loci under the Akaike information criterion (AIC), while  the HKY + I substitution model is the best fit for SSU and TEF1-α loci. BI analysis was carried out by MrBayes v. 3.2.6 (Ronquist and Huelsenbeck, 2003). Markov chain Monte Carlo sampling (MCMC) was used to decide posterior probabilities (PP) (Rannala and Yang, 1996;Zhaxybayeva and Gogarten, 2002). Six simultaneous Markov chains were run for 1,000,000 generations and trees were sampled every 100th generation. The 0.15 "temperature" value was set in the MCMC heated chain. All sampled topologies beneath the asymptote (20%) were discarded as part of a burn-in procedure, and the remaining 8,000 trees were used for calculating posterior probabilities (PP) in the 50% majority rule consensus tree (when the split frequency was lower than 0.01). Tree topology of BI analysis is represented in Supplementary Figure 2.
Tree topologies generated in this study were visualized on FigTree v. 1.4.0 3 , and Microsoft Office PowerPoint 2016 (Microsoft Inc., United States) was used to edit and redraw the phylogram. New sequences generated from the present study are deposited in GenBank ( Table 1). The final alignment and phylogram were submitted in TreeBASE 4 .
Notes: The phylogenetic affinities of Immotthia have never been investigated in previous studies, and the genus was accommodated in different families by different authors based only on morphological characteristics (Barr, 1987a(Barr, , 2002Jaklitsch et al., 2002;Akulov and Hayova, 2016;Hyde et al., 2017;Doilom et al., 2018). In the present study, Immotthia is phylogenetically close to Pseudocoleophoma in Dictyosporiaceae. The sexual morph of Immotthia differs from Pseudocoleophoma in morphology and habitat. Immotthia forms dense, superficial ascomata on hypostoma, with cylindrical to cylindric-clavate asci, ellipsoidal to fusiform, light brown to reddish brown, asymmetrical ascospores, lacking mucilaginous sheath and usually found as hyperparasites on hypoxylon-like stroma. On the other hand, Pseudocoleophoma forms scattered or in groups, immersed to erumpent ascomata, with cylindrical to clavate asci, fusiform, hyaline ascospores, surrounded by mucilaginous sheath, and mostly found as saprobes in a terrestrial environment Jayasiri et al., 2019;Tennakoon et al., 2019;Li et al., 2020). The asexual morph of Immotthia differs from Pseudocoleophoma in having multiloculate, carbonaceous conidiomata and ellipsoidal, brown, aseptate conidia (Hyde et al., 2017), whereas Pseudocoleophoma has uniloculate conidiomata and cylindrical to subcylindrical or fusiform, hyaline, 0-1septate conidia.
Notes: The type specimens of all Immotthia species were compared; I. bambusae shows the most similarity to I. hypoxylon, a synonym of I. atrograna. However, the novel taxon differs from I. atrograna (= I. hypoxylon) in having a smaller size of ascomata, asci, and ascospores. The lower cell of ascospores of I. bambusae is longer and narrower than the upper cell, while in I. atrograna (= I. hypoxylon), the upper cell of ascospores is larger than the lower cell. Immotthia bambusae was collected on the stromata of Hypoxylon associated with bamboo in Thailand (tropical region), whereas the type specimen of I. hypoxylon was collected from Hypoxylon truncatum colonizing dead canes of Rosa in Louisiana, United States (subtropical region). The other different features between I. bambusae and the other Immotthia species are provided in Table 2. Therefore, I. bambusae is introduced as a new species in this study based on both morphology and multigene phylogeny.
In this study, we sequenced five different fruiting bodies and DNA sequence similarity (ITS regions) revealed that they have identical nucleotides (100% similarity). Even our phylogeny depicts a close relationship (100% support) among these five strains. This ensures the correctness of the new generated sequences from the direct DNA extraction of fruiting bodies.

Index Fungorum Number: IF 557441
Facesoffungi Number: FoF 09539 Etymology: The generic epithet "Pseudocyclothyriella" refers to the resemblance of conidial morphology of the new genus to the genus Cyclothyriella.
Life Mode and Known Distribution: Pseudocyclothyriella is reported as a saprobe on Clematis vitalba (Ranunculaceae). The genus is presently known from Italy (Phukhamsakda et al., 2020).
