Research Article
Research Article
Morpho-molecular diversity of Linocarpaceae (Chaetosphaeriales): Claviformispora gen. nov. from decaying branches of Phyllostachys heteroclada
expand article infoXiu-Lan Xu§, Chun-Lin Yang§, Rajesh Jeewon|, Dhanushka N. Wanasinghe, Ying-Gao Liu§, Qian-Gang Xiao
‡ Chengdu Academy of Agricultural and Forestry Sciences, Chengdu, China
§ Sichuan Agricultural University, Chengdu, China
| University of Mauritius, Reduit, Mauritius
¶ Kunming Institute of Botany, Kunming, China
Open Access


In this paper, Claviformispora gen. nov. in Linocarpaceae is introduced from Phyllostachys heteroclada in Sichuan Province, China. The new genus is characterised by its distinct morphological characters, such as ostiole with periphyses, asci with a thick doughnut-shaped, J- apical ring and clavate ascospore without septum-like band and appendage. Maximum Likelihood and Bayesian Inference phylogenetic analyses, based on DNA sequence data from ITS, LSU, SSU and TEF-1α regions, provide further evidence that the fungus is a distinct genus within this family. The new genus is compared with similar genera, such as Linocarpon and Neolinocarpon. Descriptions, illustrations and notes are provided for the new taxon.


bambusicolous fungi, one new genus and species, phylogeny, taxonomy


The order Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández (Sordariomycetes) was introduced in Sordariomycetidae O.E. Erikss. & Winka, based on LSU sequence data (Huhndorf et al. 2004) and currently comprises four families viz. Chaetosphaeriaceae Réblová, M.E. Barr & Samuels, Helminthosphaeriaceae Samuels, Cand. & Magni, Leptosporellaceae S. Konta & K.D. Hyde and Linocarpaceae S. Konta & K.D. Hyde (Hongsanan et al. 2017; Konta et al. 2017; Wijayawardene et al. 2020). Recently, 43 genera were accepted within Chaetosphaeriaceae and seven genera within Helminthosphaeriaceae (Hyde et al. 2020; Wijayawardene et al. 2020). Based on morphology and combined analyses of ITS and LSU sequence data, Konta et al. (2017) accommodated Linocarpon Syd. & P. Syd. and Neolinocarpon K.D. Hyde in Linocarpaceae and Leptosporella Penz. & Sacc. in Leptosporellaceae. Leptosporellaceae and Linocarpaceae are morphologically similar in their dome-shaped ascomata and filiform ascospores (Hyde and Alias 1999; Cai et al. 2004; Konta et al. 2017). The former, however, can be delineated based on ascospores that are narrow, long filiform, with gradually tapering ends and indistinct mucilage (if present), whereas in Linocarpaceae, ascospores have a distinct appendage at the apex and are generally wider and differ in appearance at the ends (Konta et al. 2017).

The genus Leptosporella was introduced with L. gregaria Penz. & Sacc., 1897 as the type species by Penzig and Saccardo (1897). Lumbsch and Huhndorf (2010) referred it in Sordariomycetidae genera incertae sedis. Subsequently, the genus was referred to the Chaetosphaeriales, based on phylogenetic analysis of LSU sequence data (Huhndorf and Miller 2011; Dai et al. 2016; Wijayawardene et al. 2020). At present, 15 epithets of Leptosporella are recorded in Index Fungorum ( Sydow and Sydow (1917) introduced Linocarpon with L. pandani Syd. & P. Syd., 1917 as the type species. Hyde (1992a) introduced Neolinocarpon to accommodate a linocarpon-like species, N. globosicarpum K.D. Hyde, 1992. Currently in Index Fungorum (2020), 44 and 13 epithets are accommodated in Linocarpon and Neolinocarpon, respectively. Hyde (1997) and Jeewon et al. (2003) reported that Linocarpon and Neolinocarpon can be accommodated in Hyponectriaceae (Xylariales), while Bahl (2006) suggested a closer relationship with Chaetosphaeriales and Xylariales, based on their molecular data. However, with more taxon sampling and fresh collections, Konta et al. (2017) confirmed that Linocarpon and Neolinocarpon should be accommodated in a distinct family (Linocarpaceae) in Chaetosphaeriales.

