Research Article
Research Article
Lignicolous freshwater ascomycetes from Thailand: Introducing Dematipyriforma muriformis sp. nov., one new combination and two new records in Pleurotheciaceae
expand article infoDan-Feng Bao§|, Darbhe J. Bhat, Saranyaphat Boonmee§, Kevin D. Hyde§, Zong-Long Luo, Sarunya Nalumpang|
‡ Dali University, Dali, China
§ Mae Fah Luang University, Chiang Rai, Thailand
| Chiang Mai University, Chiang Mai, Thailand
¶ Unaffiliated, Goa Velha, India
Open Access


During the study of lignicolous freshwater fungi from Thailand, three pleurotheciaceous species were collected from freshwater habitats in Thailand. Two were identified as Pleurothecium aquaticum and Rhexoacrodictys fimicola, and the third is a new species Dematipyriforma muriformis sp. nov.. Rhexoacrodictys is accepted in Pleurotheciaceae based on phylogenetic analysis. Rhexoacrodictys nigrospora is transferred to Dematipyriforma based on phylogenetic analysis and morphological characters. Pleurothecium aquaticum and Rhexoacrodictys fimicola are reported from Thailand for the first time.


1 new combination, 1 new taxon, freshwater fungi, phylogeny, Pleurotheciales, taxonomy


Pleurotheciales was introduced by Réblová et al. (2016) to accommodate a single family Pleurotheciaceae. The order was originally placed in Hypocreomycetidae by Réblová et al. (2016). Hongsanan et al. (2017) showed that Pleurotheciales clustered with Conioscyphales, Fuscosporellales and Savoryellales in a monophyletic clade within Sordariomycetes. Hence, they transferred Pleurotheciales to a newly introduced subclass Savoryellomycetidae based on phylogenetic analysis and the placement has been confirmed and accepted by Dayarathne et al. (2019) and Hyde et al. (2020a).

Pleurotheciaceae was introduced by Réblová et al. (2016) with Pleurothecium Höhn. as the type genus. Currently, Adelosphaeria, Anapleurothecium, Coleodictyospora, Dematipyriforma, Helicoascotaiwania, Melanotrigonum, Neomonodictys, Phaeoisaria, Phragmocephala, Pleurotheciella, Pleurothecium, Saprodesmium, and Sterigmatobotrys are accepted in this family (Hyde et al. 2020a; Wijayawardene et al. 2020; Dong et al. 2021). The sexual morphs of Pleurotheciaceae share dark, papillate, perithecial, astromatic, immersed to superficial ascomata, unitunicate asci with a distinct non-amyloid apical annulus, and fusiform to ellipsoidal, septate, hyaline ascospores (Réblová et al. 2016; Luo et al. 2018a; Hyde et al. 2020a). The asexual morphs of Pleurotheciaceae are diverse in morphology, comprising acrodictys-like (Monotosporella), (Hyde and Yanna 2002; Sadowski et al. 2012), helicoön-like (Helicoascotaiwania, Dayarathne et al. 2019; Réblová et al. 2020), monodictys-like (Neomonodictys, Hyde et al. 2020b) and dactylaria-like taxa (Pleurotheciella, Phaeoisaria and Pleurothecium, Réblová et al. 2016; Luo et al. 2018a). Species in Pleurotheciaceae are cosmopolitan with a worldwide distribution and have been reported from both aquatic and terrestrial habitats (Réblová et al. 2016, 2020; Hernandez-Restrepo et al. 2017; Luo et al. 2018a, 2019; Hyde et al. 2020a, b).

In this study, three new collections are placed in Dematipyriforma, Rhexoacrodictys and Pleurothecium respectively. The monotypic genus Dematipyriforma was introduced to accommodate an endophytic species, D. aquilaria which was collected from wood of Aquilaria crassna (Sun et al. 2017). Dematipyriforma was originally placed in Savoryellales (Sun et al. 2017). However, Dong et al. (2021) showed that Dematipyriforma clustered within Pleurotheciales and sister to Rhexoacrodictys and Saprodesmium. In addition, the morphology of Dematipyriforma is similar to Neomonodictys in Pleurotheciales. Therefore, they transferred Dematipyriforma to Pleurotheciales based on phylogenetic analysis and morphological characteristics. Rhexoacrodictys was introduced by Baker et al. (2002) to accommodate species previously identified as Acorcdictys (i.e., A. erecta, A. fimicola, A. fuliginosa and A. queenslandica) and wherein Rhexoacrodictys erecta was designated as the type. Two additional species R. martini and R. broussonetiae were subsequently added to the genus based on morphological characteristics (Delgado 2009; Xiao et al. 2018). While R. martini and R. queenslandica were transferred to Distoseptispora and Junewangia based on phylogenetic analysis (Xia et al. 2017). Currently, four species are accepted in Rhexoacrodictys. Pleurothecium was established by Höhnel (1919) with P. recurvatum (Morgan) Höhn as type species. Pleurothecium species are characterized by macronematous, mononematous, septate, brown conidiophores, polyblastic, sympodially extended, denticulate conidiogenous cells and solitary, septate, hyaline or pigmented or bicolored conidia (Goos 1969; Matsushima 1975, 1980; Subramanian and Bhat 1989; Matsushima and Matsushima 1996; Cooper 2005; Arzanlou et al. 2007; Wu and Zhang 2009; Réblová et al. 2012; Monteiro et al. 2016; Luo et al. 2018a). Presently, 11 species are accepted in the genus. Most Pleurothecium species are reported as saprobes from freshwater or terrestrial habitats (Wu and Zhang 2009; Réblová et al. 2012; Monteiro et al. 2016; Luo et al. 2018a).