Notes: Based on morphological distinctiveness and multigene phylogenetic analyses, a monotypic genus Pseudocyclothyriella is introduced herein to accommodate a single coelomycetous species, P. clematidis which was previously described as Pseudocoleophoma clematidis by Phukhamsakda et al. (2020). Pseudocyclothyriella formed an independent clade basal to Immotthia and Pseudocoleophoma with strong statistical supports (92% ML, 0.98 PP). Pseudocyclothyriella is similar to Cyclothyriella in having cylindrical, oblong to ellipsoid, aseptate, hyaline to pigmented conidia (Jaklitsch and Voglmayr, 2016). However, Pseudocyclothyriella can be distinguished from Cyclothyriella based on the conidiomatal characteristics and phylogenetic evidence. Pseudocyclothyriella is characterized by solitary to gregarious, immersed to erumpent, black, shiny, subglobose to subconical conidiomata, with oval, papilla, ostiolar canal, and pycnidial wall composed of thick-walled, scleroplectenchymatous cells. On the other hand, Cyclothyriella has black, more or less globose pycnidia, clustered in valsoid configuration, with brightly colored, disc-like ostiole, and pycnidial wall composed of pseudoparenchymatous cells (Jaklitsch and Voglmayr, 2016). Cyclothyriella belongs to its own family Cyclothyriellaceae, whereas Pseudocyclothyriella belongs to Dictyosporiaceae.
Pseudocoleophoma differs from Pseudocyclothyriella in pycnidial wall composed of thin-walled, brown to dark brown pseudoparenchymatous cells and oblong, cylindrical to subcylindrical, or rod-shaped, 0-1-septate, smooth-walled conidia Jayasiri et al., 2019;Tennakoon et al., 2019;Li et al., 2020). Pseudocyclothyriella is morphologically similar to a presumably coniothyrium-like asexual morph of Immotthia in having black, carbonaceous pycnidia, and pigmented, aseptate, smooth-walled conidia (Hyde et al., 2017). However, Immotthia forms pycnidia in groups, or multiloculate conidiostromata associated with the ascomata of Immotthia on natural host substrates and has ellipsoidal conidia as well as having hyperparasitic life mode (Hyde et al., 2017), while Pseudocyclothyriella forms solitary to gregarious, uniloculate pycidia and has oval to oblong conidia as well as having saprobic life mode (Phukhamsakda et al., 2020). The sexual morph of Pseudocyclothyriella is undetermined; thus, the sexual morphologies of Pseudocyclothyriella and Immotthia could not be compared.
Holotype Details: Italy, Arezzo Province, Badia Tega-Ortignano Raggiolo, on dead aerial branch of Clematis vitalba,  Ecology and Known Distribution: As a saprobe, Pseudocyclothyriella clematidis has contributions in the cycle of the material. To date, the species is just reported in Italy (Phukhamsakda et al., 2020).
Notes: Pseudocyclothyriella clematidis was previously treated in Pseudocoleophoma based on phylogenetic evidence (Phukhamsakda et al., 2020). Pseudocyclothyriella clematidis differs from the other Pseudocoleophoma species in having yellowish brown, oval to oblong, aseptate conidia (Phukhamsakda et al., 2020). A morphological comparison of P. clematidis with other Pseudocoleophoma species is provided in Table 3. The two strains of Pseudocyclothyriella clematidis (MFLU 16-0280 and MFLUCC 17-2177A) formed a strongly supported clade in this study. Phukhamsakda et al. (2020) introduced Pseudocoleophoma clematidis based on phylogenetic evidence and reported the species was poorly supported in a subclade in between the main clade of Pseudocoleophoma and P. typhicola (MFLUCC 16-0123). However, in our multigene phylogeny, we recovered the same taxon as basal to Pseudoconiothyrium broussonetiae. Crous et al. (2019) mentioned that the closest hits of P. broussonetiae using the ITS sequence had highest similarity to Pseudocoleophoma typhicola (MFLUCC 16-0123). However, P. typhicola is morphologically different from P. broussonetiae, but is typical of Pseudocoleophoma. We rechecked the BLASTn search result based on ITS and LSU sequences of P. typhicola (MFLUCC 16-0123) available in GenBank and noted that the DNA sequences from the ITS regions of P. typhicola are similar to the endophytic Pleosporales sp. isolate MBD_4078 (MK595603) and the uncultured fungus clone ITS_S7_clon2 (HQ873356) with 90.50% similarities and matches with P. broussonetiae strain CBS 145036 (NR_163377) with 90.23% similarity which is far away from Pseudocoleophoma. On the other hand, the BLASTn search result based on LSU sequence showed that the species is closely related with Pseudocoleophoma, and hence, these may be erroneous. In this study, we therefore excluded the ITS sequence of P. typhicola from our aligned sequence dataset and the phylogenetic results showed that P. typhicola clusters within Pseudocoleophoma (Figure 4). However, P. typhicola needs to be resequenced as well as obtaining more reliable genes (only ITS and LSU are available in GenBank) for a better phylogenetic resolution is needed.