The present research is a part of our investigations on the taxonomic and phylogenetic circumscriptions of pathogenic and saprobic micro-fungi associated with bamboo in Sichuan Province, China. In this paper, we introduce a new genus Claviformispora in Linocarpaceae, typified by C. phyllostachydis from Phyllostachys heteroclada Oliv., 1894 (Poaceae). The morphological differences and analyses of a combined ITS, LSU, SSU and TEF-1α sequence dataset support the validity of the new genus and its placement in Linocarpaceae. The new genus is compared with other genera in the family. The comprehensive descriptions and micrographs of new taxa are provided.

Materials and methods

Specimen collection and morphological study

Bamboo materials were collected from Ya’an City, China. Single ascospore isolations were carried out following the method described by Chomnunti et al. (2014) and the germinating spores were transferred to PDA, incubated at 25 °C in the dark and cultural characteristics were determined. Ascomata were observed and photographed using a dissecting microscope NVT-GG (Shanghai Advanced Photoelectric Technology Co. Ltd, China) matched to a VS-800C micro-digital camera (Shenzhen Weishen Times Technology Co. Ltd., China). The anatomical details were visualised using Nikon ECLIPSE Ni compound microscope fitted to a Canon 600D digital camera and an OPTEC BK-DM320 microscope matched to a VS-800C micro-digital camera (Shenzhen Weishen Times Technology Co. Ltd., China). Iodine reaction of the ascus wall was tested in Melzer’s reagent (MLZ). Lactate cotton blue reagent was used to observe the number of septa. The gelatinous appendage was observed in Black Indian ink. Type specimens were deposited at the Herbarium of Sichuan Agricultural University, Chengdu, China (SICAU) and Mae Fah Luang University Herbarium (MFLU). The ex-type living cultures are deposited at the Culture Collection in Sichuan Agricultural University (SICAUCC) and the Culture Collection at Mae Fah Luang University (MFLUCC). Index Fungorum numbers ( are registered and provided.

DNA extraction, PCR amplification and DNA sequencing

Total genomic DNA was extracted from mycelium that were grown on PDA at 25 °C for two weeks using a Plant Genomic DNA extraction kit (Tiangen, China) following the manufacturer’s instructions. The primers pairs LR0R and LR5 (Vilgalys and Hester 1990), NS1 and NS4, ITS5 and ITS4 (White et al. 1990), EF1-983F and EF1-2218R (Rehner 2001) were used for the amplification of the partial large subunit nuclear rDNA (LSU), the partial small subunit nuclear rDNA (SSU), internal transcribed spacers (ITS) and translation elongation factor 1-alpha (TEF-1α), respectively.

Polymerase chain reaction (PCR) was performed in 25 μl final volumes containing 22 μl of Master Mix (Beijing TsingKe Biotech Co. Ltd.), 1 μl of DNA template, 1 μl of each forward and reverse primers (10 μM). The PCR thermal cycle programmes for LSU, SSU, ITS and TEF1-α gene were amplified as: initial denaturation 94 °C for 3 minutes, followed by 35 cycles of denaturation at 94 °C for 30 seconds, annealing at 55 °C for 50 seconds, elongation at 72 °C for 1 minute and final extension at 72 °C for 10 minutes. PCR products were sequenced with the above-mentioned primers at TsingKe Biological Technology Co. Ltd, Chengdu, China. The newly-generated sequences from the LSU, SSU, TEF-1α and ITS regions were deposited in GenBank (Table 1).

