Research Article
Print
Research Article
Three novel species and new records of Kirschsteiniothelia (Kirschsteiniotheliales) from northern Thailand
expand article infoAntonio Roberto Gomes de Farias‡, Naghmeh Afshari§|, Veenavee S. Hittanadurage Silva‡, Johnny Louangphan‡, Omid Karimi‡, Saranyaphat Boonmee‡
‡ Mae Fah Luang University, Chiang Rai, Thailand
§ Mae Fah Luang University, Chiang Mai, Thailand
| Chiang Mai University, Chiang Rai, Thailand

Corresponding author: Antonio Roberto Gomes de Farias ( antonio.gom@mfu.ac.th )

Academic editor: Samantha C. Karunarathna
© 2024 Antonio Roberto Gomes de Farias, Naghmeh Afshari, Veenavee S. Hittanadurage Silva, Johnny Louangphan, Omid Karimi, Saranyaphat Boonmee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: de Farias ARG, Afshari N, Silva VSH, Louangphan J, Karimi O, Boonmee S (2024) Three novel species and new records of Kirschsteiniothelia (Kirschsteiniotheliales) from northern Thailand. MycoKeys 101: 347-370. https://doi.org/10.3897/mycokeys.101.115286
Open Access

Abstract

Kirschsteiniothelia (Kirschsteiniotheliales, Pleosporomycetidae) includes 39 saprobic species recorded from dead or decaying wood in terrestrial and freshwater habitats. This study focuses on exploring Kirschsteiniothelia diversity in woody litter in Thailand. Wood samples were collected from forest areas in Chiang Rai and Chiang Mai Provinces in Thailand and examined for fungal fructifications. Fungal isolates were obtained and their morphological and sequence data were characterised. Micromorphology associated with multilocus phylogeny of ITS, LSU and SSU sequence data identified three isolates as novel species (Kirschsteiniothelia inthanonensis, K. saprophytica and K. zizyphifolii) besides new host records for K. tectonae and K. xishuangbannaensis. The placement of the new taxa and records are supported by morphological illustrations, descriptions and molecular phylogenies and the implications of these findings are discussed. Our findings provide information for understanding Kirschsteiniothelia diversity and ecology.

Key words

Multilocus phylogeny, new host records, saprobic fungi, three new species, woody litter

Introduction

Since its introduction by Hawksworth (1985), the taxonomic placement of Kirschsteiniothelia (Kirschsteiniotheliaceae, Pleosporales, Pleosporomycetidae) has undergone several revisions. It was introduced in Pleosporaceae, with Kirschsteiniothelia aethiops as the type species. However, Barr (1993) moved it to Pleomassariaceae based on morphology and, based on molecular phylogenetic analyses, Schoch et al. (2006) demonstrated that K. aethiops does not belong to Pleosporaceae and should be placed in a new family. Kirschsteiniotheliaceae was established by Boonmee et al. (2012) to accommodate the holomorphic genus Kirschsteiniothelia. This was due to the fact that K. elaterascus and K. maritima clustered into Morosphaeriaceae and Mytilinidiales, respectively (Schoch et al. 2009; Suetrong et al. 2009; Boonmee et al. 2012). Later, Hernández-Restrepo et al. (2017) assigned it to the newly-proposed order Kirschsteiniotheliales (Dothideomycetes) due to its phylogenetic significance. Boonmee et al. (2012) also synonymised Dendryphiopsis atra under K. atra (Corda) D. Hawksw. due to their phylogenetic and asexual morph similarity (Boonmee et al. 2012; Schoch et al. 2009). The placement of Kirschsteiniothelia in the latest Outline of fungi and fungus-like taxa (Wijayawardene et al. 2022) is Kirschsteiniotheliaceae, Kirschsteiniotheliales, Dothideomycetes order incertae sedis, Dothideomycetes, Ascomycota.

Kirschsteiniothelia sexual morphs essentially have superficial to semi-immersed, subglobose to globose, dark brown to black ascomata; cylindrical clavate, bitunicate, 8-spored asci; and brown to dark brown, ellipsoidal, septate ascospores with or without a mucilaginous sheath (Hawksworth 1985; Boonmee et al. 2012; Hyde et al. 2013). However, its asexual morphs include dendryphiopsis-like and sporidesmium-like structures, with Dendryphiopsis taxa confirmed to be linked to Kirschsteiniothelia, based on morphology and molecular evidence (Schoch et al. 2009; Boonmee et al. 2012).

Kirschsteiniothelia species are mostly saprobes on dead or decaying wood in freshwater and terrestrial habitats (Boonmee et al. 2012; Hyde et al. 2013; Su et al. 2016; Mehrabi et al. 2017; Bao et al. 2018; Dong et al. 2020; Sun et al. 2021; Liu et al. 2023). These taxa play a crucial role in nutrient cycling and decomposition processes, contributing to the breakdown of organic matter in their respective ecosystems (Bucher et al. 2004). Their ability to colonise wood in freshwater habitats further emphasises their ecological significance (Su et al. 2016). In addition, Nishi et al. (2018) reported Kirschsteiniothelia associated with ankle bursitis in a Japanese patient and Guegan et al. (2021) with foot chromoblastomycosis in an immunosuppressed patient. Besides, Poch et al. (1992) discovered new compounds in Kirschsteiniothelia species, including kirschsteinin, which showed antimicrobial activity and Bugni and Ireland (2004) reported antibacterial activity from K. maritima.

This study focuses on exploring Kirschsteiniothelia diversity in woody litter in Thailand. We introduce three new species viz. K. inthanonensis, K. saprophytica and K. zizyphifolii, along with two new host records of Kirschsteiniothelia, based on a morpho-molecular approach, expanding our knowledge of the diversity in Pleosporomycetidae.

Material and methods

Sample collection, fungal isolation and microscopic characterisation

Wood litter samples were collected from forest areas in Chiang Rai and Chiang Mai, Thailand. Morphological studies were performed following the methods described by Senanayake et al. (2020). The fungal structures were examined using a Leica EZ4 stereomicroscope. The micro-morphological features were observed and photographed using a Nikon ECLIPSE Ni compound microscope with a Canon 600 D digital camera. The Tarosoft Image Frame Work programme was used to measure specimen structures, and photo plates were prepared using the open-source Inkscape v.1.3 (https://inkscape.org/).

Pure cultures were obtained through single spore isolation on Difco potato dextrose agar (PDA) using the spore suspension method (Choi et al. 1999). Germinating spores were transferred to a new PDA plate and incubated at room temperature for seven days. Ex-type pure living cultures were deposited in the Mae Fah Luang University Culture Collection (MFLUCC) and herbarium material was deposited in the Mae Fah Luang University Fungarium (MFLU), Chiang Rai, Thailand. Faces of fungi numbers (FoF) (Jayasiri et al. 2015) and Index Fungorum numbers (Index Fungorum 2023) were obtained as instructed and the data were uploaded to the Greater Mekong Subregion in the GMS database (Chaiwan et al. 2021).

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from fresh mycelium scrapings using the EE.Z.N.A. Tissue DNA Kit from Omega Bio-tek, Inc., following the manufacturer’s instructions. PCR amplifications were performed in a 50 μl reaction volume containing 10× PCR Master Mix, forward and reverse primers, DNA template and double sterilised H2O. Amplified DNA of the ITS, LSU and SSU were obtained through a polymerase chain reaction (PCR) using the pairs of primers ITS4/ITS5 (White et al. 1990), LROR/LR5 (Vilgalys and Hester 1990) and NS1/NS4 (White et al. 1990), correspondingly. The quality of the PCR products was visualised on a 1% agarose gel and sequenced by Biogenomed Co., Ltd (South Korea).

