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. 2022 Sep 29;93:57–79. doi: 10.3897/mycokeys.93.87797

Lignicolous freshwater ascomycetes from Thailand: Introducing Dematipyriformamuriformis sp. nov., one new combination and two new records in Pleurotheciaceae

Dan-Feng Bao 1,2,3, Darbhe J Bhat 4, Saranyaphat Boonmee 2, Kevin D Hyde 2, Zong-Long Luo 1,✉, Sarunya Nalumpang 3,✉
PMCID: PMC9836478  PMID: 36761914

Abstract

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

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

Introduction

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 Aquilariacrassna (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 Rhexoacrodictyserecta 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 Pleurotheciumaquaticum, one as Rhexoacrodictysfimicola, and the third as a new species in Dematipyriforma. In addition, Rhexoacrodictysnigrospora 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 (Rhexoacrodictyserecta and Pleurotheciumaquaticum) or directly from the conidiamatal tissue thalli of fungi (Dematipyriformamuriformis) 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 (http://mafft.cbrc.jp/alignment/server/) 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
ITS LSU SSU RPB2 TEF1-α
Adelosphaeriacatenata CBS 138679 KT278721 KT278707 KT278692 KT278743 –
Anapleurotheciumbotulisporum CBS 132713 KY853423 KY853483 – – –
Ascotaiwanialignicola NIL00005 HQ446341 HQ446364 HQ446284 HQ446419 HQ446307
Ascotaiwaniasawadae SS00051 HQ446340 HQ446363 HQ446283 HQ446418 HQ446306
Bactrodesmiastrumobovatum FMR 6482 FR870264 FR870266 – – –
Bactrodesmiastrumpyriforme FMR 10747 FR870263 FR870265 – – –
Bactrodesmiumabruptum CBS 144404 MN699391 MN699408 MN699365 MN704288 MN704313
Bactrodesmiumleptopus CBS 144542 MN699388 MN699423 MN699374 MN704297 MN704321
Bactrodesmiumobovatum CBS 144077 MN699395 MN699424 MN699375 MN704298 MN704322
Canalisporiumexiguum SS00809 GQ390296 GQ390281 GQ390266 HQ446436 –
Canalisporiumgrenadoideum SS03615 – GQ390267 GQ390252 HQ446420 HQ446309
Coleodictyosporamuriformis MFLUCC 18–1243 MW981642 MW981648 MW981704 – –
Coleodictyosporamuriformis MFLUCC 18–1279 MW981643 MW981649 MW981705 – –
Conioscyphahoehnelii FMR 11592 KY853437 KY853497 HF937348 – –
Conioscyphalignicola CBS 335.93 – AY484513 JQ437439 JQ429260 –
Conioscyphaperuviana ILL41202 – KF781539 – – –
Conioscyphapleiomorpha FMR 13134 KY853438 KY853498 – – –
Dematipyriformaaquilaria CGMCC 3.17268 KJ138621 KJ138623 KJ138622 – –
Dematipyriformamuriformis * MFLU 21–0146 OM654773 OM654770 – – OM672032
Dematipyriformanigrospora MFLUCC 21-0096 MZ538524 MZ538558 – – MZ567100
Dematipyriformanigrospora MFLUCC 21-0097 MZ538525 MZ538559 MZ538574 MZ567113 MZ567101
Fuscosporellapyriformis MFLUCC 16–0570 – KX550896 KX550900 KX576872 –
Helicoascotaiwaniafarinosa ILLS 53605 – AY094189 – – –
Helicoascotaiwaniafarinosa DAOMC 241947 JQ429145 JQ429230 – – –
Helicoascotaiwanialacustris CBS 145963 – MN699430 MN699382 MN704304 MN704329
Helicoascotaiwanialacustris CBS 145964 MN699400 MN699431 MN699383 MN704305 –
Helicoascotaiwanialacustris CBS 146144 MN699401 MN699432 MN699384 MN704306 –
Leotialubrica AFTOL-ID1 DQ491484 AY544644 AY544746 DQ470876 DQ028596
Melanotrigonumovale CBS 138815 KT278722 KT278711 KT278698 KT278747 –
Microglossumrufum AFTOL-ID 1292 – DQ470981 DQ471033 DQ470933 DQ471104
Monotosporellasetosa HKUCC3713 – AF132334 – – –
Mucisporaobscuriseptata MFLUCC 15–0618 – KX550892 KX550897 – –
Mucisporaphangngaensis MFLUCC 16–0865 – MG388210 MG388207 – –