Phylogenetic Analyses
The Preliminary phylogenetic analysis based on a concatenated LSU-SSU-TEF1-α-RPB2-ITS sequence matrix of representative families in Pleosporales depicts that Immotthia belongs to the family Dictyosporiaceae (Supplementary Figure 3). The phylograms from ML and BI analyses (Figure 4 and Supplementary Figure 2) of a concatenated ITS-LSU-SSU-TEF1-α sequence matrix were similar in overall topologies and were also similar to the tree topology of a concatenated ITS-LSU-TEF1-α sequence dataset (Supplementary Figure 1). Our five new strains of I. bambusae constitute a strongly supported independent subclade basal to Pseudocoleophoma (62% ML, 0.95 PP). Two strains of Pseudocyclothyriella clematidis (≡ Pseudocoleophoma clematidis, MFLU 16-0280 and MFLUCC 17-2177A) formed an independent subclade basal to Immotthia with strong statistical supports (92% ML, 0.98 PP). Based on current phylogenetic status and morphological distinctiveness compared with the other Pseudocoleophoma species, Pseudocoleophoma clematidis is transferred to the novel genus Pseudocyclothyriella as P. clematidis.
In this study, we collected a fungus associated with Hypoxylon stromata on bamboo from northeastern Thailand. Based on morphological examination comparable with the type specimens, the species is identified as a typical Immotthia but largely different to warrant the establishment of a new species. Thus, we introduce a novel species I. bambusae and this is also the first report of Immotthia associated with Hypoxylon stromata on bamboo in Thailand. Furthermore, it is the first time that DNA sequence data of Immotthia are obtained and its phylogenetic affinity within the Dictyosporiaceae was investigated. In addition, a novel genus Pseudocyclothyriella is introduced as a monotypic genus to accommodate P. clematis during the phylogenetic investigation of Immotthia.
Multigene phylogenetic analyses showed that Immotthia formed a well-resolved clade within Dictyosporiaceae in all analyses (Figure 4 and Supplementary Figures 1-3). The genus clustered with Pseudocoleophoma in all analyses with significant support in BI analysis (0.95 PP; Figure 4), but low support in ML analysis (62% ML; Figure 4). However, Immotthia is also morphologically different from Pseudocoleophoma (see notes under generic description).
Immotthia is widely distributed from tropical to temperate regions including Austria, Belgium, China, France, Lithuania, Norway, Poland, Puerto Rico, Russia, Sweden, Switzerland, Ukraine, United States, and Venezuela (Pirozynski, 1973;Jaklitsch et al., 2002;Akulov and Hayova, 2016;Hyde et al., 2017;Doilom et al., 2018;Farr and Rossman, 2020). Immotthia does not seem to exhibit a hyperparasitic lifestyle on Hypoxylon, but species of this genus were also reported as saprobes on various decayed hardwoods (Jaklitsch et al., 2002). Immotthia bambusae did not germinate on potato dextrose agar (PDA) medium, suggesting that the species has possibly an obligate parasitic life mode, which is in agreement with Jaklitsch et al. (2002). Jaklitsch et al. (2002) treated the type species of Immotthia, I. hypoxylon, as a synonym of I. atrograna after they examined the type materials of these two species. They found that the basionym of both I. hypoxylon and I. atrograna shared similar size range of ascomata, asci, and ascospores and it does not show any convincing difference on the ascomata, although these two species occurred on different hosts and habitats (Jaklitsch et al., 2002). Hyde et al. (2017) re-examined the type material of I. hypoxylon and compared it with other collections from North America and reported similar morphology, but the latter have larger ascomata and shorter asci than the type (Hyde et al., 2017). A comparison of the type examination between I. hypoxylon (Hyde et al., 2017) and I. atrograna (Jaklitsch et al., 2002) shows that I. atrograna has a larger ascomata, smaller asci, and overlapped size range of ascospores. However, these two species still lack DNA sequence data, and hence, their taxonomic status with regards to whether they are conspecific warrants further investigation. Herein, we follow the treatment of Jaklitsch et al. (2002) until the epitypes of these two species are designated and their taxonomy is revisited.