Alignments and phylogenetic analyses

The reads were assembled using the Staden Package (Staden et al. 2003) and compared against the NCBI non-redundant GenBank database (Sayers et al. 2020) and related reference sequences downloaded (Table 1). Except for concatenation and visualisation, all the steps of phylogenetic analysis were conducted in a Windows Subsystem for Linux (Microsoft, USA). The individual datasets were aligned using MAFFT with the --auto flag and automatically trimmed using TrimAl v.1.3 with the -gt (0.3) option (Capella-Gutierrez et al. 2009). The best-fit model was selected using ModelTest-NG v.0.1.7 with the --template mrbayes option for DNA 3 schemes matrices (Darriba et al. 2020). The alignments were concatenated using SequenceMatrix and subjected to Maximum Likelihood (ML) and Bayesian Inference (BI) analyses.

Table 1.
Download as
CSV
XLSX

Names, strain numbers, and corresponding GenBank accession numbers of Kirschsteiniotheliales taxa used in the phylogenetic analyses.

Taxa Strains Accession numbers
ITS LSU SSU
Acrospermum adeanum M133 EU940180 EU940104 EU940031
Acrospermum compressum M151 EU940161 EU940084 EU940012
Acrospermum gramineum M152 EU940162 EU940085 EU940013
Aliquandostipite crystallinus R 76–1 – EF175651 EF175630
Aliquandostipite khaoyaiensis CBS 118232T – GU301796 –
Anisomeridium ubianum MPN94 – GU327709 JN887379
Dyfrolomyces rhizophorae JK5456A – GU479799 GU479766
Dyfrolomyces tiomanensis NTOU3636 – KC692156 KC692155
Flavobathelium epiphyllum MPN67 – GU327717 JN887382
Halokirschsteiniothelia maritima CBS 221.60 – AY849943 AF053726
Helicomyces roseus CBS 283.51 AY916464 AY856881 AY856928
MFLUCC 15–0343 KY320523 KY320540 –
Homortomyces combreti CPC 19808T JX517281 JX517291 –
Homortomyces tamaricis MFLUCC 13–0280 KU752184 KU561874 KU870905
MFLUCC 14–0167 KU934190 KU561875 –
MFLUCC 13–0441T NR_155161 NG_059495 –
Jahnula bipileata F49–1 T JN942353 EF175657 EF175635
Jahnula sangamonensis A402–1B JN942349 EF175661 EF175639
Jahnula seychellensis SS 2113.2 – EF175664 EF175643
Kirschsteiniothelia acutispora MFLU 21–0127T OP120780 ON980758 ON980754
Kirschsteiniothelia aquatica MFLUCC 16–1685T MH182587 MH182594 MH182618
Kirschsteiniothelia arasbaranica IRAN 2509C KX621986 KX621987 KX621988
IRAN 2508CT KX621983 KX621984 KX621985
Kirschsteiniothelia atra DEN MG602687 – –
CBS 109.53 – AY016361 AY016344
MFLUCC 16–1104 MH182583 MH182589 MH182615
S–783 MH182586 MH182595 MH182617
MFLUCC 15–0424 KU500571 KU500578 KU500585
Kirschsteiniothelia cangshanensis GZCC19–0515 – MW133829 MW134609
MFLUCC 16–1350T MH182584 MH182592 –
MFLU 23–0358T OR575473 OR575474 OR575475
Kirschsteiniothelia crustaceum MFLU 21–0129T MW851849 MW851854 –
Kirschsteiniothelia dushanensis GZCC 19–0415 OP377845 MW133830 MW134610
Kirschsteiniothelia ebriosa CBS H–23379 – LT985885 –
Kirschsteiniothelia emarceis MFLU 10–0037T NR_138375 NG_059454 –
Kirschsteiniothelia extensum MFLU 21–0130T MW851850 MW851855 –
Kirschsteiniothelia fluminicola MFLUCC 16–1263T MH182582 MH182588 –
Kirschsteiniothelia inthanonensis MFLUCC 23–0277T OR762773 OR762781 OR764784
Kirschsteiniothelia lignicola MFLUCC 10–0036T HQ441567 HQ441568 HQ441569
Kirschsteiniothelia nabanheensis HJAUP C2006 OQ023274 OQ023275 OQ023037
HJAUP C2004T OQ023197 OQ023273 OQ023038
Kirschsteiniothelia phoenicis MFLU 18–0153 NR_158532 NG_064508 –
MFLUCC 18–0216T MG859978 MG860484 MG859979
Kirschsteiniothelia puerensis ZHKUCC 22–0272 OP450978 OP451018 OP451021
ZHKUCC 22–0271T OP450977 OP451017 OP451020
Kirschsteiniothelia rostrata MFLUCC 15–0619T KY697280 KY697276 KY697278
Kirschsteiniothelia septemseptatum MFLU 21–0126T OP120779 ON980757 ON980752
Kirschsteiniothelia saprophytica MFLUCC 23–0275 T OR762774 OR762783 –
MFLUCC 23–0276 OR762775 OR762782 –
Kirschsteiniothelia spatiosum MFLU 21–0128T – OP077294 ON980753
Kirschsteiniothelia submersa S–481 – MH182591 MH182616
S–601 MH182585 MH182593 –
MFLUCC 15–0427T KU500570 KU500577 KU500584
Kirschsteiniothelia tectonae MFLUCC 12–0050 KU144916 KU764707 –
MFLUCC 13–0470 KU144924 – –
Kirschsteiniothelia tectonae MFLUCC 23–0271 OR762771 OR762779 OR764782
MFLUCC 23–0272 OR762772 OR762780 OR764783
Kirschsteiniothelia thailandica MFLUCC 20–0116T MT985633 MT984443 MT984280
Kirschsteiniothelia thujina JF13210 KM982716 KM982718 KM982717
Kirschsteiniothelia vinigena CBS H–23378T – NG_075229 –
Kirschsteiniothelia xishuangbannaensis ZHKUCC 22–0221 OP289563 OP289565 OP303182
ZHKUCC 22–0220T OP289566 OP289564 OP303181
Kirschsteiniothelia xishuangbannaensis MFLUCC 23–0273 OR762770 OR762778 OR764781
MFLUCC 23–0274 OR762769 OR762777 OR764780
Kirschsteiniothelia zizyphifolii MFLUCC 23–027T OR762768 OR762776 OR764779
Megalotremis verrucosa MPN104 – GU327718 JN887383
Phyllobathelium anomalum MPN 242 – GU327722 JN887386
Stemphylium vesicarium CBS 191.86 MH861935 GU238160 GU238232
MFLUCC 14–0920 KY659560 KY659563 KY659567
Tubeufia helicomyces CBS 271.52 AY916461 AY856887 AY856933
Tubeufia javanica MFLUCC 12–0545T KJ880034 KJ880036 KJ880035
Acrospermum adeanum M133 EU940180 EU940104 EU940031
Acrospermum compressum M151 EU940161 EU940084 EU940012

The newly-generated sequences are indicated in bold. “T” refers to holotype or ex-type strains and “–” shows unavailable data in GenBank.

Maximum Likelihood (ML) trees were generated using RAxML-HPC2 on XSEDE (8.2.8) (Stamatakis 2014) in the CIPRES Science Gateway platform (Miller et al. 2010), using 1,000 bootstraps replications and applying a partitioned model of evolution calculated by ModelTest-NG. Bayesian Inference was performed using MrBayes (Ronquist et al. 2012), with four simultaneous Markov Chain Monte Carlo (MCMC) chains and four runs for 3,000,000 million generations, sampling trees every 300th generation. The first 25% of trees were discarded as burn-in and posterior probabilities (PP) were calculated from the remaining trees. The consensus phylograms were visualised using FigTree (Rambaut 2012) and edited using the open-source Inkscape v.1.3 (https://inkscape.org/).