Phylogenetic analyses

Taxa to be used for phylogenetic analyses were selected, based on results generated from nucleotide BLAST searches online in GenBank and recent publications (Lu et al. 2016; Konta et al. 2017; Senwanna et al. 2018; Wei et al. 2018; Lin et al. 2019). Gelasinospora tetrasperma (CBS 178.33) and Sordaria fimicola (CBS 508.50) were selected as the outgroup taxa. The sequences were downloaded from GenBank ( and the accession numbers are listed in Table 1. A combined ITS, LSU, SSU and TEF-1α sequence dataset was used to construct the phylogenetic tree. DNA alignments were performed by using MAFFT v.7.429 online service (Katoh et al. 2019) and ambiguous regions were excluded with BioEdit version (Hall 1999). Multigene sequences were concatenated by using Mesquite software (Maddison and Maddison 2019). Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were performed. The best nucleotide substitution model was determined by MrModeltest v. 2.2 (Nylander 2004).

Maximum Likelihood analysis and Bayesian Inference analysis were generated by using the CIPRES Science Gateway web server (Miller 2010). RAxML-HPC2 on XSEDE (8.2.10) (Stamatakis 2014) with GTR+GAMMA substitution model with 1000 bootstrap iterations was chosen for Maximum Likelihood analysis. For BI analyses, the best-fit model GTR+I+G for ITS, LSU and SSU was selected in MrModeltest 2.2 and GTR+G for TEF. The analyses were computed with six simultaneous Markov Chain Monte Carlo (MCMC) Chains with 8,000,000 generations and a sampling frequency of 100 generations. The burn-in fraction was set to 0.25 and the run automatically ended when the average standard deviation of split frequencies reached below 0.01.

Phylogenetic trees were visualised with FigTree v.1.4.3 (Rambaut and Drummond 2016) and edited using Adobe Illustrator CS6 (Adobe Systems Inc., United States). Maximum Likelihood bootstrap values (MLBP) equal to or greater than 70% and Bayesian Posterior Probabilities (BYPP) equal to or greater than 0.95 were accepted. The finalised alignment and tree were deposited in TreeBASE (, submission ID: 25996. The new taxa introduced follow the recommendations of Jeewon and Hyde (2016).


Phylogenetic analyses

Phylogenetic analyses of a combined dataset (ITS, LSU, SSU, TEF-1α) comprises 51 taxa within the order Chaetosphaeriales (Table 1), including 24 taxa in family Chaetosphaeriaceae, nine taxa in Helminthosphaeriaceae, ten taxa in Linocarpaceae, six taxa in Leptosporellaceae and two outgroup taxa in Sordariales. The dataset consisted of 5,849 characters including gaps (LSU = 1,571, ITS = 736, SSU = 2,522, TEF = 1,020). The best scoring tree of RAxML analysis is shown in Fig. 1, with the support values of ML and BI analyses.

Table 1.

Molecular data used in this study and GenBank accession numbers.