Neomonodictysmuriformis MFLUCC 16–1136 MN644509 MN644485 – – MN646856
Obliquifusoideumguttulatum MFLUCC 18–1233 MW981645 MW981650 MW981706 – –
Parafuscosporellagarethii FF00725.01 – KX958430 KX958428 KX958432 –
Parafuscosporellamoniliformis MFLUCC 15–0626 – KX550895 KX550899 – –
Parafuscosporellamucosa MFLUCC 16–0571 – MG388211 MG388208 – –
Phaeoisariaaquatica MFLUCC 16–1298 MF399237 MF399254 – MF401406 –
Phaeoisariaclematidis MFLUCC 17–1968 MG837022 MG837017 MG837027 – –
Phaeoisariafasciculata CBS 127885 – KT278705 KT278693 KT278741 –
Phaeoisariafiliformis MFLUCC 18–0214 MK878381 MK835852 MK834785 – MN200285
Phaeoisariaguttulata MFLUCC 17–1965 MG837021 MG837016 MG837026 – –
Phaeoisariapseudoclematidis MFLUCC 11–0393 – KP744501 KP753962 – –
Phaeoisariasedimenticola CGMCC 3.14949 – JQ031561 – – –
Phaeoisariasedimenticola S-908 MK878380 MK835851 – – MN200284
Phaeoisariasparsa FMR11939 – HF677185 – – –
Phragmocephalastemphylioides DAOM 673211 KT278730 KT278717 – – –
Pleurotheciellaaquatica MFLUCC 17–0464 MF399236 MF399253 MF399220 MF401405 –
Pleurotheciellacentenaria DAOM 229631 – JQ429234 JQ429246 JQ429265 –
Pleurotheciellafusiformis MFLUCC 17–0115 MF399232 MF399249 MF399217 MF401402 –
Pleurotheciellaguttulata KUMCC 15–0296 MF399240 MF399257 MF399223 MF401409 –
Pleurotheciellakrabiensis MFLUCC 18–0852 MG837018 MG837013 MG837023 – –
Pleurotheciellalunata MFLUCC 17–0111 MF399238 MF399255 MF399221 MF401407 –
Pleurotheciellarivularia CBS 125238 – JQ429232 JQ429244 JQ429263 –
Pleurotheciellarivularia CBS 125237 – JQ429233 JQ429245 JQ429264 –
Pleurotheciellasaprophytica MFLUCC 16–1251 MF399241 MF399258 MF399224 MF401410 –
Pleurotheciellasubmersa MFLUCC 17–1709 MF399243 MF399260 MF399226 MF401412 –
Pleurotheciellasubmersa MFLUCC 17–0456 MF399244 MF399261 MF399227 MF401413 –
Pleurotheciellatropica MFLUCC 16–0867 MG837020 MG837015 MG837025 – –
Pleurotheciellauniseptata DAOM 673210 KT278729 KT278716 – – –
Pleurotheciumaquaticum MFLUCC 17–1331 MF399245 MF399263 – – –
Pleurotheciumaquaticum * KUMCC 21-0477 OM654775 OM654772 OM654807 OM672034 OM672033
Pleurotheciumfloriforme MFLUCC 15–0628 NR_156614 NG_059791 – – –
Pleurotheciumobovoideum CBS 209.95 EU041784 EU041841 – – –
Pleurotheciumpulneyense MFLUCC 16–1293 – MF399262 MF399228 MF401414 –
Pleurotheciumrecurvatum CBS 138686 – KT278715 KT278702 – –
Pleurotheciumsemifecundum CBS 131271 – JQ429240 JQ429254 JQ429270 –
Rhexoacrodictyserecta HSAUPmyr4622 KU999964 KX033556 KX033526 – –
Rhexoacrodictyserecta IFRD500–016 MT555421 MT559123 MT555735 – –
Rhexoacrodictyserecta HSAUP myr6489 KU999963 KX033555 KX033525 – –
Rhexoacrodictysfimicola HMAS 47737 KU999960 KX033553 KX033522 – –
Rhexoacrodictysfimicola HMAS 42882 KU999962 KX033554 KX033524 – –
Rhexoacrodictysfimicola HMAS 43690 KU999957 KX033550 KX033519 – –
Rhexoacrodictysfimicola * MFLUCC 18–0340 OM654774 OM654771 OM654806 – –
Saprodesmiumdematiosporium KUMCC 18–0059 MW981646 MW981647 MW981707 – –
Savoryellaaquatica SS03801 – HQ446372 HQ446292 HQ446405 HQ446326
Savoryellalignicola NF00204 – HQ446378 HQ446300 HQ446413 HQ446334
Sterigmatobotrysmacrocarpa MR2973 – GU017317 – – –
Sterigmatobotrysrudis DAOM 229838 JQ429152 JQ429241 JQ429256 JQ429272 –
Sterigmatobotrysuniseptata MFLUCC 15–0358 MK878379 MK835850 MK834784 – –
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Maximum likelihood (ML) analysis generated using the RAxML-HPC2 on XSEDE (v.8.2.8) in the CIPRES Science Gateway (https://www.phylo.org, 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.