Results

Phylogenetic analyses

The concatenated nucleotide alignment of the ITS, LSU and SSU datasets comprised 69 Kirschsteiniotheliales strains, including the outgroups (S. vesicarium MFLUCC 14–0920 and CBS191.86) and included 2,640 sites (ITS = 1–561; LSU = 562–1596; SSU = 1597–2640), of which 1,550 comprised of distinct alignment patterns (ITS = 427, LSU = 668 and SSU = 455), with of 32.01% undetermined characters or gaps. The final GAMMA-based score of the best tree was -24775.722822. Maximum Likelihood phylogeny and Bayesian analyses of single- and multi-loci had similar topologies and are combined in Fig. 1. Parameters for the models of each amplicon were described in Table 2. The Bayesian analysis tracer of the combined runs checked at six million generations had an effective sampling size for all the parameters higher than 3,000 and convergence diagnostic (PSRF = Potential Scale Reduction Factor; Gelman and Rubin (1992) of 1.0. The run resulted in 10,001 trees, of which 7,501 were sampled after 25% of the trees were discarded as burn-in. The alignment contained 1,802 unique sites (ITS = 427, LSU = 782, SSU = 593). The ML and BI analyses showed similar tree topologies.

Table 2.
Download as
CSV
XLSX

Maximum Likelihood indices of Kirschsteiniothelia tree.

Parameters ITS LSU SSU
Evolutionary model GTR+I+G4 GTR+G4 GTR+I+G4
Gamma distribution shape parameter α 0.267050 0.557118 0.228478
Estimated base frequencies
A 0.199482 0.235780 0.260410
C 0.306708 0.238788 0.213687
G 0.279166 0.322010 0.267932
T 0.214644 0.203422 0.257971
Substitution rates
AC 1.269939 0.865025 1.305091
AG 2.734587 2.259149 2.368982
AT 1.504952 1.054098 0.620257
CG 1.112253 0.931891 0.757010
CT 3.835090 5.793263 8.684577
GT 1.000000 1.000000 1.000000
Figure 1. 

Maximum Likelihood phylogenetic tree generated from ITS, LSU and SSU sequence data for selected Kirschsteiniotheliales and related Dothideomycetes orders. The tree is rooted with Stemphylium vesicarium (CBS 191.86 and MFLUCC 14–0920). Newly-generated sequences are in blue and new species are in bold. Holotype and ex-type strains are symbolic by “T”. Maximum Likelihood bootstrap (MLBS) values ≥ 70% and Bayesian posterior probabilities (BYPP) ≥ 0.95 are shown at the nodes.

Four strains (MFLUCC 23–0277, MFLUCC 23–0270 and MFLUCC 23–0275 and MFLUCC 23–0276) clustered in three independent lineages (Fig. 1). MFLUCC 23–0277 clustered sister to K. septemseptata (MFLU 21–0126) with 100% Maximum Likelihood bootstrap support (MLBS) and 1.00 Bayesian posterior probabilities (BYPP) support, while MFLUCC 23–0270 grouped as a sister of K. emarceis MFLU 10–0037, but with only 16% MLBS, 0.63 BYPP support, while MFLUCC 23–0275 and MFLUCC 23–0276 clustered with K. lignicola MFLUCC 10–0036 with 84% MLBS, 1.00 BYPP support. The other strains clustered with the known species K. tectonae (MFLUCC 23–0272 and MFLUCC 23–0271) and K. xishuangbannaensis (MFLUCC 23–0273 and MFLUCC 23–0274) with 71% MLBS and 72 MLBS/0.98 BYPP support, respectively. Based on the result of morphological evidence (Figs 2–7), three new species (K. zizyphifolii, K. inthanonensis and K. saprophytica) are proposed, along with the two new host records for K. xishuangbannaensis and K. tectonae.

Figure 2. 

Kirschsteiniothelia inthanonensis (MFLU 23–0420, holotype) a, b colonies on the host c, d conidiophores and conidia e regeneration of conidiophores f–j conidiogenous cells and conidia l–o conidia p germinating conidium q, r colony on PDA (front and reverse). Scale bars: 500 μm (b–d); 50 μm (e–j); 20 μm (k–p).

Taxonomy

Kirschsteiniothelia inthanonensis J. Louangphan & Gomes de Farias, sp. nov.

Index Fungorum number: IF901384
Facesoffungi Number: FoF14982
Fig. 3

Etymology

The name refers to the location “Doi Inthanon” where the holotype was collected.

Holotype

MFLU 23–0420

Description

Saprobic on decaying wood. Sexual morph: Not observed. Asexual morph: Hyphomycetes. Colonies on the host substrate are superficial, effuse, long hairy, fascicular, scattered, dark brown to black. Mycelium superficial and immersed, composed of branched, septate, pale brown and smooth hyphae. Conidiophores 611–1549 × 2.5–6.6 μm (x̄ = 1070 × 4.1 μm, n = 20), macronematous, synnematous, compact fasciculate, straight to flexuous, brown to dark brown, branched at the apex, multi-septate, thick and smooth-walled. Conidiogenous cells 15–45 × 6.7–10.4 μm (x̄ = 24.3 × 8 μm, n = 20), monotretic to polytretic, calyciform, integrated, discrete, terminal, darkened at the apex, proliferating portion, brown, 2–4 septate. Conidia 24–230 × 5.7–14.3 μm (x̄ = 101 × 9 μm, n = 15), acrogenous, solitary, obclavate, rostrate, straight or curved, truncate at base, grey to brown, pale at apex, partly tapering towards and rounded at the apex, 2–10– euseptate, smooth-walled.

Figure 3. 

Kirschsteiniothelia saprophytica (MFLU 23–0419, holotype) a host b, c appearance of ascomata on host surface d paraphyses e–g asci h–k ascospores l, m culture on PDA (front and reverse). Scale bars: 20 µm (d–g); 10 µm (h–k).

Culture characteristics

Conidia germinated on PDA within 48 hours. Germ tubes germinated from end cell. Colony, reaching 30–35 mm diam. after one month at room temperature, circular form, flat, undulate edges, dense velvety surface, dark green on the surface, white mycelium on the tip, dark in reverse with dark green margin.

Material examined

Thailand, Chiang Mai, Chom Thong, Doi Inthanon National Park, on twigs of Quercus oleoides, 30 November 2022, Veenavee Silva, DIFWS5-01 (MFLU 23–0420, holotype), ex-type living culture MFLUCC 23–0277.

Notes

Kirschsteiniothelia inthanonensis (MFLUCC 23–0277) resembles K. septemseptatum and K. nabanheensis in having septate, cylindrical conidiophores with branches near apex, integrated, terminal conidiogenous cells and solitary, obclavate, septate conidia without mucilaginous sheaths. However, K. inthanonensis MFLUCC 23–0277 has longer and smaller conidiophores than K. septemseptatum and K. nabanheensis (611–1549 μm vs. 250–580 μm and 320–588 μm) and (2.5–6.6 μm vs. 6.5–14.5 μm and 8–12 µm), respectively and elongated conidia (Jayawardena et al. 2022; Liu et al. 2023). In addition, our phylogenetic analyses show that K. inthanonensis forms an independent branch with 100% MLBS and 1.00 BYPP support. BLASTn base pair comparisons between K. inthanonensis (MFLUCC 23–0277) and K. septemseptatum (MFLU 21–0126) show 95% similarity of ITS (479/504, 6 gaps), 99% similarity of LSU (844/853, no gaps) and 99% similarity of SSU (787/789, 2 gaps). Kirschsteiniothelia nabanheensis (HJAUP C2004) shows 94% similarity of ITS (483/513, 7 gaps), 99% similarity of LSU (540/547, no gaps) and 98% similarity of SSU (864/883, no gaps). Based on these data, we introduce K. inthanonensis as a new species.