Species name Strain GenBank accession number
Chloridium aquaticum MFLUCC 11-0212 MH476567 MH476570 MH476573
Chloridium aseptatum MFLUCC 11-0216 MH476568 NR_158365 MH476574
Claviformispora phyllostachydis SICAUCC 16-0004 MT232720 MT232736 MT232735 MT240855
Cryptophiale hamulata MFLUCC 18-0098 MG386756 MG386757
Cryptophiale udagawae MFLUCC 18-0422 MH758211 MH758198 MH758205
MFLUCC 18-0428 MH758210 MH758197 MH758204
Dictyochaeta siamensis MFLUCC 15-0614 KX609952 KX609955
Dictyochaeta assamica CBS 242.66 MH870426 MH858788
Dictyochaeta pandanicola MFLUCC 17-0563 MH376710 MH388338 MH388307 MH388373
Dictyochaeta terminalis GZCC 18-0085 MN104624 MN104613 MN104633
Echinosphaeria canescens SMH4666 KF765605
SMH4791 AY436403
Endophragmiella dimorphospora FMR_12150 KY853502 KY853442 HF937351
Gelasinospora tetrasperma CBS 178.33 DQ470980 NR_077163 DQ471032 DQ471103
Helminthosphaeria clavariarum SMH4609 AY346283
Hilberina caudata SMH1542 KF765615
Infundibulomyces cupulata BCC11929 EF113979 EF113976 EF113982
Infundibulomyces oblongisporus BCC13400 EF113980 EF113977 EF113983
Kionochaeta castaneae GZCC 18-0025 MN104621 MN104610 MN104630
Kionochaeta microspora GZCC 18-0036 MN104618 MN104607 MN104627
Leptosporella arengae MFLUCC 15-0330 MG272246 MG272255 MG366594 MG272259
Leptosporella bambusae MFLUCC 12-0846 KU863122 KU940134
Leptosporella cocois MFLUCC 15-0816 MG272256
Leptosporella gregaria SMH4290 AY346290
SMH4673 HM171287
Leptosporella elaeidis MFLU 19-0669 MK659772 MK659767 MK659774 MN883560
Linocarpon arengae MFLUCC 15-0331 MG272247 MG366596
Linocarpon cocois MFLUCC 15-0812 MG272248 MG272257 MG272253
Menispora tortuosa DAOM 231154 AY544682 KT225527 AY544723
CBS 214.56 AF178558 AF178558
Menisporopsis anisospora CBS 109475 MH874421 MH862827
Menisporopsis breviseta GZCC 18-0071 MN104623 MN104612 MN104632
Menisporopsis dushanensis GZCC 18-0084 MN104626 MN104615 MN104635
Menisporopsis pandanicola KUMCC 17-0271 MH376726 MH388353 MH388320 MH388388
Menisporopsis theobromae MFLUCC 15-0055 KX609954 KX609957
Neolinocarpon arengae MFLUCC 15-0323 MG272249 MG272258 MG366597
Neolinocarpon rachidis MFLUCC 15-0332 MG272250 MG366598
MFLUCC 15-0814a MK106353 MK106342 MK106367
MFLUCC 15-0814b MK106354 MK106368
Neolinocarpon phayaoense MFLUCC 17-0073a MG581933 MG581936 MG739512
MFLUCC 17-0073b MG581934 MG581937 MG739513
MFLUCC 17-0074 MG581935 MG581938 MG739514
Phialosporostilbe scutiformis MFLUCC 17-0227 MH758207 MH758194 MH758201
MFLUCC 18-1288 MH758212 MH758199
Ruzenia spermoides SMH4606 AY436422
SMH4655 KF765619
Synaptospora plumbea ANM963 KF765620
SMH3962 KF765621
Sordaria fimicola CBS 508.50 MH868251 MH856730
Zanclospora iberica FMR 11584 KY853544 KY853480 HF937360
FMR 12186 KY853545 KY853481 HF937361

The best scoring RAxML tree with the final optimisation had a likelihood value of -26,415.700648. The matrix had 1,751 distinct alignment patterns and 64.64% in this alignment is the gaps and completely undetermined characters. Estimated base frequencies were as follows: A = 0.236065, C = 0.261532, G = 0.295313, T = 0.207091, with substitution rates AC = 1.062535, AG = 1.855434, AT = 0.940219, CG = 1.052604, CT = 4.590285, GT = 1.000000. The gamma distribution shape parameter α = 0.311923 and the Tree-Length = 2.281738. The Bayesian analysis resulted in 20,502 trees after 8,000,000 generations. The first 25% of trees (1,624 trees), which represent the burn-in phase of the analyses were discarded, while the remaining 4,878 trees were used for calculating posterior probabilities. Bayesian posterior probabilities were evaluated by MCMC with a final average standard deviation of split frequencies = 0.009877.

Phylogenetic trees generated from Maximum Likelihood (ML) and Bayesian Inference analyses were similar in overall topologies. Phylogeny from the combined sequence data analysis indicates that all families were monophyletic with strong bootstrap support values (Fig. 1). Phylogenetic results show that our novel species Claviformispora phyllostachydis (SICAUCC 16-0004) belongs to family Linocarpaceae with 91% ML and 1.00 BYPP support and close to genera Neolinocarpon and Linocarpon (Fig. 1). The new genus Claviformispora constituted a distinct lineage in the family Linocarpaceae (Fig. 1).

Figure 1. 