We are currently investigating the diversity of lignicolous freshwater fungi from the Greater Mekong Subregion (Hyde et al. 2016). Thailand is an area of the Greater Mekong Subregion with rich fungal biodiversity. Freshwater fungi have been studied in Thailand over several decades initiated by Tubaki et al. (1983) who found 40 Ingoldian fungi in the stream foams. Many new freshwater taxa have since been reported in Thailand, especially a large number of lignicolous freshwater ascomycetes (Sivichai et al. 1998, 2000, 2002; Jones et al. 1999; Sivichai and Boonyene 2004; Zhang et al. 2011; Luo et al. 2019; Dong et al. 2020; Calabon et al. 2021, 2022). Until 2020, more than 302 freshwater taxa had been reported from Thailand (Zhang et al. 2011; Calabon et al. 2021). In this study, we introduce three taxa of Pleurotheciaceae, collected from freshwater habitats in Thailand. With phylogenetic analysis of ITS, LSU, SSU, RPB2 and TEF1-α sequence data, they are placed in Dematipyriforma, Pleurothecium and Rhexoacrodictys within Pleurotheciaceae. Of these three species, one is identified as Pleurothecium aquaticum, one as Rhexoacrodictys fimicola, and the third as a new species in Dematipyriforma. In addition, Rhexoacrodictys nigrospora is transferred to Dematipyriforma based on morphological and phylogenetic evidence.

Materials and methods

Collection, isolation and morphology

Submerged decaying woods were collected from the streams in Thailand. The sample incubation, examination and morphological studies were referred to the methods described by Luo et al. (2018b). Single spore isolations were followed the methods outlined by Senanayake et al. (2020). Specimens (dry wood with fungal material) were deposited in the herbarium of Mae Fah Luang University (MFLU), Chiang Rai, Thailand and Herbarium of Cryptogams Kunming Institute of Botany Academia Sinica (KUN-HKAS). Pure cultures were deposited in Mae Fah Luang University Culture Collection (MFLUCC) and Kunming Institute of Botany culture collection (KUNCC). Faces of Fungi and Index Fungorum numbers were registered as outlined in Jayasiri et al. (2015) and Index Fungorum (2022). The descriptions are added to it GMS database (Chaiwan et al. 2021).

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from fungal mycelium (Rhexoacrodictys erecta and Pleurothecium aquaticum) or directly from the conidiamatal tissue thalli of fungi (Dematipyriforma muriformis) as outlined by Wanasinghe et al. (2018). The Ezup Column Fungi Genomic DNA Purification Kit (Sangon Biotech, China) was used to extract DNA following the manufacturer’s instructions. ITS, LSU, SSU, RPB2 and TEF1-α gene regions were amplified using the primer pairs ITS5/ITS4, LR0R/LR7, NS1/NS4, fRPB2-5F/fRPB2-7cR and 983F/2218R, respectively (Vilgalys and Hester 1990; White et al. 1990; Liu et al. 1999). The amplification was performed in a 25 μl reaction volume containing 9.5 μl ddH2O, 12.5 μl 2 × Taq PCR Master Mix with blue dye (Sangon Biotech, China), 1 μl of DNA template and 1 μl of each primer (10 μM). The amplification condition for ITS, LSU, SSU, RPB2 and TEF1-α were followed Luo et al. (2018b). DNA sequencing of PCR products were carried out using the above-mentioned PCR primers at Tsingke Biological Engineering Technology and Services Co. (Yunnan, P.R. China).

Phylogenetic analyses

The taxa used in the phylogenetic analysis were obtained from previous studies (Table 1) (Hernandez-Restrepo et al. 2017; Luo et al. 2018a, 2019; Dayarathne et al. 2019; Hyde et al. 2020b; Réblová et al. 2020; Boonmee et al. 2021; Dong et al. 2021) and downloaded from GenBank. SEQMAN v. 7.0.0 (DNASTAR, Madison, WI) was used to assemble the consensus sequences and MAFFT v.7 online program ( was used to align the sequences (Katoh et al. 2019). BioEdit was used to manually adjust the alignments and the alignment fasta file was converted to Phylip format by Alivew (Hall 2021; Larsson 2014).

Table 1.

Taxa used in this study; the ex-type strains were indicated in bold, newly generated sequences are indicated by * after the species name.