Kirschsteiniothelia saprophytica O. Karimi, V. Silva & Gomes de Farias, sp. nov.

Index Fungorum number: IF561030
Facesoffungi Number: FoF14983
Figs 4, 5

Etymology

The species epithet refers to the saprobic life mode of the fungus.

Holotype

MFLU 23–0419

Description

Saprobic on dead wood of undetermined host. Sexual morph: Ascomata 146.7–72.26 µm diam., superficial, solitary, globose to subglobose, dark brown to black. Pseudoparaphyses 1.2–2.7 µm wide (x̄ = 1.9, n = 20), hyaline, branched, filiform, abounded. Asci 68–125 × 18–23 µm (x̄ = 101 × 20 µm, n = 10), bitunicate, 8-spored, cylindrical-claviform, sessile or short pedicellate. Ascospores 13–25 (–40) × 7–11 (–14) µm (x̄= 24 × 9.8 µm, n = 25), ellipsoid, upper cell broader than lower cell, pale brown to dark brown, 1-septate, guttulate, smooth-walled. Asexual morph: Hyphomycetous. Colonies on host gregarious. Conidiophores 90–216 × 8–12 µm (x̄ = 165 × 10.6 µm, n = 10), macronematous, mononematous, cylindrical, straight to flexuous, branched, dark brown, multi-septate, constricted at the septa. Conidiogenous cells 6.7–35 × 5–15 µm (x̄ = 17 × 10 µm, n = 10), holoblastic, monoblastic, terminal, cylindrical, brown to dark brown. Conidia 36–69 × 19–35 µm (x̄ = 55 × 27 µm, n = 15), cylindrical rounded at ends, 2–3-septa, dark brown to black, smooth-walled.

Figure 4. 

Kirschsteiniothelia saprophytica (MFLUCC 23–0276) a host b colonies on the host, associated with asexual morph c conidiophore with conidiogenous cell and conidiospore d, e conidiophore (e – from the culture) f–j conidiospore from culture k germinating spore m, n culture on PDA (front and reverse). Scale bars: 50 µm (c–e); 20 µm (f–k).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies growing on PDA 16.8 mm diam. at room temperature after 38 days and on MEA 24 mm after 12 days. Mycelium on PDA superficial to immerse, dark olivaceous to dark brown on the top, reverse dark brown to black. Conidia germinating on PDA within 48 h. Colonies growing on PDA 17 mm diam. at room temperature after 16 days. Mycelium superficial to immerse, dark olivaceous to dark brown on the top, reverse dark brown to black.

Figure 5. 

Kirschsteiniothelia zizyphifolii (MFLU 23–0415, holotype) a colonies on wood b–d, g conidiophores and conidiogenous cells e, f conidiophores with conidia h–o conidia p germinated conidium q, r cultures on PDA from the surface and reverse. Scale bars: 200 μm (a); 100 μm (b–d); 50 μm (e, f); 20 μm (g–p).

Material examined

Thailand, Mae Fah Luang University, Chiang Rai, on dead wood of unidentified host, 20 October 2022, V. Silva, V020 (MFLU 23–0419, holotype), ex-type living culture MFLUCC 23–0275 and MFLUCC 23–0276.

Notes

Our collection (MFLUCC 23–0275) shares similar general characteristics to the type strain Kirschsteiniothelia lignicola (MFLUCC 10–0105), such as spherical and dark pigmented ascomata, cylindrical to claviform asci, ellipsoidal septate ascospores and cylindrical with brown conidia (Boonmee et al. 2012). However, our collection differs from K. lignicola in having shorter asci (68–125 × 18–23 vs. 107–163.3 × 19–28.5 µm), with shorter pedicels (5–6 vs. 14.5–24 µm), shorter conidiophores (90–216 × 8–12 vs. 287–406 × 11–13 µm) and 2–3 transverse septa. Phylogenetically, our isolate clustered with K. lignicola with 84% MLBS, 1.00 BYPP. The pairwise base comparisons of the ITS and LSU sequences between K. saprophytica and K. lignicola showed identities of 93.08% (484/520, 10 gaps) and 91.18% (806/884, 4 gaps), respectively. Based on these differences, we introduce K. saprophytica as a new species.

Kirschsteiniothelia zizyphifolii N. Afshari & Gomes de Farias, sp. nov.

Index Fungorum number: IF901382
Facesoffungi Number: FoF14981
Fig. 2

Etymology

“zizyphifolii” refers to the host species on which the fungus was found.

Holotype

MFLU 23–0415

Description

Saprobic on Nayariophyton zizyphifolium (Malvaceae) woody litter in terrestrial habitat. Sexual morph: Not observed. Asexual morph: Hyphomycetes. Colonies on the substratum are superficial, effuse, dark brown to black and hairy. Mycelia superficial, composed of septate, branched, smooth-walled, dark brown hyphae. Conidiophores 287–444.5 × 10.3 –17 (–19.7) μm (x̄ = 358.5 × 13.4 μm, n = 15), macronematous, mononematous, erect, with several short branches near the apex, irregular, solitary, cylindrical, flexuous, sometimes slightly straight, dark brown to black, paler towards the apex, septate, smooth-walled. Conidiogenous cells 11–20.4 × 5.8–10.6 μm (x̄ = 14.6 × 7.6 μm, n = 25), tretic, occasionally percurrent, integrated, terminal or intercalary, cylindrical or doliiform, brown, smooth-walled. Conidia (29.5–) 37.6–46.5 × 13.5–19 μm (x̄ = 43 × 16 μm, n = 20), acrogenous, solitary, cylindrical to rarely clavate, rounded at the apex, straight or moderately curved, brown dark to brown, 2–3-septate, constricted and pigmented at the septa, smooth-walled.

Culture characteristics

Ascospores germinating on PDA within 24 hours, reaching up to 30 mm diam. after one week at room temperature. Germ tubes germinated from both end cells. Colony dense, circular, velvety, narrow towards the edge, from front, grey at centre, black towards edge, from reverse, black.

Material examined

Thailand, Chiang Rai, Mae Fa Luang, Doi Tung Forest, on dead wood of Nayariophyton zizyphifolium, 26 March 2022, N. Afshari 1C1T2R4b (MFLU23–0415, holotype), ex-type living culture MFLUCC 23–0270.

Notes

Kirschsteiniothelia zizyphifolii (MFLUCC 23–0270) resembles K. lignicola (MFLUCC 10–0036) and K. emarceis (MFLU 10–0037) in having erect and branched conidiophores with apical dark brown conidia. However, it differs from K. lignicola in the sizes of conidiophores and conidia. Furthermore, BLASTn search of ITS and LSU sequences showed that K. zizyphifolii was closest to K. emarceis with similarity values of 90% (472/522, 12 gaps) and 84% (708/842, 22 gaps), respectively. Furthermore, our isolate (MFLUCC 23–0270) was close to K. lignicola (MFLUCC 10–0036) with similarity values of 89% (ITS = 474/532, 19 gaps), 99% (LSU = 844/853, 2 gaps) and 99% (SSU = 643/648, 2 gaps). Based on these phylogenetic data, we introduce K. zizyphifolii as a new species.

