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Research Article
Identification of three novel species and one new record of Kirschsteiniothelia (Kirschsteiniotheliaceae, Kirschsteiniotheliales) from Jiangxi, China
expand article infoXing-Xing Luo‡, Ming-Gen Liao‡, Ya-Fen Hu‡, Xiu-Guo Zhang§, Zhao-Huan Xu‡, Jian Ma‡
‡ Jiangxi Agricultural University, Nanchang, China
§ Shandong Agricultural University, Taian, China

Corresponding author: Jian Ma ( jxaumj@126.com )

Academic editor: Samantha C. Karunarathna
© 2025 Xing-Xing Luo, Ming-Gen Liao, Ya-Fen Hu, Xiu-Guo Zhang, Zhao-Huan Xu, Jian Ma.
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: Luo X-X, Liao M-G, Hu Y-F, Zhang X-G, Xu Z-H, Ma J (2025) Identification of three novel species and one new record of Kirschsteiniothelia (Kirschsteiniotheliaceae, Kirschsteiniotheliales) from Jiangxi, China. MycoKeys 112: 277-306. https://doi.org/10.3897/mycokeys.112.142028
Open Access

Abstract

Fungal diversity is rapidly expanding, with numerous species being discovered worldwide. While plant debris is a habitat favoring the survival and multiplication of various microbial species. In this study, several kirschsteiniothelia-like isolates were collected from dead branches of unidentified perennial dicotyledonous plants. Based on morphological examination and phylogenetic analyses of combined ITS, LSU, and SSU sequences data using maximum-likelihood and Bayesian inference, three new species of Kirschsteiniothelia, namely K. ganzhouensis, K. jiangxiensis, and K. jiulianshanensis, were introduced, and one known species, K. inthanonensis, was recorded for the first time from China. To improve our comprehensive knowledge of the species diversity of Kirschsteiniothelia, all accepted Kirschsteiniothelia species with morphological characteristics, sequence data, asexual morphs, habitat, host, and locality are listed.

Key words

Dothideomycetes, new species, phylogeny, saprobic fungi, taxonomy

Introduction

Fungi are a diverse group of organisms that widely exist in nature and play an important role in ecosystem processes and functioning (Schimann et al. 2017). To date, approximately 165,000 fungal species have been recorded (Hyde 2022; Phukhamsakda et al. 2022; Index Fungorum 2024), but this is only a tiny fraction of the 2 to 11 million estimated species (Phukhamsakda et al. 2022; Niskanen et al. 2023), and many hidden species are still waiting to be explored. In recent years, the fungal diversity in China has drawn the attention of taxonomists, and a total of 27,807 fungal species and subspecies have been recorded by the Catalogue of Life China 2024 Annual Checklist. The recorded database strongly suggests that more research on fungal diversity in China is needed.

The genus Kirschsteiniothelia D. Hawksw. was introduced by Hawksworth (1985) for six combinations derived from Microthelia Körb. [= Anisomeridium (Müll. Arg.) M. Choisy] and Sphaeria Haller (= Hypoxylon Bull.) and was mainly characterized by superficial to semi-immersed, globose or subglobose, dark brown to black ascomata with fissitunicate, cylindrical or clavate, bitunicate, 8-spored asci and brown to dark brown, ellipsoidal, smooth-walled, 1(–2)-septate ascospores with or without a mucilaginous sheath (Hawksworth 1985; Boonmee et al. 2012; Hyde et al. 2013, Mehrabi et al. 2017). The generic type species, K. aethiops (Sacc.) D. Hawksw., has been linked with the asexual fungus Dendryphiopsis atra (Corda) S. Hughes (generic type) based on pure culture and sequence data (Hughes 1978; Hawksworth 1985; Boonmee et al. 2012). Wijayawardene et al. (2014) further proposed to use the name Kirschsteiniothelia over Dendryphiopsis S. Hughes, considering the requirement for fewer name changes, and made the correct name Kirschsteiniothelia atra (Corda) D. Hawksw. [≡ Dendryphiopsis atra (Corda) S. Hughes] as the type species. Su et al. (2016) first reported the sporidesmium-like asexual morph (K. submersa Hong Y. Su & K.D. Hyde) in Kirschsteiniothelia based on molecular evidence, and later the sporidesmium-like asexual morphs were frequently reported in Kirschsteiniothelia with undetermined sexual morphs (Li et al. 2016; Hyde et al. 2017; Bao et al. 2018; Sun et al. 2021; Jayawardena et al. 2022; Hyde et al. 2023; Liu et al. 2023; Xu et al. 2023; Yang et al. 2023; Zhang et al. 2023; de Farias et al. 2024; Sruthi et al. 2024). Thus, Kirschsteiniothelia has two types of asexual morphs, namely dendryphiopsis-like and sporidesmium-like. The dendryphiopsis-like asexual morph is characterized by macronematous, branched at the apex, forming a stipe and head, brown to dark brown, determinate or percurrently extending conidiophores with mono- to polytretic, integrated, terminal and lateral conidiogenous cells that produce acrogenous, solitary or catenate, septate conidia. The sporidesmium-like asexual morph has macronematous, unbranched conidiophores with integrated, terminal, monoblastic or monotretic, determinate or irregularly extending conidiogenous cells that produce acrogenous, solitary or catenate, septate conidia with or without a mucilaginous sheath (Sun et al. 2021; Liu et al. 2023; Xu et al. 2023; Sruthi et al. 2024; Tang et al. 2024).

Kirschsteiniothelia is a genus of the Dothideomycetes O.E. Erikss. & Winka, of which familial placements have undergone several revisions. It was originally assigned to the family Pleosporaceae Nitschke by Hawksworth (1985) and later transferred to Pleomassariaceae M.E. Barr by Barr (1993) based on host, morphology, and mimicry. Schoch et al. (2006) revealed that K. aethiops (generic type) was not phylogenetically close to Pleosporaceae based on molecular data and should be placed in a separate family. Schoch et al. (2009) and Suetrong et al. (2009) further showed that two other Kirschsteiniothelia species, K. elaterascus Shearer and K. maritima (Linder) D. Hawksw., clustered into Morosphaeriaceae Suetrong, Sakay., E.B.G. Jones & C.L. Schoch and Mytilinidiaceae Kirschst., respectively. On this basis, Boonmee et al. (2012) introduced a new family, Kirschsteiniotheliaceae Boonmee & K.D. Hyde, to accommodate taxa grouping with K. aethiops based on combined ITS, LSU, and SSU sequence data, and transferred K. elaterascus and K. maritima to Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L. Schoch and Halokirschsteiniothelia Boonmee & K.D. Hyde, respectively. Later, Hernandez-Restrepo et al. (2017) treated Kirschsteiniotheliaceae in a new order, Kirschsteiniotheliales Hern.-Restr., R.F. Castañeda, Gené & Crous, based on its distant relationship to other lineage representatives of different orders in Dothideomycetes. Hongsanan et al. (2020) further showed that Kirschsteiniotheliales cluster with Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss., but diverged around 221 MYA.

Jiangxi Province is located in the southeast of China. Its rich vegetation and subtropical climatic regimes favor the survival and multiplication of various microbial species. However, its mycobiota, especially of mitosporic fungi, is relatively backward. During our continuing survey of saprophytic microfungi from this region, several interesting hyphomycetes were collected on dead branches of unidentified plants. Both molecular analyses and morphological data placed four species within the genus Kirschsteiniothelia. Three of these, namely K. ganzhouensis, K. jiangxiensis, and K. jiulianshanensis, are introduced as new to science, while the fourth is K. inthanonensis J. Louangphan & Gomes de Farias, a new record from China.