Species Strain number GenBank accession number
Adelosphaeria catenata CBS 138679 KT278721 KT278707 KT278692 KT278743
Anapleurothecium botulisporum CBS 132713 KY853423 KY853483
Ascotaiwania lignicola NIL00005 HQ446341 HQ446364 HQ446284 HQ446419 HQ446307
Ascotaiwania sawadae SS00051 HQ446340 HQ446363 HQ446283 HQ446418 HQ446306
Bactrodesmiastrum obovatum FMR 6482 FR870264 FR870266
Bactrodesmiastrum pyriforme FMR 10747 FR870263 FR870265
Bactrodesmium abruptum CBS 144404 MN699391 MN699408 MN699365 MN704288 MN704313
Bactrodesmium leptopus CBS 144542 MN699388 MN699423 MN699374 MN704297 MN704321
Bactrodesmium obovatum CBS 144077 MN699395 MN699424 MN699375 MN704298 MN704322
Canalisporium exiguum SS00809 GQ390296 GQ390281 GQ390266 HQ446436
Canalisporium grenadoideum SS03615 GQ390267 GQ390252 HQ446420 HQ446309
Coleodictyospora muriformis MFLUCC 18–1243 MW981642 MW981648 MW981704
Coleodictyospora muriformis MFLUCC 18–1279 MW981643 MW981649 MW981705
Conioscypha hoehnelii FMR 11592 KY853437 KY853497 HF937348
Conioscypha lignicola CBS 335.93 AY484513 JQ437439 JQ429260
Conioscypha peruviana ILL41202 KF781539
Conioscypha pleiomorpha FMR 13134 KY853438 KY853498
Dematipyriforma aquilaria CGMCC 3.17268 KJ138621 KJ138623 KJ138622
Dematipyriforma muriformis* MFLU 21–0146 OM654773 OM654770 OM672032
Dematipyriforma nigrospora MFLUCC 21-0096 MZ538524 MZ538558 MZ567100
Dematipyriforma nigrospora MFLUCC 21-0097 MZ538525 MZ538559 MZ538574 MZ567113 MZ567101
Fuscosporella pyriformis MFLUCC 16–0570 KX550896 KX550900 KX576872
Helicoascotaiwania farinosa ILLS 53605 AY094189
Helicoascotaiwania farinosa DAOMC 241947 JQ429145 JQ429230
Helicoascotaiwania lacustris CBS 145963 MN699430 MN699382 MN704304 MN704329
Helicoascotaiwania lacustris CBS 145964 MN699400 MN699431 MN699383 MN704305
Helicoascotaiwania lacustris CBS 146144 MN699401 MN699432 MN699384 MN704306
Leotia lubrica AFTOL-ID1 DQ491484 AY544644 AY544746 DQ470876 DQ028596
Melanotrigonum ovale CBS 138815 KT278722 KT278711 KT278698 KT278747
Microglossum rufum AFTOL-ID 1292 DQ470981 DQ471033 DQ470933 DQ471104
Monotosporella setosa HKUCC3713 AF132334
Mucispora obscuriseptata MFLUCC 15–0618 KX550892 KX550897
Mucispora phangngaensis MFLUCC 16–0865 MG388210 MG388207
Neomonodictys muriformis MFLUCC 16–1136 MN644509 MN644485 MN646856
Obliquifusoideum guttulatum MFLUCC 18–1233 MW981645 MW981650 MW981706
Parafuscosporella garethii FF00725.01 KX958430 KX958428 KX958432
Parafuscosporella moniliformis MFLUCC 15–0626 KX550895 KX550899
Parafuscosporella mucosa MFLUCC 16–0571 MG388211 MG388208
Phaeoisaria aquatica MFLUCC 16–1298 MF399237 MF399254 MF401406
Phaeoisaria clematidis MFLUCC 17–1968 MG837022 MG837017 MG837027
Phaeoisaria fasciculata CBS 127885 KT278705 KT278693 KT278741
Phaeoisaria filiformis MFLUCC 18–0214 MK878381 MK835852 MK834785 MN200285
Phaeoisaria guttulata MFLUCC 17–1965 MG837021 MG837016 MG837026
Phaeoisaria pseudoclematidis MFLUCC 11–0393 KP744501 KP753962
Phaeoisaria sedimenticola CGMCC 3.14949 JQ031561
Phaeoisaria sedimenticola S-908 MK878380 MK835851 MN200284
Phaeoisaria sparsa FMR11939 HF677185
Phragmocephala stemphylioides DAOM 673211 KT278730 KT278717
Pleurotheciella aquatica MFLUCC 17–0464 MF399236 MF399253 MF399220 MF401405
Pleurotheciella centenaria DAOM 229631 JQ429234 JQ429246 JQ429265
Pleurotheciella fusiformis MFLUCC 17–0115 MF399232 MF399249 MF399217 MF401402
Pleurotheciella guttulata KUMCC 15–0296 MF399240 MF399257 MF399223 MF401409
Pleurotheciella krabiensis MFLUCC 18–0852 MG837018 MG837013 MG837023
Pleurotheciella lunata MFLUCC 17–0111 MF399238 MF399255 MF399221 MF401407
Pleurotheciella rivularia CBS 125238 JQ429232 JQ429244 JQ429263
Pleurotheciella rivularia CBS 125237 JQ429233 JQ429245 JQ429264
Pleurotheciella saprophytica MFLUCC 16–1251 MF399241 MF399258 MF399224 MF401410
Pleurotheciella submersa MFLUCC 17–1709 MF399243 MF399260 MF399226 MF401412
Pleurotheciella submersa MFLUCC 17–0456 MF399244 MF399261 MF399227 MF401413
Pleurotheciella tropica MFLUCC 16–0867 MG837020 MG837015 MG837025
Pleurotheciella uniseptata DAOM 673210 KT278729 KT278716
Pleurothecium aquaticum MFLUCC 17–1331 MF399245 MF399263
Pleurothecium aquaticum * KUMCC 21-0477 OM654775 OM654772 OM654807 OM672034 OM672033
Pleurothecium floriforme MFLUCC 15–0628 NR_156614 NG_059791
Pleurothecium obovoideum CBS 209.95 EU041784 EU041841
Pleurothecium pulneyense MFLUCC 16–1293 MF399262 MF399228 MF401414
Pleurothecium recurvatum CBS 138686 KT278715 KT278702
Pleurothecium semifecundum CBS 131271 JQ429240 JQ429254 JQ429270
Rhexoacrodictys erecta HSAUPmyr4622 KU999964 KX033556 KX033526
Rhexoacrodictys erecta IFRD500–016 MT555421 MT559123 MT555735
Rhexoacrodictys erecta HSAUP myr6489 KU999963 KX033555 KX033525
Rhexoacrodictys fimicola HMAS 47737 KU999960 KX033553 KX033522
Rhexoacrodictys fimicola HMAS 42882 KU999962 KX033554 KX033524
Rhexoacrodictys fimicola HMAS 43690 KU999957 KX033550 KX033519
Rhexoacrodictys fimicola * MFLUCC 18–0340 OM654774 OM654771 OM654806
Saprodesmium dematiosporium KUMCC 18–0059 MW981646 MW981647 MW981707
Savoryella aquatica SS03801 HQ446372 HQ446292 HQ446405 HQ446326
Savoryella lignicola NF00204 HQ446378 HQ446300 HQ446413 HQ446334
Sterigmatobotrys macrocarpa MR2973 GU017317
Sterigmatobotrys rudis DAOM 229838 JQ429152 JQ429241 JQ429256 JQ429272
Sterigmatobotrys uniseptata MFLUCC 15–0358 MK878379 MK835850 MK834784