Kirschsteiniothelia tectonae Doilom, Bhat & K.D. Hyde, 2016

Index Fungorum number: IF551992
Facesoffungi Number: FoF01883
Fig. 6

Description

Saprobic on Microcos paniculata (Malvaceae) woody litter in terrestrial habitats. Sexual morph: Not observed. Asexual morph: Hyphomycetes. Colonies on the substrate, hairy, superficial, dark brown, scattered, partially grouped. Conidiophores 59–90 × 8.6–12 μm (x̄ = 75 × 10.7 μm, n = 10), superficial, simple, macronematous, mononematous, cylindrical, straight to slightly curved, branched or unbranched, septate, dark brown to black. Conidiogenous cells 7–9.4 × 6–7.3 μm (x̄ = 8 × 6.7 μm, n = 5), monoblastic, determinate, integrated, terminal. Conidia 62.5– 133 × 11 – 18.5(–21) μm (x̄ = 94 × 16 μm, n = 30), cylindrical-obclavate, elongate, straight to slightly curved, rounded being slightly paler at the apex, obconically truncate at the base, 7–12–septa, olivaceous green to brown, smooth–walled.

Figure 6. 

Kirschsteiniothelia tectonae (MFLUCC 23–0271, new record) a, b colonies on wood c–e conidiophores with conidia and conidiogenous cells f–k conidia l germinated conidium m, n culture on PDA (front and reverse). Scale bars: 100 μm (a, b); 50 μm (c–e, l); 20 μm (f–k).

Culture characteristics

Conidia germinating on PDA within 24 hours, reaching up to 15–20 mm diam. after one week at room temperature. Germ tubes generated from basal cells. Colony on PDA, dense, circular, flat or effuse, velvety, from front brown at the centre and black at the edge, from reverse, dark brown.

Material examined

Thailand, Chiang Rai, Mae Fa Luang, Doi Tung, on dead wood of Microcos paniculata, 6 June 2022, N. Afshari 3C2T3R5 (MFLU 23–0416), living culture MFLUCC 23–0272. On dead wood of Dalbergia cana, 3 March 2022, N. Afshari 4C1T2R3 (MFLU 23–0417), living culture MFLUCC 23–0272.

Known distribution

Thailand (Li et al. 2016; this study)

Known hosts

Tectona grandis (Li et al. 2016), Microcos paniculata and Dipterocarpus alatus (this study)

Kirschsteiniothelia xishuangbannaensis R.F. Xu & Tibpromma

Index Fungorum number: IF559433
Facesoffungi Number: FoF12758
Fig. 7

Description

Saprobic on Microcos paniculata (Malvaceae) woody litter in terrestrial habitats. Sexual morph: Not observed. Asexual morph: Hyphomycetes. Colonies effuse on the substrate, hairy, solitary or scattered, dark brown. Conidiophores 135–178 × 7.7–11 μm (x̄ = 151 × 9 μm, n = 10), macronematous, straight to curved, solitary, brown, slightly larger at base, narrowing towards apex, septate. Conidiogenous cells 14.4–27.4 × 7.8–11 μm (x̄ = 22 × 10 μm, n = 10), holoblastic, monoblastic, integrated, smooth, terminal, determinate, cylindrical or lageniform, brown. Conidia 70–141 × 14.5–19 μm (x̄ = 100 × 17 μm, n = 20), solitary, acrogenous, obclavate, rostrate, straight or slightly curved, truncate at the base, olivaceous green to brown, subhyaline at the apex, 5–10-septate, large guttulate.

Figure 7. 

Kirschsteiniothelia xishuangbannaensis (MFLUCC 23–0273, new record) a, b colonies on wood c, d conidiophores and conidiogenous cells e conidiophore with conidium f–o conidia p germinated conidium q, r culture on PDA (front and reverse). Scale bars: 200 μm (a); 100 μm (b); 50 μm (c, d, p); 30 μm (e); 20 μm (f–o).

Culture characteristics

Conidia germinating on PDA within 24 hours reaching up to 2 cm diam. after one week at room temperature. Germ tubes generated from both end cells. Colony on PDA, dense, circular, flat or effuse, velvety, from front, brown at the centre and dark brown at edge, from reverse, black to pale brown radiating.

Material examined

Thailand, Chiang Rai, Mae Fa Luang, Doi Tung, on dead wood of Microcos paniculata, 6 June 2022, N. Afshari 3C2T1R1, living culture MFLUCC 23–0273. On dead wood of Dipterocarpus alatus, 27 September 2022, N. Afshari 2C3T1R3c (MFLU 23–0418), living culture MFLUCC 23–0274.

Known distribution

China (Xu et al. 2023), Thailand (this study).

Known hosts

Hevea brasiliensis (Xu et al. 2023), Microcos paniculata and Dalbergia cana (this study).

Discussion

This study introduces three new species and new host records of Kirschsteiniothelia from dead wood from Chiang Rai Province, Thailand, based on morphological and molecular analyses (Figs 1–7). Kirschsteiniothelia species have been found almost worldwide, including in the United States of America (Hawksworth 1985; Hyde 1997; Wang et al. 2004; Su et al. 2016), Iran (Mehrabi et al. 2017), Switzerland (Hawksworth 1985; Wang et al. 2004), Thailand (Boonmee et al. 2012; Li et al. 2016; Bao et al. 2018; Hyde et al. 2018; Sun et al. 2021; Jayawardena et al. 2022), South Africa (Marincowitz et al. 2008), China (Chen et al. 2006; Su et al. 2016; Bao et al. 2018; Liu et al. 2023; Yang et al. 2023; Xu et al. 2023), Canada (Hawksworth 1985), Italy (Wang et al. 2004), Spain (Rodríguez-Andrade et al. 2019) and India (Bao et al. 2018). Most of the species (K. acutispora, K. chiangmaiensis, K. crustacea, K. emarceis, K. extensa, K. lignicola, K. phoenicis, K. rostrata, K. septemseptata, K. spatiosa, K. tectonae and K. thailandica) have been reported from Thailand (Boonmee et al. 2012; Li et al. 2016; Bao et al. 2018; Hyde et al. 2018; Sun et al. 2021; Jayawardena et al. 2022), representing more than 25% of the species in this genus. Our results expand the knowledge of the diversity of this genus, especially in Thailand.

This genus is also prone to be highly speciose, given the recent introduction of ten new species (Jayawardena et al. 2022; Hyde et al. 2023; Liu et al. 2023; Louangphan et al. 2023 (under review); Xu et al. 2023). With the introductions of the present study (K. inthanonensis, K. saprophytica, K. paniculata and K. zizyphifolii), 32.5% of the species will have been introduced within two years, mainly as saprobes in woody litter. Besides, most Kirschsteiniothelia species have been reported from terrestrial environments, with only a few (K. cangshanensis, K. fluminicola and K. rostrata) reported from freshwater habitats (Bao et al. 2018). Their ecological significance also relies on their ability to infect humans (Nishi et al. 2018; Guegan et al. 2021). This demonstrates the potential for further discoveries on the diversity and lifestyles within Kirschsteiniothelia. Thus, exploring its diversity, especially in woody litter in protected environments and other tropical areas, will reveal the vast diversity within Kirschsteiniotheliaceae. For example, frequent incursions into fungal diversity have established Thailand as a hotspot for its diversity (Hyde et al. 2018).

Furthermore, Kirschsteiniothelia species appear to not have host specificity, as from our results, the same species were found associated with different hosts: K. xishuangbannaensis, previously reported from dead branches of Hevea brasiliensis (Xu et al. 2023), was recorded from Microcos paniculata (MFLUCC 23–0273) and Dipterocarpus alatus (MFLUCC 23–0274); K. paniculata was isolated from Microcos paniculata (MFLUCC 23–0271) and Dalbergia cana (MFLUCC 23–0272). In this regard, the host-specificity or host-recurrence of saprobic fungi has been discussed over the last two decades (Hooper et al. 2000; Zhou and Hyde 2001; Santana et al. 2005; Kodsueb et al. 2008; Tennakoon et al. 2022). However, saprotrophs seem to be less host-specific when compared with other trophic modes (Zhou and Hyde 2001). This may be because different hosts have different chemical compositions, which may affect the fungi of a particular species (Hyde et al. 2007). This hypothesis suggests that woody litter may harbour many species yet to be discovered (Kodsueb et al. 2008).