Dictyosporiaceae comprises 17 genera, including Immotthia and Pseudocyclothyriella. Asexual morph of Immotthia is recognized as chaetophoma-, coniothyrium-, microsphaeropsis-, or pyrenochaeta-like (Jaklitsch et al., 2002;Akulov and Hayova, 2016;Hyde et al., 2017;Doilom et al., 2018), which is also similar to coelomycetous asexual morph of Roussoella (Hyde et al., 2017). Hyde et al. (2017) tentatively placed Immotthia in Roussoellaceae based on the morphological similarity to Roussoella. However, Immotthia is phylogenetically distinct from Coniothyrium (Coniothyriaceae) and Roussoella (Roussoellaceae). The genus is transferred from Roussoellaceae to Dictyosporiaceae in this study. The coelomycetous asexual morph of Immotthia was found on natural substrates associated with the sexual morph. However, the link between sexual and asexual morph of Immotthia has not yet been proven, although Jaklitsch et al. (2002) attempted to elucidate the connection based on cultural experiments. The ascospores of Immotthia do not germinate on artificial media due to its obligate parasitic life mode. The connection of the sexualasexual morph needs to be confirmed based on DNA sequence data obtained from direct DNA extraction of fruiting bodies as well as on culture-based studies.
Asexual morphs of most genera in Dictyosporiaceae are hyphomycetes, except for Immotthia, Pseudocoleophoma, Pseudocyclothyriella, and Pseudoconiothyrium which are coelomycetous asexual morphs Crous et al., 2019;Hongsanan et al., 2020). Based on morphological characteristics, Immotthia is most similar to Pseudoconiothyrium in having hyaline, smooth, doliiform to ampulliform, phialidic conidiogenous cells and ellipsoidal, brown conidia (Crous et al., 2019). However, the genus can be distinguished from Pseudoconiothyrium by multiloculate, smaller conidiomata, smooth-walled conidia, whereas Pseudoconiothyrium has uniloculate, larger conidiomata, verrucose conidia (Crous et al., 2019). Two strains of Pseudocoleophoma clematidis formed a stable clade basal to Immotthia and separated from the main clade of Pseudocoleophoma in all analyses (Figure 4 and Supplementary Figures 1-3). We, therefore, re-examined the holotype specimen of P. clematidis and found that the species was morphologically different from the other Pseudocoleophoma (see Table 3). Thus, Pseudocyclothyriella is introduced herein based on the evidence from both morphology and phylogeny. Pseudocyclothyriella is also similar to Pseudoconiothyrium but differs in having thick-walled, scleroplectenchymatous cells of pycnidial wall. Based on our current phylogenetic results, Immotthia and Pseudocyclothyriella are clearly distinct from Pseudoconiothyrium.
Only four genera in Dictyosporiaceae have been reported for their sexual morphs, viz. Dictyosporium Corda, Gregarithecium Kaz. Tanaka and K. Hiray., Immotthia, and Pseudocoleophoma. Immotthia is morphologically different from these three genera in having ellipsoidal to fusiform, light brown to reddish brown, asymmetrical ascospores, lacks mucilaginous sheath, and exhibits a hyperparasitic life mode (Jaklitsch et al., 2002;Hyde et al., 2017). On the other hand, the other three genera have hyaline, fusiform to narrowly fusiform ascospores, with or without mucilaginous sheath and have been reported as saprobes in terrestrial and freshwater habitats Boonmee et al., 2016). The familial descriptions for Dictyosporiaceae have previously been restricted to hyphomycetous asexual morphs (Boonmee et al., 2016;Hongsanan et al., 2020). We recommend that the descriptions and illustrations of Dictyosporiaceae should include both coelomycetous and hyphomycetous asexual morphs.

DATA AVAILABILITY STATEMENT
The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found in the article/Supplementary Material.

AUTHOR CONTRIBUTIONS
H-BJ and RP: conceptualization, data curation, and formal analysis. SL, SCK, and RP: funding acquisition. H-BJ, RP, CP, and MD: investigation, methodology, and writing-original draft. RP and NS: project administration. SL, SCK, RJ, and PK: supervision. NS, PK, RJ, SL, and SCK: writing-review and editing. All authors contributed to the article and approved the submitted version.