Materials and methods

Collections and examination of specimens

The samples of dead branches were collected randomly from the forest ecosystem of Guanshan and Jiulianshan National Nature Reserves, Jiangxi Province, China, placed in Ziplock plastic bags with collection information (Rathnayaka et al. 2024), and taken to the laboratory of conservation and utilization of fungal resources. Samples were processed and examined following the methods described in Ma et al. (2011). Colonies present on the surface of dead branches were examined and observed visually using a stereomicroscope (Motic SMZ-168, Xiamen, China) at varying magnifications ranging from 0.75 to 5 times. Fresh colonies were isolated with a sterile needle at 5 × magnification under a stereomicroscope, mounted on a slide with a drop of lactic acid-phenol solution (lactic acid, phenol, glycerol, and sterile water in proportions of 1:1:2:1), and subsequently scrutinized under an Olympus BX 53 light microscope equipped with an Olympus DP 27 digital camera (Olympus Optical Co., Tokyo, Japan) for microscopic morphological characterization. The conidia of the target colony were directly collected from the specimen using the tip of a sterile toothpick dipped in 40% sterile glycerin water. These conidia were then placed on the surface of PDA (20% potato + 2% dextrose + 2% agar, wt/vol) and incubated at 25 °C. The individual germinated conidia were transferred to fresh PDA plates and incubated in an incubator maintained in darkness at 25 °C. Culture characteristics were meticulously examined and recorded after 5 days. Colony colors were evaluated according to Rayner’s charts (Rayner 1970). All fungal strains have been preserved in sterilized glycerin at a concentration of 10% and stored at temperatures of approximately 4 °C for future studies. The specimens and cultures studied have been archived within the Herbarium of Jiangxi Agricultural University, Plant Pathology, Nanchang, China (HJAUP). The names assigned to new taxa have been officially registered within MycoBank (http://www.mycobank.org).

DNA extraction, PCR amplification, and sequencing

Total genomic DNA was extracted from fungal cultures grown on PDA plates for 2 weeks at 25 °C using the Solarbio Fungal Genomic DNA Extraction Kit (Beijing Solarbio Science & Technology Co., Ltd., Beijing, China). Three different gene regions, ITS, LSU, and SSU, were selected for this study. Primer pairs ITS5/ITS4 (White et al. 1990), 28S1-F/28S3-R, and 18S-F/18S-R (Xia et al. 2017) were used to amplify parts of the ITS, LSU, and SSU loci, respectively. The final volume of the PCR reaction was carried out in a 20 μL reaction volume containing 10 μL of 2 × Power Taq PCR MasterMix, 0.8 μL each of forward and reverse primer, 1 μL of DNA template, and 7.4 µL of ddH2O. The PCR thermal cycling conditions of ITS, LSU, and SSU were initialized at 94 °C for 3 min, followed by 35 cycles of denaturation at 94 °C for 15 s, annealing at 54 °C for 15 s, elongation at 72 °C for 30 s, a final extension at 72 °C for 10 min, and finally kept at 4 °C. The PCR products were visualized on 1% agarose gel electrophoresis stained with ethidium bromide. Sequencing was performed bidirectionally by Hunan Youkanglai Biotechnology Co., Ltd., Changsha, China. Newly obtained sequences in this study have been deposited in NCBI GenBank (www.ncbi.nlm.nih.gov, accessed on 25 May 2024; Table 1).

Table 1.
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Names, strain numbers, and corresponding GenBank accessions of Kirschsteiniothelia taxa used in the phylogenetic analyses. New sequences are indicated in bold.

Species Strain Number GenBank Accession Numbers
ITS LSU SSU
Acrospermum adeanum M133 EU940180 EU940104 EU940031
A. compressum M151 EU940161 EU940084 EU940012
A. gramineum M152 EU940162 EU940085 EU940013
Anisomeridium ubianum MPN94 – GU327709 JN887379
Flavobathelium epiphyllum MPN67 – GU327717 JN887382
Kirschsteiniothelia acutispora MFLU 21-0127 OP120780 ON980758 ON980754
K. agumbensis NFCCI 5714 T PP029048 – PP029049
K. aquatica MFLUCC 17-1685 T MH182587 MH182594 MH182618
K. arasbaranica IRAN 2509C KX621986 KX621987 KX621988
K. arasbaranica IRAN 2508C T KX621983 KX621984 KX621985
K. atra CBS 109.53 – AY016361 AY016344
K. atra MFLUCC 15-0424 KU500571 KU500578 KU500585
K. bulbosapicalis GZCC 23-0732 T PQ248937 PQ248933 PQ248929
K. cangshanensis MFLUCC 16-1350 T MH182584 MH182592 –
K. chiangmaiensis MFLU 23-0358 T OR575473 OR575474 OR575475
K. crustacea MFLU 21-0129 T MW851849 MW851854 –
K. dendryphioides KUNCC 10431 T OP626354 PQ248935 PQ248931
K. dendryphioides KUNCC 10499 PQ248938 – –
K. dushanensis 18D-43 T OP377845 – –
K. ebriosa CBS H-23379 – LT985885 –
K. emarceis MFLUCC 10-0037 T HQ441570 HQ441571 HQ441572
K. esperanzae T. Raymundo 6581 T OQ877253 OQ880482 –
K. extensa MFLU 21-0130 T MW851850 MW851855 –
K. fluminicola MFLUCC 16-1263 T MH182582 MH182588 –
K. ganzhouensis HJAUP C1209 T PP505546 PP506568 PP527763
K. ganzhouensis HJAUP C1210 PQ456024 PQ443751 PQ443763
K. ganzhouensis HJAUP C1211 PQ456025 PQ443752 PQ443764
K. guangdongensis ZHKUCC 22-0233 T – OR164974 –
K. inthanonensis MFLUCC 23–0277 T OR762773 OR762781 OR764784
K. inthanonensis HJAUP C1502 PQ456029 PQ443756 PQ443768
K. inthanonensis HJAUP C1503 PQ456030 PQ443757 PQ443769
K. jiangxiensis HJAUP C1273 T PP505548 PP506566 PP506565
K. jiangxiensis HJAUP C1274 PQ456026 PQ443753 PQ443765
K. jiangxiensis HJAUP C1275 PQ456027 PQ443754 PQ443766
K. jiulianshanensis HJAUP C1313 T PP505549 PP506562 PP506563
K. jiulianshanensis HJAUP C1314 PQ456028 PQ443755 PQ443767
K. laojunensis KUN L88727 T PP081658 – PP081651
K. lignicola MFLUCC 10-0036 T HQ441567 HQ441568 HQ441569
K. longirostrata GZCC 23-0733 T PQ248939 PQ248934 PQ248930
K. longisporum UESTCC 24.0190 T PQ038266 PQ038273 PQ046108
K. nabanheensis HJAUP C2004 T OQ023197 OQ023273 OQ023038
K. nabanheensis HJAUP C2006 OQ023274 OQ023275 OQ023037
K. phoenicis MFLUCC 18-0216 T MG859978 MG860484 MG859979
K. pini UESTCC 24.0131 T PP835321 PP835315 PP835318
K. puerensis ZHKUCC 22-0271 T OP450977 OP451017 OP451020
K. puerensis ZHKUCC:22-0272 OP450978 OP451018 OP451021
K. ramus GZCC:23-0596 T OR098711 OR091333 –
K. rostrata MFLUCC 15-0619 T KY697280 KY697276 KY697278
K. rostrata MFLUCC 16-1124 – MH182590 –
K. saprophytica MFLUCC 23–0275 T OR762774 OR762783 –
K. saprophytica MFLUCC 23–0276 OR762775 OR762782 –
K. septemseptata MFLU 21-0126 T OP120779 ON980757 ON980752
K. sichuanensis UESTCC 24.0127 T PP785368 PP784322 –
K. spatiosa MFLU 21-0128 T OP077294 – ON980753
K. submersa MFLUCC 15-0427 T KU500570 KU500577 KU500584
K. submersa S-481 – MH182591 MH182616
K. tectonae MFLUCC 12-0050 T KU144916 KU764707 –
K. tectonae MFLUCC 23-0272 OR762772 OR762780 OR764783
K. thailandica MFLUCC 20-0116 T MT985633 MT984443 MT984280
K. thujina JF 13210 T KM982716 KM982718 KM982717
K. vinigena CBS H-23378 T – LT985883 –
K. xishuangbannaensis ZHKUCC 22-0220 T OP289566 OP303181 OP289564
K. xishuangbannaensis ZHKUCC 22-0221 OP289563 OP303182 OP289565
K. zizyphifolii MFLUCC 23–0270 T OR762768 OR762776 OR764779
Megalotremis verrucosa MPN104 – GU327718 JN887383
Phyllobathelium anomalum MPN 242 – GU327722 JN887386
P. firmum ERP 3175 – GU327723 –
Pseudorobillarda eucalypti MFLUCC 12-0422 KF827451 KF827457 KF827463
Ps. phragmitis CBS 398.61 MH858101 EU754203 EU754104
Strigula guangxiensis HMAS-L0138040 T KY100301 MK206256 –
S. nemathora MPN 72 – JN887405 JN887389
Tenuitholiascus porinoides HMAS-L0139638 T – MK206259 MK352441
T. porinoides HMAS-L0139639 – MK206258 MK352442
T. porinoides HMAS-L0139640 – MK206260 MK352443