A combined approach should be employed to resolve the taxonomic placement of new species in this genus. This approach should include at least molecular phylogeny and morphological characters (Chethana et al. 2021; Maharachchikumbura et al. 2021). It should also include the linking of sexual and asexual morphologies, which are important factors in the taxonomy of Ascomycota, as pleomorphism can bias the morphological characters (Maharachchikumbura et al. 2021). However, only a few of the 39 Kirschsteiniothelia species, specifically K. atra and K. recessa (Hawksworth 1985) and K. lignicola and K. emarceis (Boonmee et al. 2012), are known from both their sexual and asexual morphs.

The findings of this study underscore the importance of integrating multiple types of evidence for the identification and classification of fungal species and they demonstrate the potential for further discoveries within Kirschsteiniothelia. The discovery of new species and host records has significant implications for our understanding of the ecological roles and interactions of this genus. In particular, identifying new host records provides valuable insights into the host range and specificity of Kirschsteiniothelia species, which may help elucidate the mechanisms underlying these interactions. Further research is necessary to fully explore the ecological significance of these findings and determine the potential impacts of Kirschsteiniothelia species on their hosts and ecosystems.

Acknowledgements

Antonio R. Gomes de Farias thanks Thailand Science Research and Innovation (TSRI) and National Science Research and Innovation Fund (NSRF) (Fundamental Fund: Grant no. 662A16047) entitled “Biodiversity, ecology, and applications of plant litter-inhabiting fungi for waste degradation”. Saranyaphat Boonmee thanks the National Research Council of Thailand (NRCT: Project no. P-19–52624) project entitled “Comparison of diversity and biogeographical distribution of Ascomycetous fungi from two protected areas in Turkey and Thailand” under the Doi Inthanon National Park permission No.0402/2804. The authors would like to thank Dr. Shaun Pennycook for checking and suggesting Latin names of the new taxa, Martin van de Bult, Narong Apichai and the Doi Tung Development Project for sample collection (permission number 7700/17142 with the title “The diversity of saprobic fungi on selected hosts in forest northern Thailand”), Chiang Rai, Thailand.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was partially supported by the National Science and Technology Development Agency (NSTDA: Project No. P–19–52624), under the National Park Permission No. 0907.4/8218 and No. 0907.4/19647 and Thailand Science Research and Innovation (TSRI) and National Science Research and Innovation Fund (NSRF) Fundamental Fund grant (Grant no. 662A16047), entitled “Biodiversity, ecology and applications of plant litter-inhabiting fungi for waste degradation”.

Author contributions

Antonio Roberto Gomes de Farias: Conceptualization and design of the study, Funding acquisition, Writing – original draft; Naghmeh Afshari: Methodology, Writing – original draft; Veenavee S. Hittanadurage Silva: Methodology, Writing – original draft; Johnny Louangphan: Methodology, Writing – original draft; Omid Karimi: Writing – original draft; Saranyaphat Boonmee: Funding acquisition, Methodology, Supervision, Writing – revision.

Author ORCIDs

Antonio Roberto Gomes de Farias https://orcid.org/0000-0003-4768-1547

Veenavee S. Hittanadurage Silva https://orcid.org/0000-0001-8921-1370

Omid Karimi https://orcid.org/0000-0001-9652-2222

Saranyaphat Boonmee https://orcid.org/0000-0001-5202-2955

Data availability

All of the data that support the findings of this study are available in the main text.