Maximum likelihood (ML) analysis generated using the RAxML-HPC2 on XSEDE (v.8.2.8) in the CIPRES Science Gateway (, Stamatakis 2006; Stamatakis et al. 2008; Miller et al. 2010) with rapid bootstrap analysis, followed by 1000 bootstrap replicates, using the GTR+I+G model of evolution.

Bayesian analysis was performed by MrBayes v. 3.2 (Ronquist et al. 2012), best-fit model of DNA evolution for the Bayesian inference analysis was estimated by MrModeltest v. 2.2 (Nylander 2004) and the GTR+I+G model was selected for LSU, ITS, RPB2 and TEF1-α, GTR+G model was selected for SSU. Posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002) was defined by Bayesian Markov Chain Monte Carlo (BMCMC) sampling method in MrBayes v. 3.0b4 (Huelsenbeck and Ronquist 2001). Six simultaneous Markov Chains were run for 50,000,000 generations and trees were sampled every 500th generation (resulting in 100,000 trees). The first 20% trees that represented the burn-in phase were discarded and the remaining 80% (post burn-in) trees used for calculating posterior probabilities (PP) for the majority rule consensus tree.

Phylogenetic trees were visualized with FigTree v. 1.4.2 (Rambaut 2014) and edited in Microsoft Office PowerPoint 2019 (Microsoft Inc., United States). Newly generated sequences in this study were deposited in GenBank.


Phylogenetic analyses

The dataset of combined ITS, LSU, SSU, RPB2 and TEF1-α sequence data comprises 81 strains with 4257 characters including gaps (ITS: 509 bp, LSU: 1006 bp, SSU: 862 bp, RPB2: 1032 bp, TEF1-α: 848 bp). Leotia lubrica (AFTOL-ID1) and Microglossum rufum (AFTOL-ID 1292) were used as outgroup taxa. RAxML and Bayesian analyses were conducted and resulted in generally congruent topologies. The best RAxML tree with a final likelihood value of –45872.924927 is presented. The matrix had 2433 distinct alignment patterns, with 44.65% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.234712, C = 0.261626, G = 0.290634, T = 0.213028; substitution rates AC = 1.347806, AG = 2.754719, AT = 1.490447, CG = 1.095887, CT = 6.696475, GT = 1.000000; gamma distribution shape parameter α = 0.316898.