Notes: “T” indicates ex-type strain. “—” stands for unavailability of sequence data in GenBank.

Phylogenetic analyses

Novel sequences were generated from ten strains in this study, and all available reference sequences of Kirschsteiniothelia species were downloaded from GenBank. All sequences in this study included in the phylogenetic analyses are summarized in Table 1. Each gene region was independently aligned using the online version of MAFFT v.7 (Katoh and Standley 2013) on the web server (http://maffth.cbrc.jp/alignment/server/, accessed on 10 December 2024). The alignment was reviewed in MEGA v.7, followed by minor manual adjustments to ensure character homology between taxa. A matrix was formed with 60 strains (514 characters) for ITS, 70 strains (581 characters) for LSU, and 55 strains (1,239 characters) for SSU. The aligned matrices were concatenated into a single matrix (74 strains, 2334 characters). These sequence data were concatenated by Phylosuite software v1.2.1 within “Concatenate Sequence” (Zhang et al. 2020), and the concatenated aligned dataset was analyzed separately using maximum-likelihood (ML) and Bayesian inference (BI). The best evolutionary model for each alignment dataset was constructed using ModelFinder (Kalyaanamoorthy et al. 2017). Maximum-likelihood phylogenies were inferred using IQ-TREE (Nguyen et al. 2015) within 10,000 ultrafast bootstraps (Minh et al. 2013) under the best partitioned model. The optima trees were inferred using the heuristic search option with 1000 random sequence additions. The best-fit model was TN+F+I+G4 for ITS and LSU and TNe+I+G4 for SSU alignments. Bayesian inference phylogenies were inferred using MrBayes 3.2.6 (Ronquist et al. 2012) based on the partition model (2 parallel runs, 2,000,000 generations), and the best nucleotide substitution model for each locus was identified using ModelFinder of Phylosuite software v1.2.1 to be GTR+F+I+G4 for ITS and LSU and SYM+I+G4 for SSU. The resulting trees were visualized using FigTree v.1.4.2 (Zhang et al. 2020) (http://tree.bio.ed.ac.uk/software/figtree, accessed on 10 December 2024) and further edited in Adobe Illustrator 2021. The alignments and trees were deposited in TreeBASE: S31882 (http://treebase.org/treebase-web/home.html).

Results

Molecular phylogeny

The phylogenetic tree inferred from maximum-likelihood and Bayesian inference analyses based on combined ITS, LSU, and SSU sequence data consisted of four orders (Acrosperales, Kirschsteiniotheliales, Monoblastiales, and Strigulales). The concatenated sequence matrix comprised 74 sequences with 2334 total characters in the combined dataset (ITS: 1–514, LSU: 515–1095, SSU: 1096–2334), 1151 distinct patterns, 670 parsimony informative sites (ITS: 270, LSU: 237, SSU: 163), 349 singleton sites, and 1315 constant sites. Pseudorobillarda eucalypti (MFLUCC 12-0422) and Ps. phragmitis (CBS 398.61) were regarded as the outgroup. The phylogenetic trees have a similar topology, obtained from the combined dataset of maximum-likelihood and Bayesian inference analyses. The best-scoring ML concatenated tree (lnL = –18756.227) with superimposed posterior probabilities from MrBayes analysis is shown in Fig. 1. Phylogenetic analyses of the ITS+LSU+SSU concatenated datasets showed that these ten strains nested within the genus Kirschsteiniothelia, representing four independent lineages (Fig. 1). Kirschsteiniothelia ganzhouensis (HJAUP C1209, HJAUP C1210, and HJAUP C1211) clustered sister to K. fluminicola (MFLUCC 16–1263) with 100% ML/0.99 BI bootstrap support. Kirschsteiniothelia jiangxiensis (HJAUP C1273, HJAUP C1274, and HJAUP C1275) formed an independent lineage basal to Clade 1 with 86% ML/0.90 BI bootstrap support. Kirschsteiniothelia jiulianshanensis (HJAUP C1313 and HJAUP C1314) forms a distinct clade sister to the clade containing K. thujina (JF 13210) and K. laojunensis (KUN-L 88727) with 100% ML/1.00 BI bootstrap support. In addition, our new collection (HJAUP C1502 and HJAUP C1503) clustered together with the known species K. inthanonensis (MFLUCC 23-0277) with 100% ML/1.00 BI bootstrap support, indicating they represent the same species.

Figure 1. 

Maximum-likelihood phylogenetic tree of Kirschsteiniothelia based on the combined ITS, LSU, and SSU sequence data. The ML and BI bootstrap support values equal to or above 80% and 0.90 are given above the nodes. Bar = 0.06 substitutions per nucleotide position. The tree was rooted to Pseudorobillarda eucalypti (MFLUCC 12-0422) and Ps. Phragmitis (CBS 398.61). Strains of the species from the present study were marked in red. Orders were indicated on the right side of the tree in blocks. Some branches are shortened according to the indicated multipliers to fit the page size, and these are indicated by the symbol (//).

Taxonomy

Kirschsteiniothelia ganzhouensis Y.F. Hu & Jian Ma, sp. nov.

MycoBank No: 856638
Fig. 2

Type

China • Jiangxi Province, Ganzhou City, Longnan County, Jiulianshan National Nature Reserve, on dead branches of an unidentified broadleaf tree, 29 June 2022, Y.F. Hu (HJAUP M1209, holotype), ex-type living culture, HJAUP C1209 = HJAUP C1210 = HJAUP C1211.

Etymology

The name refers to the type locality “Ganzhou City”.

Description

Saprobic on decaying wood in terrestrial habitats. Asexual morph: Hyphomycetes. Colonies on natural substratum effuse, dark brown, hairy. Mycelium superficial and immersed, composed of branched, dark brown to black, septate, smooth-walled hyphae. Conidiophores macronematous, mononematous, erect, straight or flexuous, irregular or subscorpioid branched near the apex, cylindrical, smooth, septate, dark brown to black, 146.8–200 × 7.1–10.1 μm (x̄ = 176.1 × 8.0 μm, SD = 21 × 1, n = 15). Conidiogenous cells monotretic, integrated, terminal or intercalary, cylindrical, pale brown to brown, determinate, or sometimes with several cylindrical, enteroblastic percurrent extensions. Conidia acrogenous, solitary, obclavate, straight or slightly curved, sometimes rostrate, smooth, subhyaline to pale brown, 2–7(–14)-distoseptate, 20.3–65.8(–164) × 3.0–5.3 μm (x̄ = 36.4 × 4.7 μm, SD = 12 × 0.36, n = 20), tapering to 1.3–2.6 μm near the apex, 3.0–5.3 μm wide at the base, and rounded at the apex. Sexual morph: Undetermined.