References

  • Bao D, Luo Z, Liu J, Bhat D, Sarunya N, Li W, Su H, Hyde K (2018) Lignicolous freshwater fungi in China III: Three new species and a new record of Kirschsteiniothelia from northwestern Yunnan Province. Mycosphere: Journal of Fungal Biology 9(4): 755–768. https://doi.org/10.5943/mycosphere/9/4/4
  • Barr ME (1993) Notes on the Pleomassariaceae. Mycotaxon 49(1): 129–142.
  • Boonmee S, Ko Ko T, Chukeatirote E, Hyde K, Chen H, Cai L, Mckenzie E, Jones E, Kodsueb R, Bahkali A (2012) Two new Kirschsteiniothelia species with Dendryphiopsis anamorphs cluster in Kirschsteiniotheliaceae fam. nov. Mycologia 104(4): 698–714. https://doi.org/10.3852/11-089
  • Bucher VVC, Hyde KD, Pointing SB, Reddy CA (2004) Production of wood decay enzymes, loss of mass, and lignin solubilisation in wood by diverse tropical freshwater fungi. Microbial Ecology 15(3): 1–14. https://doi.org/10.1007/s00248-003-0132-x
  • Bugni TS, Ireland CM (2004) Marine-derived fungi: A chemically and biologically diverse group of microorganisms. Natural Product Reports 21(1): 143–163. https://doi.org/10.1039/b301926h
  • Capella-Gutierrez S, Silla-Martínez J, Gabaldón T (2009) TrimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25(15): 1972–1973. https://doi.org/10.1093/bioinformatics/btp348
  • Chaiwan N, Gomdola D, Wang S, Monkai J, Tibpromma S, Doilom M, Wanasinghe DN, Mortimer PE, Lumyong S, Hyde KD (2021) An online database providing updated information of microfungi in the Greater Mekong Subregion. Mycosphere: Journal of Fungal Biology 12(1): 1513–1526. https://gmsmicrofungi.org. https://doi.org/10.5943/mycosphere/12/1/19
  • Chen CY, Wang CL, Huang JW (2006) Two new species of Kirschsteiniothelia from Taiwan. Mycotaxon 98: 153–158.
  • Chethana KT, Manawasinghe IS, Hurdeal VG, Bhunjun CS, Appadoo MA, Gentekaki E, Raspé O, Promputtha I, Hyde KD (2021) What are fungal species and how to delineate them? Fungal Diversity 109(1): 1–25. https://doi.org/10.1007/s13225-021-00483-9
  • Choi YW, Hyde KD, Ho WH (1999) Single spore isolation of fungi. Fungal Diversity 3: 19–38.
  • Darriba D, Posada D, Kozlov AM, Stamatakis A, Morel B, Flouri T (2020) ModelTest-NG: A new and scalable tool for the selection of DNA and protein evolutionary models. Molecular Biology and Evolution 37(1): 291–294. https://doi.org/10.1093/molbev/msz189
  • Dong W, Wang B, Hyde KD, McKenzie EH, Raja HA, Tanaka K, Abdel-Wahab MA, Abdel-Aziz FA, Doilom M, Phookamsak R, Hongsanan S, Wanasinghe DN, Yu XD, Wang GN, Yang H, Yang J, Thambugala KM, Tian Q, Luo ZL, Yang JB, Miller AN, Fournier J, Boonmee S, Hu DM, Nalumpang S, Zhang H (2020) Freshwater Dothideomycetes. Fungal Diversity 105(1): 319–575. https://doi.org/10.1007/s13225-020-00463-5
  • Guegan H, Cailleaux M, Le Gall F, Robert-Gangneux F, Gangneux JP (2021) chromoblastomycosis due to a never-before-seen dematiaceous fungus in a kidney transplant patient. Microorganisms 9(10): e2139. https://doi.org/10.3390/microorganisms9102139
  • Hernández-Restrepo M, Gené J, Castañeda-Ruiz RF, Mena-Portales J, Crous PW, Guarro J (2017) Phylogeny of saprobic microfungi from Southern Europe. Studies in Mycology 86: 53–97. https://doi.org/10.1016/j.simyco.2017.05.002
  • Hooper DU, Bignell DE, Brown VK, Brussard L, Dangerfield JM, Wall DH, Wardle DA, Coleman DC, Giller KE, Lavelle P, Van Der Putten WH (2000) Interactions between aboveground and belowground biodiversity in terrestrial ecosystems: Patterns, mechanisms, and feedbacks: We assess the evidence for correlation between aboveground and belowground diversity and conclude that a variety of mechanisms could lead to positive, negative, or no relationship-depending on the strength and type of interactions among species. Bioscience 50(12): 1049–1061. https://doi.org/10.1641/0006-3568(2000)050[1049:IBAABB]2.0.CO;2
  • Hyde KD (1997) Ascomycetes described on Freycinetia. Sydowia 49: 1–20.
  • Hyde KD, Bussaban B, Paulus B, Crous PW, Lee S, Mckenzie EH, Photita W, Lumyong S (2007) Diversity of saprobic microfungi. Biodiversity and Conservation 16(1): 7–35. https://doi.org/10.1007/s10531-006-9119-5
  • Hyde KD, Jones EBG, Liu J-K, Ariyawansa H, Boehm E, Boonmee S, Braun U, Chomnunti P, Crous PW, Dai D-Q, Diederich P, Dissanayake A, Doilom M, Doveri F, Hongsanan S, Jayawardena R, Lawrey JD, Li Y-M, Liu Y-X, Lücking R, Monkai J, Muggia L, Nelsen MP, Pang KL, Phookamsak R, Senanayake IC, Shearer CA, Suetrong S, Tanaka K, Thambugala KM, Wijayawardene NN, Wikee S, Wu HX, Zhang Y, Aguirre-Hudson B, Alias SA, Aptroot A, Bahkali AH, Bezerra JL, Bhat DJ, Camporesi E, Chukeatirote E, Gueidan C, Hawksworth DL, Hirayama K, De Hoog S, Kang JC, Knudsen K, Li WJ, Li XH, Liu ZY, Mapook A, McKenzie EHC, Miller AN, Mortimer PE, Phillips AJL, Raja HA, Scheuer C, Schumm F, Taylor JE, Tian Q, Tibpromma S, Wanasinghe DN, Wang Y, Xu JC, Yacharoen S, Yan JY, Zhang M (2013) Families of Dothideomycetes. Fungal Diversity 63(1): 1–313. https://doi.org/10.1007/s13225-013-0263-4
  • Hyde KD, Chaiwan N, Norphanphoun C, Boonmee S, Camporesi E, Chethana KW, Dayarathne MC, De Silva NI, Dissanayake AJ, Ekanayaka AH, Hongsanan S, Huang SK, Jayasiri SC, Jayawardena R, Jiang HB, Karunarathna A, Lin CG, Liu JK, Liu NG, Lu YZ, Luo ZL, Maharachchimbura SSN, Manawasinghe IS, Pem D, Perera RH, Phukhamsakda C, Samarakoon MC, Senwanna C, Shang QJ, Tennakoon DS, Thambugala KM, Tibpromma S, Wanasinghe DN, Xiao YP, Yang J, Zeng XY, Zhang JF, Zhang SN, Bulgakov TS, Bhat DJ, Cheewangkoon R, Goh TK, Jones EBG, Kang JC, Jeewon R, Liu ZY, Lumyong S, Kuo CH, McKenzie EHC, Wen TC, Yan JY, Zhao Q (2018) Mycosphere notes 169–224. Mycosphere: Journal of Fungal Biology 9(2): 271–430. https://doi.org/10.5943/mycosphere/9/2/8
  • Jayasiri SC, Hyde KD, Ariyawansa HA, Bhat J, Buyck B, Cai L, Dai YC, Abd-Elsalam KA, Ertz D, Hidayat I, Jeewon R, Jones EBG, Bahkali AH, Karunarathna SS, Liu JK, Luangsa-ard JJ, Lumbsch HT, Maharachchikumbura SSN, McKenzie EHC, Moncalvo JM, Ghobad-Nejhad M, Nilsson H, Pang KL, Pereira OL, Phillips AJL, Raspé O, Rollins AW, Romero AI, Etayo J, Selçuk F, Stephenson SL, Suetrong S, Taylor JE, Tsui CKM, Vizzini A, Abdel-Wahab MA, Wen TC, Boonmee S, Dai DQ, Daranagama DA, Dissanayake AJ, Ekanayaka AH, Fryar SC, Hongsanan S, Jayawardena RS, Li WJ, Perera RH, Phookamsak R, de Silva NI, Thambugala KM, Tian Q, Wijayawardene NN, Zhao RL, Zhao Q, Kang JC, Promputtha I (2015) The Faces of Fungi database: Fungal names linked with morphology, phylogeny and human impacts. Fungal Diversity 74(1): 3–18. https://doi.org/10.1007/s13225-015-0351-8