In the phylogenetic analysis, Dematipyriforma muriformis (MFLU 21–0146) clustered with the ex-type strain of D. aquilaria (CGMCC 3.17268) with low support (Fig. 1). The new isolate of Rhexoacrodictys fimicola (MFLUCC 18–0340) clustered with three strains of R. fimicola (HMAS 42882, HMAS 43690 and HMAS 47737) with 100% ML/1.00 PP support (Fig. 1). Pleurothecium aquaticum (KUNCC 21–0477) clustered with the ex-type strain of P. aquaticum (MFLUCC 17–1331) with 100% ML/1.00 PP support (Fig. 1).

Figure 1. 

Phylogram based on a combined ITS, LSU SSU, RPB2 and TEF1-α sequence data of selected members of four orders of the Savoryellomycetidae. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities (PP) greater than 0.95 are given as ML/PP above the nodes. Newly obtained sequences are indicated in red and ex-type strains are in bold.


Dematipyriforma muriformis D.F. Bao, K.D. Hyde & Z.L. Luo, sp. nov.

Fig. 2


Referring to the muriform conidia of this species.


MFLU 21–0146.


Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph: Colonies on substratum superficial, scattered, black, shining, granulate. Mycelium immersed, composed of hyaline, branched, septate, smooth, hyphae. Conidiomata sporodochial, subhyaline. Conidiophores 10–26.5 × 2–3 μm (x‒ = 18.2 × 2.3 μm, n = 20), micronematous to semi-macronematous, mononematous, fasciculate, simple or branched, hyaline, cylindrical, smooth. Conidiogenous cells monoblastic, integrated, terminal, determinate, hyaline, smooth. Conidia 23–26 × 15.5–18 μm (x‒ = 24.6 × 16.7 μm, n = 30), acrogenous, solitary, smooth, thick-walled, ellipsoidal to obovoid, muriform, rounded at apex, pointed at base, with 3–5 transverse septa, 1-longitudinal septum in all cells and rarely in end cells, slightly constricted at septa, subhyaline to pale olivaceous when young, olive to dark brown at maturity.

Material examined

Thailand, Bangkok Province, Bang Kapi District, on decaying wood submerged in a freshwater stream, 3 October 2017, Z.L. Luo, Bsite 4–3–1 (MFLU 21–0146, holotype; KUN-HKAS 122858, isotype).


In the phylogenetic analysis, Dematipyriforma muriformis clustered with the ex-type strain of D. aquilaria (CGMCC 3.17268) within Pleurotheciaceae with low support (Fig. 1). The ITS blast result in NCBI GenBank showed that D. muriformis (MFLU 21–0146) is 92.36% and 91.92% similar to D. nigrospora (MFLUCC 21-0097) and D. aquilaria (CGMCC 3.17268) respectively.

Dematipyriforma muriformis resembles D. aquilaria in having micronematous, mononematous, smooth septate conidiophores, monoblastic, integrated, terminal, determinate conidiogenous cells and solitary, muriform conidia. However, D. muriformis differs from D. aquilaria in having hyaline conidiophores and slightly smaller conidia (23–26 × 15.5–18 vs. 25–37.5 × 15–22.5 μm). In addition, conidia of D. muriformis are subhyaline to pale olivaceous when young, olive to dark brown at maturity, with 3–5 transverse septa, 1-longitudinal septum in all cells and rarely in end cells. Whereas, D. aquilaria has pale grey olivaceous to pale brown conidia with 4–5 transverse septa and 0–2 longitudinal septa (Sun et al. 2017).

Dematipyriforma muriformis shares some similar characteristics with Neomonodictys taxa in Pleurotheciaceae, such as monoblastic, integrated, terminal, determinate conidiogenous cells and muriform conidia. Neomonodictys, however, lacks sporodochial conidiomata and conidia of Neomonodictys are subglobose to globose, while, Dematipyriforma muriformis has ellipsoidal to obovoid conidia (Hyde et al. 2020b).

Figure 2. 

Dematipyriforma muriformis (MFLU 21–0146, holotype) a, b colonies on wood c–d conidiomata e–i conidiophore with conidia j–q conidia. Scale bars: 30 μm (c–d, o–q); 20 μm (e–n).

Dematipyriforma nigrospora (Boonmee, D.F. Bao & K.D. Hyde) D.F. Bao, K.D. Hyde & Z.L. Luo, comb. nov.

Rhexoacrodictys nigrospora Boonmee, D.F. Bao & K.D. Hyde, in Boonmee et al., Fungal Diversity 111: 200 (2021).


Thailand, Phetchabun Province, on decaying bark, 25 July 2019, S. Boonmee, LSP03 (MFLU 21–0073).

Descriptions and illustrations

See Boonmee et al. (2021).


Rhexoacrodictys nigrospora was introduced by Boonmee et al. (2021) based on morphological characters and phylogenetic analysis. In our phylogenetic analysis, R. nigrospora clustered with two Dematipyriforma species (D. aquilaria and D. muriformis) in a distinct clade within Pleurotheciaceae (Fig. 1). Therefore, we transfer Rhexoacrodictys nigrospora to Dematipyriforma, as Dematipyriforma nigrospora comb. nov.