Figure 2. 

Kirschsteiniothelia ganzhouensis (HJAUP M1209, holotype) a, b colonies after 4 weeks on PDA (front and reverse) c, d conidiophores and conidiogenous cells e, f conidiophores, conidiogenous cells, and conidia g conidia.

Culture characteristics

Colonies growing on PDA medium reaching 30–35 mm diam. after 4 weeks at 25 °C in darkness, irregular circular, surface yellow-brown with fluffy hyphae, reverse dark brown to black.

Note

The phylogenetic tree showed that K. ganzhouensis (HJAUP C1209, HJAUP C1210, and HJAUP C1211) clusters with K. fluminicola (MFLUCC 16-1263). Based on the BLASTn results, ITS and LSU gene sequences of K. ganzhouensis (HJAUP C1209) showed 93% (484/520, 3 gaps) and 99% (518/525, 0 gap) similarities to K. fluminicola (MFLUCC 16-1263), respectively. Moreover, K. ganzhouensis differs morphologically from K. fluminicola Z.L. Luo, K.D. Hyde & H.Y. Su (Bao et al. 2018) in having monotretic conidiogenous cells, shorter conidiophores (146.8–200 μm vs. 209–286 μm), and smaller conidia (20.3–65.8 × 3.0–5.3 μm vs. 47.5–86.5 × 8–10 μm). In addition, K. ganzhouensis further differs from K. fluminicola in that it occurs in a terrestrial habitat and not in a freshwater habitat.

Kirschsteiniothelia inthanonensis J. Louangphan & Gomes de Farias, 2024

Index Fungorum: IF901384
Facesoffungi Number: FoF14982
Fig. 3

Description

Saprobic on decaying wood in terrestrial habitats. Asexual morph: Hyphomycetes. Colonies on natural substratum effuse, dark brown, hairy. Mycelium immersed and superfcial, composed of branched, septate, dark brown to black, smooth-walled hyphae. Conidiomata synnematous, solitary, erect, cylindrical, dark brown to black, becoming narrower toward the apex, up to 1266 μm high, 110–330 μm wide at the swollen base. Conidiophores distinct, macronematous, erect, straight or flexuous, closely fasciculate, branched near the apex, septate, smooth, cylindrical, brown to dark brown, up to 1266 μm long, 4.8–8 μm wide, diverging laterally and terminally. Conidiogenous cells monotretic, integrated, terminal, cylindrical, smooth, brown, determinate, or sometimes with several cylindrical, enteroblastic percurrent extensions. Conidia acrogenous, solitary or catenate, obclavate, straight or slightly curved, smooth, olivaceous brown to brown, 2–5-euseptate, 20–48 × 8–13.3 μm (x̄ = 31.4 × 9.8 μm, SD = 9 × 1, n = 30), partly tapering towards and rounded at the apex. Sexual morph: Undetermined.

Figure 3. 

Kirschsteiniothelia inthanonensis (HJAUP M1502, holotype) a, b colonies after 4 weeks on PDA (front and reverse) c, d synnemata with conidiophores, conidiogenous cells, and conidia e conidiogenous cells and conidia f conidia.

Culture characteristics

Colonies growing on PDA medium reaching 85–90 mm diam. after 4 weeks at 25 °C in darkness, circular, surface velvety, with reddish-brown to brown mycelium, reverse brown to dark brown.

Material examined

China • Jiangxi Province, Ganzhou City, Longnan County, Jiulianshan Town, Guanshan National Nature Reserve, on dead branches of an unidentified broadleaf tree, 27 June 2021, Y.F. Hu (HJAUP M1502, holotype), living culture, HJAUP C1502 = HJAUP C1503.

Note

Kirschsteiniothelia inthanonensis was originally described with an asexual morph on the twigs of Quercus oleoides in Thailand (de Farias et al. 2024) and was known only from its type collection. Morphologically, our new collection shows high morphological similarity to K. inthanonensis except for its wider conidiophores (4.8–8 μm vs. 2.5–6.6 μm), shorter conidia (20–48 μm vs. 24–230 μm) with fewer septa (2–5-euseptate vs. 2–10-euseptate) (de Farias et al. 2024). In addition, the phylogenetic tree showed that our new collection (HJAUP C1502 and HJAUP C1503) clustered with K. inthanonensis (MFLUCC 23-0277). Based on pairwise nucleotide comparisons of ITS, LSU, and SSU, their nucleotide differences (0/517 in ITS, 2/565 in LSU, and 0/1022 in SSU) are minor. Therefore, we identified our new collection as K. inthanonensis, and it is a new record for China.

Kirschsteiniothelia jiangxiensis Y.F. Hu & Jian Ma, sp. nov.

MycoBank No: 856639
Fig. 4

Type

China • Jiangxi Province, Ganzhou City, Longnan County, Jiulianshan National Nature Reserve, on dead branches of an unidentified broadleaf tree, 29 June 2022, Y.F. Hu (HJAUP M1273, holotype), ex-type living culture, HJAUP C1273 = HJAUP C1274 = HJAUP C1275.

Etymology

The name refers to the locality “Jiangxi Province”, from where the fungus was collected.

Description

Saprobic on decaying wood in terrestrial habitats. Asexual morph: Hyphomycetes. Colonies on natural substratum effuse, dark brown, hairy. Mycelium superficial and immersed, composed of branched, septate, dark brown to black, smooth-walled hyphae. Conidiophores macronematous, mononematous, simple or branched, erect, straight or flexuous, cylindrical, smooth, septate, dark brown to black, 32.9–90.4 × 7.3–12.9 μm (x̄ = 48.7 × 9.1 μm, SD = 17 × 2, n = 15). Conidiogenous cells monoblastic, integrated, terminal, cylindrical, smooth, brown to dark brown, determinate, or sometimes with several cylindrical, enteroblastic percurrent extensions. Conidia solitary, acrogenous, obclavate, straight or curved, smooth, brown, 7–10-euseptate, 75.9–103.8 × 8.9–15.2 μm (x̄ = 90.8 × 10.9 μm, SD = 8 × 2, n = 20), tapering to 2.5–5.6 μm at the apex, 5.3–7.6 μm wide at the truncate base, and rounded at the apex. Sexual morph: Undetermined.

Figure 4. 

Kirschsteiniothelia jiangxiensis (HJAUP M1273, holotype) a, b colonies after 4 weeks on PDA (front and reverse) c conidia d, e conidiophores, conidiogenous cells, and conidia f conidiophores and conidiogenous cells.

Culture characteristics

Colonies growing on PDA medium reaching 85–90 mm diam. after 4 weeks at 25 °C in darkness, irregular circular, surface velvety, grey-white in center and brown at margin with dense mycelium, reverse dark brown to black.

Note

The phylogenetic tree showed that K. jiangxiensis (HJAUP C1273, HJAUP C1274, and HJAUP C1275) belongs to Kirschsteiniothelia and forms a distinct lineage sister to Clade 1. However, K. jiangxiensis (HJAUP C1273) differs from the morphologically most similar species, K. spatiosa (MFLU 21-0128) (Jayawardena et al. 2022), in having shorter conidiophores [32.9–90.4 μm (x̄ = 48.7) vs. 70–128 µm (x̄ = 100)] and smaller conidia [75.9–103.8 × 8.9–15.2 μm (x̄ = 90.8 × 10.9 μm) vs. 90–139 μm × 9.5–16.5 µm (x̄ = 113 × 14 μm)] with fewer septa (7–10 vs. 8–23), and further from K. spatiosa by 96 nucleotides (67/380 in ITS and 26/1032 in SSU). In addition, K. jiangxiensis also differs from other taxa in Clade 1 in the size of conidiophores and conidia.

Kirschsteiniothelia jiulianshanensis Y.F. Hu & Jian Ma, sp. nov.