Phylogram of RAxML analysis based on a combined ITS, LSU, SSU and TEF-1α sequence dataset within order Chaetosphaeriales. Bootstrap support values for maximum likelihood (ML, left) greater than 70% and Bayesian posterior probabilities (PP, right) equal to or greater than 0.95 are indicated at the nodes. The tree is rooted to Gelasinospora tetrasperma (CBS 178.33) and Sordaria fimicola (CBS 508.50). All sequences from ex-type strains are in bold. The newly-generated sequence is in red.


Linocarpaceae S. Konta & K.D. Hyde, Mycosphere 8(10): 1962 (2017) emend.

Type genus

Linocarpon Syd. & P. Syd.


Saprobic and endophytic fungi on monocotyledons and rarely dicotyledons. Sexual morph: Ascomata solitary or aggregated, superficial or immersed comprising black, dome-shaped or subglobose, slightly raised blistering areas with a central ostiole or immersed with a black shiny papilla. Peridium composed of dark brown to black cells of textura angularis. Hamathecium comprising septate paraphyses that are longer than asci, wider at the base, tapering towards the apex. Asci 8-spored, unitunicate, cylindrical, with a J-, apical ring, developing from the base and periphery of the ascomata. Ascospores parallel or spiral in asci, hyaline or pale yellowish in mass, filiform or claviform, straight or curved, unicellular with or without refringent bands, with or without polar appendages. Asexual morph: Phialophora-like spp. were found in Linocarpon appendiculatum and L. elaeidis cultures (Hyde 1992b), but no records are available for other species.


Linocarpaceae was introduced as a new family to accommodate Linocarpon and Neolinocarpon species, based on morphology and phylogeny (Konta et al. 2017). Appressoria were first recorded from Neolinocarpon rachidis (Hyde et al. 2019). The new genus Claviformispora, which is well-supported within Linocarpaceae suggests that there is a need to amend the morphological circumscriptions of the family given that the ascomata (subglobose) and ascospore (claviform) characters are so different from the other two genera.

Claviformispora X. L. Xu & C. L. Yang, gen. nov.

Type species

Claviformispora phyllostachydis X. L. Xu & C. L. Yang


Name reflects the claviform ascospores.


Saprobic on dead branches. Sexual morph: Stromata solitary or gregarious, black, erumpent. Ascomata solitary or aggregated, immersed, subglobose, slightly raised blistering areas with a central ostiole with periphyses. Peridium outer cells merging with the host tissues, composed of pale to dark brown cells of textura angularis. Hamathecium comprising hyaline, septate paraphyses, longer than asci, wider at the base, tapering towards the apex. Asci 8-spored, cylindrical to cylindric-clavate, unitunicate, short pedicellate, apically rounded, with a doughnut-shaped, refractive, J- apical ring. Ascospores overlapping uniseriate or 2-seriate, clavated with a thin pedicellate, 1-celled, hyaline, without appendage and refringent bands, smooth-walled. Asexual morph: Undetermined.


Claviformispora resembles Neolinocarpon in having immersed ascomata and ostiole with periphyses, but differs in forming aggregated ascomata, cylindric-clavate, short pedicellate asci, clavate ascospores with thin pedicel and without septa-like bands and appendages, whereas the ascospores of Neolinocarpon and Linocarpon (Linocarpaceae) species are usually filiform with refringent bands and appendages (Hyde 1992b, 1997; Konta et al. 2017). The nature of the ascospore appendages appears to be phylogenetically significant for intergeneric delineation as has been seen in other studies (Poonyth et al. 2000; Jeewon et al; 2003, Thongkantha et al. 2003; Cai et al. 2004; Konta et al. 2017), but this warrants further investigations with more sampling and fresh collections of Neolinocarpon and Linocarpon. Differences in morphology between these genera in Linocarpaceae are summarised in Table 2.

Table 2.

Morphological comparison of Linocarpon, Neolinocarpon and Claviformispora.