Dematipyriforma nigrospora resembles D. muriformis in having micronematous or semi-macronematous, mononematous conidiophores and monoblastic, polyblastic, integrated, terminal conidiogenous cells. However, D. nigrospora differs from D. muriformis in having brown to dark brown conidiophores and globose to subglobose, dark brown to black conidia (Boonmee et al. 2021). Conidiophores of D. muriformis are hyaline and conidia are ellipsoidal to obovoid, muriform, and subhyaline to pale olivaceous when young, olive to dark brown at maturity.

Rhexoacrodictys fimicola (M.B. Ellis & Gunnell) W.A. Baker & Morgan-Jones, in Baker, Partridge & Morgan-Jones, Mycotaxon 82: 103 (2002)

Fig. 3


Maya, Perak, on elephant dung, September 1958, A.H.S, Onions, IMI 76413.


Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph: Colonies on the substratum superficial, effuse, hairy or velvety, black. Mycelium mostly immersed, composed of branched, septate, smooth, pale brown hyphae. Conidiophores (17.5–)20–44.5 (–65.5) × 2.5–4.0 μm (x‒ = 32.2 × 3.4 μm, n = 20), macronematous, mononematous, erect, straight or slightly flexuous, thick-walled, smooth, orange-brown or brown, 3–7-septate. Conidiogenous cells monoblastic, integrated, terminal. Conidia 16.5–24 × 11–15 μm (x‒ = 20.3 × 13 μm, n = 30), solitary, dry, acrogenous, broadly oval to subglobose, muriform, transversely and longitudinally septate, with transverse septa typically spanning the whole conidial width, with longitudinal septa typically incomplete, short; dark-blackish brown to black, smooth, narrowly truncate at the base.

Figure 3. 

Rhexoacrodictys fimicola (MFLU 21–0147, new record) a–c colonies on wood d–j conidiophores with conidia k–m conidi n germinating conidium o–r re-produced asexual morph of Rhexoacrodictys fimicola s–t culture on PDA from surface and reverse. Scale bars: 20 μm (d–j, o–r); 10 μm (k–n).

Cultural characteristics

Conidia germinating on PDA within 24 h. Germ tubes produced from the basal cell. Colonies on PDA reaching 3 cm diameter in 30 days at 20–25 °C, pale brown, with dense, tight mycelia on the surface, sparse at the margin, reverse dark brown, with smooth margin. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, monoblastic, integrated, hyaline to pale brown, smooth. Conidia broad oval to subglobose, muriform, strongly constricted at all the septa, hyaline when young, brown to grayish-brown when aged, smooth-walled.

Material examined

Thailand, Bangkok Province, Bang Kapi District, on decaying wood submerged in a freshwater stream, 3 October 2017, Z.L. Luo, Bsite 4–3–2 (MFLU 21–0147 = KUN-HKAS 122859), living culture, MFLUCC 18–0340.


In the phylogenetic analysis, our new isolate MFLUCC 18–0340 clustered with three strains of Rhexoacrodictys fimicola (HMAS 42882, HMAS 43690 and HMAS 47737) with strong support (100% ML/ 1.00 PP). The nucleotide BLASTn search of ITS showed that our new strain (MFLUCC 18–0340) has 99.7%, 99.3% and 99.1% similarities with strain HMAS 43690, HMAS 47737 and HMAS 42882 of Rhexoacrodictys fimicola, respectively. Morphologically, our new collection is similar to R. fimicola in having macronematous, mononematous, indeterminate conidiophores, integrated, terminal, monoblastic, pale brown conidiogenous cells and broadly oval to subglobose, transversely and longitudinally septate, smooth, brown to black conidia, with the size of conidia and conidiophores are overlapping (Ellis 1961; Baker et al. 2002). Based on both phylogeny and morphology, we identified our species as R. fimicola.

Rhexoacrodictys fimicola was originally introduced by Ellis (1961) as Acrodictys fimicola. Baker et al. (2002) transferred A. fimicola to Rhexoacrodictys based on morphological characteristics. Rhexoacrodictys fimicola has been reported on Bambusa vulgaris and elephant dung from Africa and Malaysia respectively. Our collection, on the other hand, was collected from freshwater habitats and represents the first time it was reported from Thailand.

Pleurothecium aquaticum Z.L. Luo, H.Y. Su & K.D. Hyde, in Luo, Hyde, Bhat, Jeewon, Maharachchikumbura, Bao, Li, Su, Yang & Su, Mycol. Prog. 17(5): 526 (2018)

Fig. 4


Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph: colonies on substratum, effuse, shining, dark brown to black. Mycelium partly immersed, composed of septate, branched, smooth, dark brown hyphae. Conidiophores 84–110 × 3–4 μm (x‒ = 97 × 3.4 μm, n = 10), macronematous, mononematous, erect, simple, unbranched, straight or slightly flexuous, 5–8-septate, dark brown, pale towards apex, smooth. Conidiogenous cells integrated, polyblastic, terminal, hyaline, denticulate, smooth. Conidia 18–22 × 4–5 μm (x‒ = 20 × 4.5 μm, SD = 4 n = 30), acrogenous, solitary, clavate, mostly curved, rounded at apex, tapering at base, hyaline, 3-septate, with guttulate cells, smooth.