MycoBank No: 856640
Fig. 5

Type

China • Jiangxi Province, Ganzhou City, Longnan County, Jiulianshan National Nature Reserve, on dead branches of an unidentified broadleaf tree, 29 June 2022, Y.F. Hu (HJAUP M1313, holotype), ex-type living culture, HJAUP C1313 = HJAUP C1314.

Etymology

The name refers to Jiulianshan National Nature Reserve, the locality where the fungus was collected.

Description

Saprobic on decaying wood in terrestrial habitats. Asexual morph: Hyphomycetes. Colonies on natural substratum effuse, dark brown, hairy. Mycelium immersed and superficial, composed of branched, dark brown to black, septate, smooth-walled hyphae. Conidiophores macronematous, mononematous, unbranched, erect, straight or flexuous, cylindrical, smooth, dark brown to black, 7–17-septate, 128.6–291.4(–430) × 7.1–10 μm (x̄ = 217.4 × 8.46 μm, SD = 85 × 0.7, n = 15). Conidiogenous cells monotretic, integrated, terminal, cylindrical, brown, determinate, or sometimes with several cylindrical, enteroblastic percurrent extensions. Conidia acrogenous, solitary, obclavate, straight or slightly curved, pale brown to brown, 4–7-euseptate, 31.4–57.1 × 10–11.4 μm (x̄ = 41.6 × 11.04 μm, SD = 9 × 0.5, n = 20), tapering to 4.6–6 μm at the apex, 2.3–3.4 μm wide at the truncate base, and rounded at the apex. Sexual morph: Undetermined.

Figure 5. 

Kirschsteiniothelia jiulianshanensis (HJAUP M1313, holotype) a, b colonies after 4 weeks on PDA (front and reverse) c–g conidiophores, conidiogenous cells, and conidia h conidia.

Culture characteristics

Colonies growing on PDA medium reaching 85–90 mm diam. after 4 weeks at 25 °C in darkness, circular, surface velvety, grey-white in center and brown at margin with dense mycelium, reverse dark brown to black.

Note

The phylogenetic tree showed that K. jiulianshanensis (HJAUP C1313 and HJAUP C1314) clusters with K. laojunensis (KUN-L 88727) and K. thujina (JF 13210). Based on nucleotide comparisons, K. jiulianshanensis (HJAUP C1313) and K. laojunensis (KUN-L 88727) showed 63 bp differences (3%, including eight gaps) in ITS and SSU regions; K. jiulianshanensis (HJAUP C1313) and K. thujina (JF 13210) showed 75 bp differences (7%, including twelve gaps) in ITS, LSU, and SSU regions. Moreover, K. jiulianshanensis was found only in its asexual morph, while K. laojunensis Q.F. Meng & S.B. Fu (Meng et al. 2024) and K. thujina (Peck) D. Hawksw. (Hawksworth 1985) are known only as the sexual morph. In addition, K. jiulianshanensis (HJAUP C1313) can be distinguished from the morphologically most similar species, K. crustacea S. Wang, Q. Zhao & K.D. Hyde (Jayawardena et al. 2022), in having longer conidiophores (128.6–291.4 μm vs. 60–170 μm) and smaller conidia (31.4–57.1 × 10–11.4 μm vs. 45–75 × 10–18 μm), and further from K. crustacea by 90 nucleotides (79/491 in ITS and 11/545 in SSU).

Discussion

Saprobic fungi are highly diverse in freshwater and terrestrial habitats, and a large number of novel taxa have been reported from many natural substrates such as submerged wood, dead branches, bark, culms, and leaves (Ellis 1971, 1976; Wu and Zhuang 2005; Hernandez-Restrepo et al. 2017; Luo et al. 2019; Hyde et al. 2023; Dissanayake et al. 2024; Tang et al. 2024). In our study, numerous hyphomycetes were collected on dead branches from terrestrial habitats in Jiangxi Province, China. Based on morphological characteristics and multi-locus (ITS, LSU, and SSU) phylogenetic analyses, three new species of Kirschsteiniothelia, viz. K. ganzhouensis, K. jiangxiensis, and K. jiulianshanensis, and one new Chinese record, K. inthanonensis, were identified, which contributed to our understanding of the species diversity of this genus.

Kirschsteiniothelia was established by Hawksworth (1985), with K. aethiops as the type species. To date, 62 epithets of Kirschsteiniothelia have been recorded (Index Fungorum 2024; Sruthi et al. 2024; Tian et al. 2024), but K. elaterascus and K. maritima were respectively transferred to Neohelicascus W. Dong, H. Zhang, K.D. Hyde & Doilom and Halokirschsteiniothelia Boonmee & K.D. Hyde based on phylogenetic analyses (Boonmee et al. 2012; Dong et al. 2020). Wijayawardene et al. (2014) further proposed to use Kirschsteiniothelia over Dendryphiopsis in the context of the one fungus, one name initiative and synonymized K. aethiops with K. atra (Corda) D. Hawksw. Accordingly, K. incrustans, derived from Microthelia incrustans (Ellis & Everh.) Corlett & S. Hughes (a synonym of K. aethiops), was also treated as the synonym of K. atra (Wijayawardene et al. 2014; Mehrabi et al. 2017; Index Fungorum 2024; Jin et al. 2024). Mehrabi et al. (2017) provided an identification key to 20 well-documented Kirschsteiniothelia species and listed their principal synonyms. Sun et al. (2021) provided a synopsis of 35 Kirschsteiniothelia species with distribution, habitat, host, and morphology type of each species, but K. elaterascus, K. incrustans, and K. maritima were previously rejected out of Kirschsteiniothelia, and the other five species, viz. K. arbuscula, K. binsarensis, K. biseptata, K. fascicularis, and K. goaensis in Kirschsteiniothelia, were invalid (Turland et al. 2018: Art. F.5.1: no identifier number cited, and Art. 41.1: lacking a full and direct basionym reference) until Sruthi et al. (2024) legitimately placed them as five new combinations of Kirschsteiniothelia. Xu et al. (2023) summarized the morphological, host, and location information of 29 Kirschsteiniothelia species. Subsequently, Sruthi et al. (2024) listed 34 asexual morphs under Kirschsteiniothelia, and Tang et al. (2024) provided a checklist for 59 Kirschsteiniothelia species with their host, habitat, country, and reported morph, but K. dujuanhuensis was unpublished. Thus, Kirschsteiniothelia currently comprises 58 valid species. All species are known for their asexual or sexual morphs, and only five species, namely K. atra, K. emarceis, K. lignicola, K. recessa, and K. saprophytica, are known from both morphs.

Traditionally, Kirschsteiniothelia species have been characterized and identified based on morphological characteristics, but the lack of molecular data made it difficult to evaluate the phylogenetic relationships and taxonomic placements of some doubtful or morphological similarity species. With the development of molecular technology, multi-gene combined analysis has gradually occupied a dominant status in fungal taxonomy. To date, there are 39 Kirschsteiniothelia species with molecular data (Tang et al. 2024), and recent studies indicated that the concatenated dataset of ITS, LSU, and SSU sequences shows good resolution in revealing the phylogeny of Kirschsteiniothelia. Since 2012, all described Kirschsteiniothelia species were identified using ITS, LSU, and SSU except for K. ebriosa and K. vinigena using LSU (Rodríguez-Andrade et al. 2020), K. esperanzae, K. guangdongensis, K. laojunensis, and K. ramus using ITS and LSU (Raymundo et al. 2023; Senanayake et al. 2023; Zhang et al. 2023; Meng et al. 2024). In this study, we also conducted phylogenetic analyses using ITS, LSU, and SSU sequences, and our newly obtained ten strains nested within the genus Kirschsteiniothelia formed four independent lineages with reliable support value and can be recognized as three new phylogenetic species, namely K. ganzhouensis, K. jiangxiensis, and K. jiulianshanensis, and one known species, K. inthanonensis.