  • Jayawardena RS, Hyde KD, Wang S, Sun YR, Suwannarach N, Sysouphanthong P, Abdel-Wahab MA, Abdel-Aziz FA, Abeywickrama PD, Abreu VP, Armand A, Aptroot A, Bao DF, Begerow D, Bellanger JM, Bezerra JDP, Bundhun D, Calabon MS, Cao T, Cantillo T, Carvalho JLVR, Chaiwan N, Chen CC, Courtecuisse R, Cui BK, Damm U, Denchev CM, Denchev TT, Deng CY, Devadatha B, de Silva NI, dos Santos LA, Dubey NK, Dumez S, Ferdinandez HS, Firmino AL, Gafforov Y, Gajanayake AJ, Gomdola D, Gunaseelan S, He S, Htet ZH, Kaliyaperumal M, Kemler M, Kezo K, Kularathnage ND, Leonardi M, Li JP, Liao CF, Liu S, Loizides M, Luangharn T, Ma J, Madrid H, Maharachchikumbura SSN, Manamgoda DS, Martín MP, Mekala N, Moreau PA, Mu Y, Pahoua P, Pem D, Pereira OL, Phonrob W, Phukhamsakda C, Raza M, Ren GC, Rinaldi AC, Rossi W, Samarakoon BC, Samarakoon MC, Sarma VV, Senanayake IC, Singh A, Souza MF, Souza-Motta CM, Spielmann AA, Su W, Tang X, Tian XG, Thambugala KM, Thongklang N, Tennakoon DS, Wannathes N, Wei DP, Welti S, Wijesinghe SN, Yang H, Yang Y, Yuan HS, Zhang H, Zhang J, Balasuriya A, Bhunjun CS, Bulgakov TS, Cai L, Camporesi E, Chomnunti P, Deepika YS, Doilom M, Duan WJ, Han SL, Huanraluek N, Jones EBG, Lakshmidevi N, Li Y, Lumyong S, Lu ZL, Khuna S, Kumla J, Manawasinghe IS, Mapook A, Punyaboon W, Tibpromma S, Lu YZ, Yan JY, Wang Y (2022) Fungal diversity notes 1512–1610: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 117(1): 1–272. https://doi.org/10.1007/s13225-022-00513-0
  • Kodsueb R, McKenzie EH, Lumyong S, Hyde KD (2008) Fungal succession on woody litter of Magnolia liliifera (Magnoliaceae). Fungal Diversity 31: 30(5): 55–72.
  • Li GJ, Hyde KD, Zhao RL, Hongsanan S, Abdel-Aziz FA, Abdel-Wahab MA, Alvarado P, Alves-Silva G, Ammirati JF, Ariyawansa HA, Baghela A (2016) Fungal diversity notes 253–366: taxonomic and phylogenetic contributions to fungal taxa. Fungal diversity 78: 1–237. https://doi.org/10.1007/s13225-016-0366-9
  • Liu J, Hu Y, Luo X, Castañeda-Ruíz RF, Xia J, Xu Z, Cui R, Shi X, Zhang L, Ma J (2023) Molecular phylogeny and morphology reveal four novel species of Corynespora and Kirschsteiniothelia (Dothideomycetes, Ascomycota) from China: A checklist for Corynespora reported worldwide. Journal of Fungi 9(1): е107. https://doi.org/10.3390/jof9010107
  • Maharachchikumbura SS, Chen Y, Ariyawansa HA, Hyde KD, Haelewaters D, Perera RH, Samarakoon MC, Wanasinghe DN, Bustamante DE, Liu JK, Lawrence DP (2021) Integrative approaches for species delimitation in Ascomycota. Fungal Diversity 09(1): 155–179. https://doi.org/10.1007/s13225-021-00486-6
  • Marincowitz S, Crous PW, Groenewald JZ (2008) Microfungi occurring on Proteaceae in the fynbos. CBS Biodiversity Series 7: 1–166.
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov 2010, New Orleans, 8 pp. https://doi.org/10.1109/GCE.2010.5676129
  • Nishi M, Okano I, Sawada T, Hara Y, Nakamura K, Inagaki K, Yaguchi T (2018) Chronic Kirschsteiniothelia infection superimposed on a pre-existing non-infectious bursitis of the ankle: the first case report of human infection. BMC Infectious Diseases 8(1): 1–5. https://doi.org/10.1186/s12879-018-3152-3
  • Poch GK, Gloer JB, Shearer CA (1992) New bioactive metabolites from a freshwater isolate of the fungus Kirschsteiniothelia sp. Journal of Natural Products 55(8): 1093–1099. https://doi.org/10.1021/np50086a010
  • Rodríguez-Andrade E, Stchigel AM, Guarro J, Cano-Lira JF (2019) Fungal diversity of deteriorated sparkling wine and cork stoppers in Catalonia, Spain. Microorganisms 8(1): 1–12. https://doi.org/10.3390/microorganisms8010012
  • Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Schoch CL, Crous PW, Groenewald JZ, Boehm EWA, Burgess TI, de Gruyter J, de Hoog GS, Dixon LJ, Grube M, Gueidan C, Harada Y, Hatakeyama S, Hirayama K, Hosoya T, Huhndorf SM, Hyde KD, Jones EBG, Kohlmeyer J, Kruys Å, Li YM, Lücking R, Lumbsch HT, Marvanová L, Mbatchou JS, McVay AH, Miller AN, Mugambi GK, Muggia L, Nelsen MP, Nelson P, Owensby CA, Phillips AJL, Phongpaichit S, Pointing SB, Pujade-Renaud V, Raja HA, Rivas ER, Robbertse B, Ruibal C, Sakayaroj J, Sano T, Selbmann L, Shearer CA, Shirouzu T, Slippers B, Suetrong S, Tanaka K, Volkmann-Kohlmeyer B, Wingfield MJ, Wood AR, Woudenberg JHC, Yonezawa H, Zhang Y, Spatafora JW (2009) A class-wide phylogenetic assessment of Dothideomycetes. Studies in Mycology 64: 1–15. https://doi.org/10.3114/sim.2009.64.01
  • Senanayake IC, Rathnayaka AR, Marasinghe DS, Calabon MS, Gentekaki E, Lee HB, Hurdeal VG, Pem D, Dissanayake LS, Wijesinghe SN, Bundhun D (2020) Morphological approaches in studying fungi: Collection, examination, isolation, sporulation and preservation. Mycosphere: Journal of Fungal Biology 11(1): 2678–2754. https://doi.org/10.5943/mycosphere/11/1/20
  • Staden R, Judge DP, Bonfield JK (2003) Analysing sequences using the Staden package and EMBOSS. In Introduction to Bioinformatics: A Theoretical and practical approach. Humana Press, Totowa, 393–410. https://doi.org/10.1007/978-1-59259-335-4_24
  • Su H, Hyde KD, Maharachchikumbura SSN, Ariyawansa HA, Luo Z, Promputtha I, Tian Q, Lin C, Shang Q, Zhao Y, Chai H, Su H, Hyde KD, Maharachchikumbura SS, Ariyawansa HA, Luo Z, Promputtha I, Tian Q, Lin C, Shang Q, Zhao Y, Chai H, Liu X, Bahkali AH, Bhat JD, McKenzie EHC, Zhou D (2016) The families Distoseptisporaceae fam. nov., Kirschsteiniotheliaceae, Sporormiaceae and Torulaceae, with new species from freshwater in Yunnan Province, China. Fungal Diversity 80(1): 375–409. https://doi.org/10.1007/s13225-016-0362-0
  • Suetrong S, Schoch CL, Spatafora JW, Kohlmeyer J, Volkmann-Kohlmeyer B, Sakayaroj J, Phongpaichit S, Tanaka K, Hirayama K, Jones EB (2009) Molecular systematics of the marine Dothideomycetes. Studies in Mycology 64(1): 155–173. https://doi.org/10.3114/sim.2009.64.09
  • Tennakoon DS, Kuo CH, Purahong W, Gentekaki E, Pumas C, Promputtha I, Hyde KD (2022) Fungal community succession on decomposing leaf litter across five phylogenetically related tree species in a subtropical forest. Fungal Diversity 115(1): 73–103. https://doi.org/10.1007/s13225-022-00508-x
  • Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172(8): 4238–4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990
  • Wang YZ, Aptroot A, Hyde KD (2004) Revision of the genus Amphisphaeria. Fungal Diversity Research Series 13: 1–168.
  • Wijayawardene NN, Hyde KD, Dai DQ, Sánchez-García M, Goto BT, Saxena RK, Erdoğdu M, Selçuk F, Rajeshkumar KC, Aptroot A, Błaszkowski J, Boonyuen N, da Silva GA, de Souza FA, Dong W, Ertz D, Haelewaters D, Jones EBG, Karunarathna SC, Kirk PM, Kukwa M, Kumla J, Leontyev DV, Lumbsch HT, Maharachchikumbura SSN, Marguno F, Martínez-Rodríguez P, Mešić A, Monteiro JS, Oehl F, Pawłowska J, Pem D, Pfliegler WP, Phillips AJL, Pošta A, He MQ, Li JX, Raza M, Sruthi OP, Suetrong S, Suwannarach N, Tedersoo L, Thiyagaraja V, Tibpromma S, Tkalčec Z, Tokarev YS, Wanasinghe DN, Wijesundara DSA, Wimalaseana SDMK, Madrid H, Zhang GQ, Gao Y, Sánchez-Castro I, Tang LZ, Stadler M, Yurkov A, Thines M (2022) Outline of Fungi and fungus-like taxa – 2021. Mycosphere: Journal of Fungal Biology 13(1): 53–453. https://doi.org/10.5943/mycosphere/13/1/2
  • Xu RF, Phukhamsakda C, Dai DQ, Karunarathna SC, Tibpromma S (2023) Kirschsteiniothelia xishuangbannaensis sp. nov. from pará rubber (Hevea brasiliensis) in Yunnan, China. Current Research in Environmental & Applied Mycology 13(1): 34–56. https://doi.org/10.5943/cream/13/1/3 [Journal of Fungal Biology]
  • Yang J, Liu LL, Jones EG, Hyde KD, Liu ZY, Bao DF, Liu NG, Li WL, Shen HW, Yu XD, Liu JK (2023) Freshwater fungi from karst landscapes in China and Thailand. Fungal Diversity 119(1): 1–212. https://doi.org/10.1007/s13225-023-00514-7
login to comment