Cultural characteristics

Conidia germinating on PDA within 24 h. Germ tubes produced from the basal and apical cells. Colonies on PDA reaching 2.3 cm diameter in 30 days at 20–25 °C, with dense mycelia, dry, rigid, rugose, dark brown, reverse dark brown.

Material examined

Thailand, Prachuap Khan, on submerged decaying wood, 15 August 2017, V. Kumar, site1–24–2 (MFLU 21–0148 = KUN-HKAS 122857), living culture, KUNCC 21–0477.


In the phylogenetic analysis, our new collection KUNCC 21–0477 clustered with the ex-type strain of Pleurothecium aquaticum (MFLUCC 17–1331) with high (100% ML/1.00 PP). In addition, the ITS and LSU BLASTn search on NCBI GenBank showed that our new strain is 99.88% and 97.45% similarities to the ex-type of P. aquaticum (MFLUCC 17–1331). The new collection is morphologically similar to P. aquaticum in having macronematous, mononematous, septate, brown, pale brown towards the apex conidiophores, integrated, terminal, polyblastic, denticulate conidiogenous cells and hyaline, cylindrical or clavate, rounded at the apex, obtuse and tapering towards base, 3-septate conidia. We therefore identified our new collection as P. aquaticum. Pleurothecium aquaticum was introduced by Luo et al. (2018a) collected from freshwater habitats in China. Our new collection, on the other hand, was collected from Thailand and is a new record for Thailand.

Figure 4. 

Pleurothecium aquaticum (MFLU 21–0148, new record) a, b colonies on wood c, d conidiophores e, f conidiogenous cells g–i conidia m germinating conidium n, o culture on PDA from surface and reverse. Scale bars: 30 μm (c, d); 10 μm (e–m).


Pleurotheciaceae is a diverse family. The sexual morphs of Pleurotheciaceae are quite similar and difficult to distinguish without molecular data (Réblová et al. 2016; Hyde et al. 2020a). However, the asexual morphs in the family are morphologically diverse. Most genera have mononematous, macrounematous conidiophores (Anapleurothecium, Pleurothecium, Pleurotheciella and Rhexoacrodictys) (Réblová et al. 2016; Luo et al. 2018a, 2019; Hyde et al. 2020a), whereas some genera have synnematous conidiophores (Phaeoisaria and Phragmocephala) (Höhnel 1919; Mason and Hughes 1951; Seifert et al. 2011; Wijayawardene et al. 2012; Su et al. 2015; Réblová et al. 2016; Luo et al. 2018a), and others with micronematous or reduced conidiophores (Neomonodictys and Sterigmatobotrys). (Hyde et al. 2020b). Conidiogenous cells of Anapleurothecium, Pleurothecium, Phaeoisaria and Pleurotheciella are polyblastic and denticulate (Réblová et al. 2012, 2016; Monteiro et al. 2016; Luo et al. 2018a). Phragmocephala and Monotosporella have monoblastic conidiogenous cells (Mason and Hughes 1951; Hyde and Yanna 2002; Wijayawardene et al. 2012; Su et al. 2015). Conidia of Pleurotheciaceae are diverse in their shape, color and septation. Conidia of Sterigmatobotrys are fusiform and in persistent chains (Heuchert et al. 2018); Helicoascotaiwania has helicosporous conidia (Dayarathne et al. 2019); Anapleurothecium, Melanotrigonum, Pleurothecium, Phaeoisaria and Pleurotheciella have clavate, ellipsoidal, obovoidal, fusiform-cylindrical, hyaline or brown, aseptate or transversely septate conidia (Réblová et al. 2012, 2016; Monteiro et al. 2016; Hernandez-Restrepo et al. 2017; Luo et al. 2018a); Monotosporella, Neomonodictys and Phragmocephalahave ellipsoidal or subglobose to globose conidia (Mason and Hughes 1951; Hyde and Yanna 2002;Wijayawardene et al. 2012; Su et al. 2015; Hyde et al. 2020b). However, conidia of Neomonodictys are muriform (Hyde et al. 2020b), whereas, Phragmocephala and Monotosporella have transversely septate conidia.

In this study, we introduced a new asexual species, Dematipyriforma muriformis based on both morphology and phylogeny. Dematipyriforma was introduced by Sun et al. (2017) with a single species D. aquilaria which was reported as an endophyte from Aquilaria crassna in China. While our new species is a saprobe isolated on submerged wood from freshwater habitats in Thailand. In addition, Rhexoacrodictys nigrospora is transferred to Dematipyriforma in this study. Currently, three species are accepted in the genus. Morphologically, the muriform conidia of Dematipyriforma are similar to Neomonodictys, Saprodesmium and Coleodictyospora. However, Dematipyriforma can be distinguished from Neomonodictys by the shape of conidia (ellipsoidal to obovoid vs. subglobose to globose) and conidiophores (semi-micronematous to macronematous vs. micronematous or lacking conidiophores, Hyde et al. 2020b). Dematipyriforma differs from Coleodictyospora in the conidia lacking a semi-gelatinous sheath (Dong et al. 2021). Dematipyriforma is distinct from Saprodesmium by the micronematous to semi-macronematous, simple or branched, hyaline, cylindrical, conidiophores, whereas, conidiophores of Saprodesmium are micronematous, unbranched, consisted of 1–4 subglobose smooth, hyaline cells (Dong et al. 2021).