Kirschsteiniothelia is widely distributed in tropical and subtropical regions. Most species of this genus are known from dead woods or twigs in terrestrial and freshwater habitats, but occasionally, some species have been reported to be associated with orchid symbiosis, bioactive metabolites, and human infection (Poch et al. 1992; Nishi et al. 2018; Chen et al. 2022). Currently, with the addition of our species, there are 61 species in Kirschsteiniothelia (Tables 2–4), including 38 asexual species (Table 2), 18 sexual species (Table 3), and 5 species known for both morphs (Table 4), and 23 of which have dendryphiopsis-like asexual morphs and 20 have sporidesmium-like asexual morphs. In addition, on the basis of the typification, we found that members in Kirschsteiniothelia are mainly reported from China (22 species), Thailand (15 species), the USA (10 species), and India (4 species), whereas most regions are still essentially unrecorded. Thus, further research with the morpho-molecular approach is necessary to explore the hidden species diversity of Kirschsteiniothelia from different geographic regions and further focus on the correction of their asexual and sexual morphs, which will make significant contributions to the taxonomy of this genus and be necessary to quantify their roles in natural ecosystems.

Table 2.
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Synopsis of morphological characteristics, sequence data, type of asexual morph, habitat, host, and locality compared across Kirschsteiniothelia species with asexual morphs.

Species Conidiophores (μm) Conidiogenous cells Conida Sequence data Type of asexual morph Habitat/Host/ Locality References
Size (µm) Morphology
Kirschsteiniothelia acutispora 180–260 × 7–12.5 Monoblastic 75–120 × 10.5–19.5 Obclavate to obspathulate, rostrate, mid to dark brown, becoming pale brown towards the apex, 7–12-euseptate Present Sporidesmium-like Terrestrial/On dead branches/Thailand Jayawardena et al. (2022)
K. agumbensis 9.05–14.95 × 7.25–8.5 Monoblastic 228–450.5 × 15–23.5 Cylindrical, rostrate, dark brown to brown, pale brown at apex, coarsely verrucose to granulate to punctate, 18–41-euseptate Present Sporidesmium-like Terrestrial/On decaying wood of Garcinia sp./India Sruthi et al. (2024)
K. aquatica 114–151 × 7–8 Monoblastic 35–46 × 7.5–8.5 Obclavate, subhyaline, dark brown at base, septate, sometimes percurrently proliferate at broken ends Present Sporidesmium-like Freshwater/On submerged wood/China Bao et al. (2018)
K. arbuscula 240–580 × 10–13 Monotretic 42–64 × 12–14 Subfusiform, fusiform, or obclavate, blackish olivaceous, 3–7 septate Absent Dendryphiopsis-like Terrestrial/On bark of many plants/USA Ellis (1976); Pratibha et al. (2010); Sruthi et al. (2024)
K. binsarensis 280–520 × 6.5–8 Monotretic 36–44 × 8–10 Obclavate to obclavate-fusiform, brown, 4–5-septate Absent Dendryphiopsis-like Terrestrial/On dead branches/India Subramanian and Srivastva (1994); Sruthi et al. (2024)
K. biseptata Up to 180 × 8–10 Monotretic 28–39 × 19–22 Ellipsoidal or obovate, brown, 2-septate Absent Dendryphiopsis-like Terrestrial/On dead twig/South Africa Morgan-Jones et al. (1983); Sruthi et al. (2024)
K. bulbosapicalis (47–)58–128(–199) × 7.5–12.5(–16.5) Monoblastic 118–236.5 × 15–27 Cylindrical, ovoid to obclavate, rostrate, olivaceous to reddish-brown to dark brown, 8–13-septate, with a spherical hyaline mucilaginous sheath Present Sporidesmium-like Terrestrial/On unidentified decaying wood/China Tang et al. (2024)
K. cangshanensis 105.5–135.5 × 6–8 Monoblastic 33–43 × 7.5–8.5 Obclavate, pale brown to brown, with a gelatinous sheath at apex, septate Present Sporidesmium-like Freshwater/On submerged wood/China Bao et al. (2018)
K. crustacea 60–170 × 6.5–10.5 Monoblastic 45–75 × 10–18 Obclavate to obspathulate, rostrate, mid to dark brown and hyaline to light brown towards the apex, 5–6-euseptate Present Sporidesmium-like Freshwater/On decaying bamboo/Thailand Jayawardena et al. (2022)
K. dendryphioides 179–467 × 4.5–8 Monotretic 30–55 × 9–13.5 Cylindrical, oblong, and occasionally clavate, brown, 2–4-septate Present Dendryphiopsis-like Freshwater/On decaying wood/China Tang et al. (2024)
K. dushanensis 160–307 × 6.5–13 Monoblastic 62–81 × 12.5–18 Fusiform lower part and euseptate, narrower cylindrical upper part, rostrate, olivaceous brown to soot brown, pale brown or subhyaline at the apex, truncate and darkened at the base, sometimes with a mucilaginous sheath surrounding the tail-like upper part or the apex, 5–11-septate, with distoseptate Present Sporidesmium-like Freshwater/On submerged wood/China Yang et al. (2023)
K. ebriosa 40–150 × 4 Mono- to polytretic 8–14 × 4–5 Cylindrical with rounded ends, mostly catenate, brown to dark brown, 1–2(–5)-septate Present Dendryphiopsis-like Freshwater/From sparkling wine/Spain Rodríguez-Andrade et al. (2020)
K. extensa 80–230 × 6.5–9.5 Monoblastic 45–120 × 5–12 Obclavate, becoming pale brown to pale towards the apex, 5–8 euseptate Present Sporidesmium-like Terrestrial/On decaying wood/Thailand Jayawardena et al. (2022)
K. fascicularis 200–450 × 9–11 Monotretic 48–90 × 5–10 Long-clavate, strongly attenuated at the base, 3–8 septate Absent Dendryphiopsis-like Terrestrial/On bark of Liquidambar/USA Hughes (1958); Sruthi et al. (2024)
K. fluminicola 209–286 × 7–9 Monoblastic 47.5–86.5 × 8–10 Solitary to short-catenate, obclavate, rostrate, subhyaline to dark brown, with conspicuous, spherical guttules in almost all cells, multi-septate Present Sporidesmium-like Freshwater/ Unidentified decaying wood/China Bao et al. (2018)