Results

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). Leotialubrica (AFTOL-ID1) and Microglossumrufum (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, Dematipyriformamuriformis (MFLU 21–0146) clustered with the ex-type strain of D.aquilaria (CGMCC 3.17268) with low support (Fig. 1). The new isolate of Rhexoacrodictysfimicola (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). Pleurotheciumaquaticum (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.

Figure 1.

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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.

Taxonomy

. Dematipyriforma muriformis

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

9771DF07-5B27-5031-9B8E-090279283AA0

http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=553383

https://www.facesoffungi.org/?s=FoF10414

Fig. 2

Figure 2.

Figure 2.

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Dematipyriformamuriformis (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).

Etymology.

Referring to the muriform conidia of this species.

Holotype.

MFLU 21–0146.

Description.

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).

Notes.

In the phylogenetic analysis, Dematipyriformamuriformis 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.

Dematipyriformamuriformis 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).

Dematipyriformamuriformis 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, Dematipyriformamuriformis has ellipsoidal to obovoid conidia (Hyde et al. 2020b).

. Dematipyriforma nigrospora

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

A6148170-3B36-5B96-9677-CAD862AE897E

http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=553384

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

Holotype.

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

Descriptions and illustrations.

See Boonmee et al. (2021).

Notes.

Rhexoacrodictysnigrospora 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 Rhexoacrodictysnigrospora to Dematipyriforma, as Dematipyriformanigrospora comb. nov.

Dematipyriformanigrospora 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)

3D33BD6A-675E-542C-8B19-614ECB4162C6

Fig. 3

Figure 3.

Figure 3.

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Rhexoacrodictysfimicola (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 Rhexoacrodictysfimicolas–t culture on PDA from surface and reverse. Scale bars: 20 μm (d–j, o–r); 10 μm (k–n).

Holotype.

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

Description.

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.

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.

Notes.

In the phylogenetic analysis, our new isolate MFLUCC 18–0340 clustered with three strains of Rhexoacrodictysfimicola (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 Rhexoacrodictysfimicola, 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.

Rhexoacrodictysfimicola was originally introduced by Ellis (1961) as Acrodictysfimicola. Baker et al. (2002) transferred A.fimicola to Rhexoacrodictys based on morphological characteristics. Rhexoacrodictysfimicola has been reported on Bambusavulgaris 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)

7644B4BA-2F53-5575-A238-F9CE05C64759

Fig. 4

Figure 4.

Figure 4.

Open in a new tab

Pleurotheciumaquaticum (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).

Description.

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.

Notes.

In the phylogenetic analysis, our new collection KUNCC 21–0477 clustered with the ex-type strain of Pleurotheciumaquaticum (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. Pleurotheciumaquaticum 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.

Discussion

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, Dematipyriformamuriformis 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 Aquilariacrassna in China. While our new species is a saprobe isolated on submerged wood from freshwater habitats in Thailand. In addition, Rhexoacrodictysnigrospora 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. Rhexoacrodictyserecta 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 Rhexoacrodictyserecta (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, Rhexoacrodictyserecta and R.fimicola clustered with Monotosporellasetosa 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 Monotosporellasetosa (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 Monotosporellasetosa. 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, Pleurotheciumobovoideum was placed distant from Pleurothecium and close to Neomonodictysmuriformis and Coleodictyosporamuriformis which is consistent with recent studies (Luo et al. 2018a, 2019; Hyde et al. 2020b). Pleurotheciumobovoideum was introduced by Arzanlou et al. (2007) based on morphological characters. However, their analysis showed that P.obovoideum clustered with Ascotaiwaniahughesii 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.

Supplementary Material

XML Treatment for Dematipyriforma muriformis
XML Treatment for Dematipyriforma nigrospora
XML Treatment for Rhexoacrodictys fimicola
XML Treatment for Pleurothecium aquaticum

Acknowledgements

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.

Citation

Bao D-F, Bhat DJ, Boonmee S, Hyde KD, Luo Z-L, Nalumpang S (2022) Lignicolous freshwater ascomycetes from Thailand: Introducing Dematipyriforma muriformis sp. nov., one new combination and two new records in Pleurotheciaceae. MycoKeys 93: 57–79. https://doi.org/10.3897/mycokeys.93.87797

Contributor Information

Zong-Long Luo, Email: 514992672@qq.com.

Sarunya Nalumpang, Email: sarunya.v@cmu.ac.th.

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Associated Data

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

Supplementary Materials

XML Treatment for Dematipyriforma muriformis
mycokeys.93.87797-treatment1.xml (22.6KB, xml)
XML Treatment for Dematipyriforma nigrospora
mycokeys.93.87797-treatment2.xml (19.1KB, xml)
XML Treatment for Rhexoacrodictys fimicola
mycokeys.93.87797-treatment3.xml (13.1KB, xml)
XML Treatment for Pleurothecium aquaticum
mycokeys.93.87797-treatment4.xml (11KB, xml)

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