Rhexoacrodictys comprises six species of which four species (R. erecta, R. fimicola, R. martini and R. queenslandica) have sequence data available in the GenBank. Among them, R. martini and R. queenslandica were transferred to Distoseptispora and Junewangia based on phylogenetic analysis (Xia et al. 2017). However, sequence data of R. martini are doubted by several studies (Sun et al. 2020; Shen et al. 2021), as its morphology does not fit with the characters of Distoseptispora. Rhexoacrodictys erecta and R. fimicola clustered within Pleurotheciaceae (Luo et al. 2019; Dong et al. 2021). The placement of Rhexoacrodictys was questionable since it was established. Baker et al. (2002) established the genus; however, they did not mention the placement of the genus. Xia et al. (2017) firstly provided sequence data for Rhexoacrodictys erecta (Type species of Rhexoacrodictys) and R. fimicola based on their fresh collections, their phylogenetic analysis showed that R. erecta and R. fimicola clustered within Savoryellaceae. However, they did not include the related orders (Conioscyphales, Fuscosporellales and Pleurotheciales) in Savoryellomycetidae. Luo et al. (2019) found that R. erecta and R. fimicola grouped in Pleurotheciaceae. Recently, Dong et al. (2021) obtained the same result as Luo et al. (2019). However, Boonmee et al. (2021) and Wijayawardene et al. (2022) placed Rhexoacrodictys in Savoryellaceae (Savoryellales). Our result is consistent with Luo et al. (2019) and Dong et al. (2021), the two species clustered within Pleurotheciaceae (Fig. 1). On the other hand, the morphology of Rhexoacrodictys is similar to Dematipyriforma, Neomonodictys and Saprodesmium, in having muriform conidia, micronematous conidiophores and holoblastic, monoblastic conidiogenous cells. Therefore, we formally accepted Rhexoacrodictys in Pleurotheciaceae (Pleurotheciales) based on morphological characters and phylogenetic analysis.

In our phylogenetic analysis, Rhexoacrodictys erecta and R. fimicola clustered with Monotosporella setosa which is the type species of Monotosporella. Morphologically, R. erecta and R. fimicola fit well within the genus concept of Monotosporella in having macronematous, mononematous, brown, septate conidiophores, monoblastic, percurrent conidiogenous cells and acrogenous, brown septate conidia (Hughes 1958; Baker et al. 2002; Hyde and Yanna 2002). However, the strain of Monotosporella setosa (HKUCC 3713) lacks a morphological description. Therefore, further study is necessary to clarify the relationship between Rhexoacrodictys and Monotosporella, whether they should be combined into one genus or not. In addition, our phylogenetic analysis showed that three strains of R. erecta clustered with Monotosporella setosa. However, M. erecta differs from M. setosa in having transverse and longitudinal septation, while, conidia of M. setosa only have transverse septa. Only LSU sequence data is available for M. setosa, which is not significant to distinguish in the phylogenetic tree, but morphologically they are quite distinct. Hence, we maintain them as two distinct species, however, further morphological and phylogenetic analysis is required to clarify the relationship between these two species.

In our phylogenetic analysis, Pleurothecium obovoideum was placed distant from Pleurothecium and close to Neomonodictys muriformis and Coleodictyospora muriformis which is consistent with recent studies (Luo et al. 2018a, 2019; Hyde et al. 2020b). Pleurothecium obovoideum was introduced by Arzanlou et al. (2007) based on morphological characters. However, their analysis showed that P. obovoideum clustered with Ascotaiwania hughesii and with more sequence data now available for Pleurothecium species, P. obovoideum is shown phylogenetically distinct from Pleurothecium. Morphologically, P. obovoideum is similar to Pleurothecium in having distinct brown conidiophores, polyblastic, denticulate conidiogenous cells and pale brown, ellipsoidal to obovate conidia. However, conidia of P. obovoideum are aseptate and solitary or in short chains whereas the conidia of Pleurothecium are solitary and unicellular or septate. Thus, the placement of P. obovoideum needs revision in the future with more evidence.


We would like to thank the National Natural Science Foundation of China (Project ID: 32060005 and 31970021) for financial support. This study was also supported by the Yunnan Fundamental Research Project (grant NO. 202101AU070137, 202201AW070001) and Thailand research fund “Macrofungi diversity research from the Lancang-Mekong Watershed and Surrounding areas (Grant no. DBG6280009)”. Dan-Feng Bao would like to thank Shaun Pennycook from Landcare Research, Auckland, New Zealand, for advising on the taxon names. Wen-Li Li is acknowledged for her help with DNA extraction and PCR amplification.


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