K. ganzhouensis 146.8–200 × 7.1–10.1 Monotretic 20.3–65.8(–164) × 3.0–5.3 Obclavate, rostrate, subhyaline to brown, pale at apex, 2–7(–14)-distoseptate Present Dendryphiopsis-like Terrestrial/On decaying wood/China This study
K. guangdongensis 250–350 × 10–18 Monoblastic 290–300 × 42–50 Elongated, flask-shaped, blackish brown to black, apical cell paler than others, with a thin sheath at apex, 13-septate with one longitudinal septum in 5 basal cells Present Sporidesmium-like Terrestrial/On plant twigs/China Senanayake et al. (2023)
K. goaensis 85–230 × 4–6 Monotretic 20–40 × 5–7.5 Cylindrical, rounded at both ends, dark brown, 3–5-septate Absent Dendryphiopsis-like Terrestrial/On dead and decaying bark/India Pratibha et al. (2010)
K. inthanonensis 611–1549 × 2.5–6.6 Mono- to polytretic 24–230 × 5.7–14.3 Obclavate, rostrate, grey to brown, pale at apex, 2–10-euseptate Present Dendryphiopsis-like Terrestrial/On twigs of Quercus oleoides /Thailand de Farias et al. (2024)
K. jiangxiensis 32.9–90.4 × 7.3–12.9 Monoblastic 75.9–103.8 × 8.9–15.2 Obclavate, rostrate, pale brown to brown, 7–10-euseptate Present Sporidesmium-like Terrestrial/On decaying wood/China This study
K. jiulianshanensis 128.6–291.4(–430) × 7.1–10 Monotretic 31.4–57.1 × 10–11.4 Obclavate, rostrate, subhyaline to dark brown, 4–7-euseptate Present Sporidesmium-like Terrestrial/On decaying wood/China This study
K. longirostrata 80–252 × 4.5–9.5 Monoblastic 36.5–109(–160) × 8–16 Cylindrical, obpyriform to obclavate, rostrate, guttulate, 6–18-septate, proliferating, pale brown to brown, with a mucilaginous sheath surrounding the upper part of the apex Present Sporidesmium-like Freshwater/On decaying wood/China Tang et al. (2024)
K. longisporum 115–285 × 6.5–14 Holoblastic 35–130 × 8.5–15 Cylindrical-obclavate, elongated, brown, 3–15-distoseptate, verruculose Present Dendryphiopsis-like Terrestrial/On dead branches of Pinus taeda/China Tian et al. (2024)
K. nabanheensis 320–588 × 8–12 Monotretic 32–112 × 8–12 Obclavate or fusiform, sometimes rostrate, dark brown to brown, 3–7 euseptate Present Dendryphiopsis-like Terrestrial/On dead branches/China Liu et al. (2023)
K. pini 69–124 × 3.5–7 Monoblastic 22–45 × 5–10 Obclavate, becoming brown to pale towards the apex, 3–6-euseptate Present Dendryphiopsis-like Terrestrial/On decaying branches of Pinus/China Jin et al. (2024)
K. puerensis 100–250 × 5–12 Monoblastic 60–140 × 5–20 Obclavate, pale-brown to brown, pale-brown at the apex, sometimes with 1–2 hyaline sheaths around the tip, 5–12-septate Present Sporidesmium-like Terrestrial/On dead wood of Coffea/China Hyde et al. (2023)
K. ramus 102–248 × 5–11 Monotretic 42–56 × 15–22 Cylindrical, pale olivaceous when young, brown when mature, 2–3-septate, verruculose Present Dendryphiopsis-like Freshwater/On decaying wood/China Zhang et al. (2023)
K. rostrata 190–450 × 9–15 Monoblastic 80–150 × 10–20 Obclavate, rostrate, olivaceous brown to brown, pale at apex, sometimes with a mucilaginous sheath at apex, 8–13-septate Present Sporidesmium-like Freshwater/On decaying wood/Thailand Hyde et al. (2017)
K. septemseptata 250–580 × 6.5–14.5 Mono- to polytretic 25–55 × 6.5–12.5 Obclavate, rostrate, olivaceous brown to brown, pale at apex, 5–8 euseptate Present Dendryphiopsis-like Terrestrial/On decaying wood/Thailand Jayawardena et al. (2022)
K. shimlaensis 110–268 × 12–19 Monotretic 41–81 × 13–17.5 Obovoid, oblong, broad clavate or cylindrical, dark brown or black, microguttulate, lumen aspect granulose, 2–5(–6) septate Absent Dendryphiopsis-like Terrestrial/On decaying stump of Cedrus deodara/India Verma et al. (2021)
K. sichuanensis 82–194 × 5–10 Monoblastic 34–54 × 8–14 Obclavate, becoming brown to pale towards the apex, 2–7 euseptate Present Dendryphiopsis-like Terrestrial/On dead wood/China Jin et al. (2024)
K. spatiosa 70–128 × 7.5–12.5 Monoblastic 90–139 × 9.5–16.5 Obclavate, rostrate, olivaceous brown to brown, pale at apex, sometimes with a mucilaginous sheath at apex, 8–23-euseptate Present Sporidesmium-like Terrestrial/On decaying wood/Thailand Jayawardena et al. (2022)
K. submersa 220–280 × 6–7 Monoblastic 37.5–51.5 × 8.5–9.5 Obclavate, brown to pale brown, hyaline and thinner at the tip, 4–6-septate Present Sporidesmium-like Freshwaterl/On decaying wood/China Su et al. (2016)
K. tectonae Up to 200 × 4–8 Monoblastic 135–150 × 16–17 Cylindric-obclavate, elongate, dark reddish brown, with sheath at apex, 9–25 or more septa Present Sporidesmium-like Terrestrial/On dead bark of Tectona grandis/Thailand Li et al. (2016)
K. thailandica 55–93 × 7–10 Monoblastic 74–110 × 13–20 Obclavate, olivaceous or brown, hyaline at apex, with a conspicuous, gelatinous, hyaline sheath around tip, 6–8-distoseptate Present Sporidesmium-like Terrestrial/On twigs of Ficus microcarpa/ Thailand Sun et al. (2021)
K. vinigena 100–150 × 3 Mono- to polytretic 8–80 × 4–5 Solitary or catenate, cylindrical, dark brown, smooth-walled to coarsely verrucose, 1–2(–7)-septate Present Dendryphiopsis-like Terrestrial/ From cork stopper / Spain Rodríguez-Andrade et al. (2020)
K. xishuangbannaensis 35–150 × 5–15 Monoblastic 30–150 × 5–20 Obclavate, rostrate, yellow-brown to brown, subhyaline or pale-brown at apex, some have guttules, 1–2 hyaline globose to ampulliform, mucilaginous sheaths around the tip, 3–8-septate Present Sporidesmium-like Terrestrial/On dead branches of Hevea brasiliensis/China Xu et al. (2023)
K. zizyphifolii 287–444.5 × 10.3–17(–19.7) Tretic 37.6–46.5 × 13.5–19 Cylindrical to rarely clavate, brown dark to brown, 2–3-septate Present Dendryphiopsis-like Terrestrial/On dead wood of Nayariophyton zizyphifolium/Thailand de Farias et al. (2024)

Notes: All conidia are solitary and smooth except where indicated.
Table 3.
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Synopsis of morphological characteristics, sequence data, habitat, host, and locality compared across Kirschsteiniothelia species with sexual morphs.

Species Asci (μm) Ascospores Sequence data Habitat/Host/Locality References
Size (μm) Characteristics
Kirschsteiniothelia abietina 100–110 × 20 23–28 × 6–10 Irregularly biseriate, elliptical, rounded at the ends, slightly or not at all constricted at the septum, young hyaline, nucleosomes or granular, becoming brown Absent Terrestrial/On bark of Tsuga canadensis/USA Wang et al. (2004)
K. acerina 85–95 × 20–24 22–26 × 8–11 Ellipsoid, 1-septate, the lower cell often somewhat smaller, slightly constricted at the septum, brown to dark brown, verruculose Absent Terrestrial/On absorbing mycorrhizal rootlets of Acer saccharum/USA Hawksworth (1985)
K. arasbaranica 120–180 × 30–40 (30–)34–42(–44) × (12–)13–16(–18) Narrowly to broadly ellipsoidal with rounded apex, brown to dark brown at maturity, verrucose to finely spinulose, covered with a mucilaginous sheath, guttulate, 1-septate, septum deeply constricted and submedian, the upper cell distinctly larger than the lower cell Present Terrestrial/On dead branch of Quercus petraea/Iran Mehrabi et al. (2017)
K. atkinsonii 70–90 × 9–16 14–16 × 5–6 2-seriate, clavate, light brown, upper cell wider than elongated lower cell, verrucose, 1-septate Absent Terrestrial/On leaves of Freycinetia arnotti/USA Hyde (1997)
K. chiangmaiensis 76–119 × 24–30 20–31 × 9–12 Bi- or tri-seriate in the middle and uniseriate in the top of the ascus, ellipsoid to fusiform, narrowly to broadly ellipsoidal with rounded or slightly pointed at the ends, 1-septate, septum submedian and deeply constricted, the upper cell distinctly larger than the lower cell, guttulate, brown, smooth, with a mucilaginous sheath Present Freshwater/On decaying wood/ Thailand Louangphan et al. (2024)
K. dolioloides 125–145 × 25–34 84–39 × 14–15 2-celled, with a slight constriction at the septum, young olivine, old dark brown Absent Terrestrial/On bark of Pinus/Switzerland Wang et al. (2004)
K. esperanzae (168–)178–203 × 32–35 40–50(–53) × 14–17 Ellipsoid or soleiform, 1-septate, slightly constricted at the septum, light brown to olive-brown, smooth Present Terrestrial/On decaying wood/Mexico Raymundo et al. (2023)
K. laojunensis (105–)130–162(–180) × (17–)20–30 (34–)35–55(–56) × (10–)11–14(–16) Fusiform, usually1-septate, slightly constricted at the septum, asymmetric with a slightly larger upper cell, both ends slightly subacute, guttulate, hyaline when young and turning dark brown with greenish or bluish coloration at maturity Present Terrestrial/On the bark of Abies fabri/China Meng et al. (2024)
K. phileura – 22 × 10 Ellipsoid, 1-septate, the upper cell usually larger than the lower cell, somewhat constricted at the septum, brown Absent Terrestrial/On the bark of Tilia american/USA Barr (1993)
K. phoenicis 70–112 × 14–24 18–27 × 5–7.5 Ellipsoid, rounded or slightly pointed at the ends, brown, 1-septate, septum submedian and constricted, upper cell broader than the lower cell, guttulate, smooth, with a mucilaginous sheath Present Freshwater/On rachis of Phoenix paludosa/Thailand Hyde et al. (2018)
K. populi 80–90 × 8 12 × 6 Ovate, ends rounded, fuliginous, equally uniseptate, much constricted Absent Terrestrial/On decorticated branches of Populus angustifolia/USA Wang et al. (2004)
K. proteae 54–72 × 6–8 (11–)13–17(–20) × 3–4(–5) Fusoid, 1-septate, median or submedian, smooth, with germ pore at ascospore ends, at times cells become biguttulate, pale brown to brown Absent Terrestrial/On decorticated twig litter of Protea cynaroides/South Africa Marincowitz et al. (2008)
K. reticulata 90–110 × 23–27 17–23 × 7–10 Ellipsoid, dark brown, 1-septate, septum constricted, median or occasionally submedian, with reticulate ornamentation on surface, covered with mucilaginous sheath Absent Terrestrial/On dead twigs/China Chen et al. (2006)
K. smilacis 75–100 × 16–21 20–24 × 6–8 Ellipsoid, pale brown, 1-septate, slightly constricted at septum, wall finely and inconspicuously verrucose, covered with mucilaginous sheath Absent Terrestrial/On stem of Smilax sp./China Chen et al. (2006)
K. striatispora 65–75 × 9–11 (14–)15–18(–19) × 5–6.5 Ellipsoid to somewhat soleiform, 1-septate, the cells equal in size or the lower slightly smaller, apices rounded, reddish-brown, slightly granular at first but finally with up to five longitudinal or sinuate furrows Absent Terrestrial/On dead twigs of Juniperus communis subsp. nana/Switzerland Hawksworth (1985)
K. thujina 100–140 × 17–22 (29–)36–50(–55) × (12–) 15–17(–19) Elongate-ellipsoid, slightly attenuated towards the apices, 1-septate, dark brown, apparently smooth walled, often guttulate Present Terrestrial/On decaying wood of Thuja occidentalis/USA Hawksworth (1985); Zhang and Fournier (2015)
K. umbrinoidea – 23–28 × 75–9 Oblong-fusiform, hyaline, two guttulate Absent Terrestrial/On bark of Aesculus hippocastanum/Italy Wang et al. (2004)
K. xera 70–75 × – 17–23 × 6–7 1-septate, constricted at the septum, cells somewhat unequal, guttulate, with granulate contents, uniseriate or partly biseriate Absent Terrestrial/On bark of Prunus/USA Wang et al. (2004)

Notes: All ascospores are smooth except where indicated.
Table 4.
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Synopsis of morphological characteristics, sequence data, type of asexual morph, habitat, host, and locality compared across Kirschsteiniothelia species with asexual and sexual morphs.

Species Teleomorph Anamorph Sequence Data Habitat/Host/Locality References
Asci (μm) Ascosporous morphology Morphology Type of asexual morph
Kirschsteiniothelia atra 70–90 × 12–15 25–33 × 8.5–12 μm, ellipsoidal, rounded or somewhat constricted at the apices, 1-septate, the upper cell usually larger in size, somewhat constricted at the septum, brown, sometimes appearing almost smooth Conidiophores 245–355 × 8–10 μm; Conidiogenous cells tretic, integrated, sometimes percurrent, terminal, becoming intercalary, new cell developing from apical or subapical part of subtending cell; Conidia 54–63 × 14–18 μm, solitary, cylindrical, rounded at the apex, and narrowly truncate at the base, brown, 3–4-septate Dendryphiopsis-like Present Terrestrial/On dead wood/Czech Republic Hawksworth (1985); Wijayawardene et al. (2014); Su et al. (2016)
K. emarceis 88–140 × 18–24 25–28 × 8–9 µm, biseriate, ellipsoidal, septum median to supra-median, dull green, becoming brown to dark brown at maturity, 1-septate, smooth Conidiophores 162–271 × 7–14 μm; Conidia 45–56 × 14–15 µm, oblong to clavate, grayish, brown to dark brown, 3–4(–5)-septate, constricted at septa, smooth Dendryphiopsis-like Present Terrestrial/On dead wood/Thailand Boonmee et al. (2012)
K. lignicola 107–163 × 19–28.5 27–30 × 10–12 µm, biseriate, ellipsoidal, 1(–2) septate, with median septum or in lower part, some ascospores with secondary septum, dull green, brown to dark brown at maturity Conidiophores 287–406 × 11–13 μm; Conidia 39–48 × 21–25 μm, obovoid to broadly, 1–2-septate, constricted at septa, smooth, dark brown, rounded at ends Dendryphiopsis-like Present Terrestrial/On decaying wood/Thailand Boonmee et al. (2012)
K. recessa 90 × 10 15–17.5 × 5–6.5 μm, elongate-ellipsoid, rounded at the apices, 1-septate, cells equal in size or the lower slightly smaller, slightly constricted at septum, pale brown, almost smooth or with a very weak verruculose ornamentation Conidiophores 3.5–4.0 μm wide, red–brown, thick-walled, septate Dendryphiopsis-like Absent Terrestrial/On rotten wood/USA Hawksworth (1985)
K. saprophytica 8–125 × 18–23 13–25(–40) × 7–11(–14) µm, ellipsoid, upper cell broader than lower cell, pale brown to dark brown, 1-septate, guttulate, smooth Conidiophores 90–216 × 8–12 μm; Conidiogenous cells monoblastic, terminal, cylindrical, brown to dark brown; Conidia 36–69 × 19–35 μm, cylindrical rounded at ends, 2–3 septa, dark brown to black, smooth Dendryphiopsis-like Present Terrestrial/On dead wood/Thailand de Farias et al. (2024)

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the National Natural Science Foundation of China (Nos. 32160006, 31970018).

Author contributions

Sampling: Y.F.H.; Fungal isolation: M.G.L.; Microscopy: M.G.L.; Description and phylogenetic analyses: X.X.L. and Y.F.H.; Writing – original draft preparation: X.X.L.; Writing – review and editing, X.G.Z., Z.H.X., and J.M. All authors read and approved the final manuscript.

Author ORCIDs

Ming-Gen Liao https://orcid.org/0009-0001-9537-1773

Xiu-Guo Zhang https://orcid.org/0000-0001-9733-8494

Zhao-Huan Xu https://orcid.org/0009-0008-2641-7783

Jian Ma https://orcid.org/0000-0001-9783-1860

Data availability

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

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Supplementary material

Supplementary material 1 

The concatenated ITS, LSU and SSU sequences

Xing-Xing Luo, Ming-Gen Liao, Ya-Fen Hu, Xiu-Guo Zhang, Zhao-Huan Xu, Jian Ma

Data type: fas

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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