Morphology Linocarpon Neolinocarpon Claviformispora
(Type: L. pandani) (Type: N. globosicarpum) (Type: C. phyllostachydis)
Stromata Absent Absent Solitary or aggregated, comprising elliptical areas and large black areas, with slit-like openings
Ascomata Solitary, superficial, subglobose, comprising black, dome-shaped, raised blistering areas, central ostiole Solitary, deeply immersed, oval to globose, with central raised, dark, shiny papilla, central ostiole with periphyses Solitary or aggregated, deeply immersed, subglobose, slightly raised blistering areas, central ostiole with periphyses
Peridium Textura angularis Textura angularis Textura angularis
Hamathecium Hyaline, septate paraphyses, longer than asci Hyaline, septate paraphyses, longer than asci Hyaline, septate paraphyses, longer than asci
Asci Cylindrical, unitunicate, a small non-amyloid apical ring Long cylindrical, pedicellate, unitunicate, an oblong to wedge-shaped, refractive, apical ring and some with a refractive circular body below Cylindrical to cylindric-clavate, unitunicate, pedicellate, doughnut-shaped, refractive, J- apical ring
Ascospores Filiform, aseptate, hyaline or pale-yellowish in mass, parallel or spiral, with appendage and refringent septum-like bands or absent Filiform, aseptate, hyaline or pale-yellowish in mass, parallel or spiral, with apical appendages and refringent bands or absent Clavate, thin pedicellate, aseptate, hyaline, parallel, no appendage and refringent band
Asexual morph Only found in L. appendiculatum and L. elaeidis, conidiophore arising from the aerial mycelium, conidiogenous cells phialidic, smooth, translucent brown, conidia clavate to fusiform, straight or slightly curved or slightly sinuous, unicellular, smooth, colourless Undetermined Undetermined
Others Colonies on MEA and PDA growing slowly Colonies on MEA growing slowly. Ascospores on MEA produced appressoria-like structures at each tip of germ tube, only found in N. rachidis Colonies on PDA grow faster
References Hyde (1992b), Konta et al. (2017), Thongkantha et al. (2003) Hyde et al. (2019), Senwanna et al. (2018), Hyde et al. (1998) This study

Claviformispora phyllostachydis X. L. Xu & C. L. Yang, sp. nov.

Fig. 2


China, Sichuan Province, Ya’an City, Yucheng Distinct, Kongping Township, alt. 1133 m, 29°50.14'N, 103°03'E, on dead branches of Phyllostachys heteroclada Oliv. (Poaceae), 11 December 2016, C. L. Yang and X. L. Xu, YCL201612002 (SICAU 16-0007, holotype; MFLU 18-1217, isotype), ex-type living culture, SICAUCC 16-0004 = MFLUCC 18-1230.

Figure 2. 

Claviformispora phyllostachydis (SICAU 16-0007, holotype) a, b Stromata on host substrate c section through ascoma with ascomata d ostiole with periphyses e peridium f paraphyses g–j asci k–o ascospores p germinated ascospore q, r colony on PDA after 7 days. Scale bars: 2 mm (a), 500 μm (b), 100 μm (c), 20 μm (d, e), 10 μm (f–p).


The specific epithet refers to the host genus Phyllostachys.


Saprobic on dead branches of Phyllostachys heteroclada Oliv. Sexual morph: Stromata solitary comprising elliptical areas or aggregated in large black areas, slightly raised with slit-like openings presenting on host surface. Ascomata 120–240 μm high × 220–490 μm diameter (x̄ = 189 × 345 μm, n = 20), perithecial, immersed, central, papillate ostiole with periphyses, oval-globose in section, the cells between the perithecia are composed with tissue of stromata and host. Peridium 20–40 μm wide (x̄ = 33 μm, n = 10), outer cells merging with the host tissues, composed of pale to dark brown cells of textura angularis. Hamathecium comprising hyaline, hypha-like, septate paraphyses, occasionally branched, longer than asci, wider at the base, 2–4 μm diameter (x̄ = 2.7 μm, n = 20) tapering towards the apex, 0.78–1.20 μm diameter (x̄ = 0.98 μm, n = 20). Asci 90–160 × 9–15 μm (x̄ = 118 × 13 μm, n = 20), 8-spored, cylindrical to cylindric-clavate, unitunicate, short pedicellate, apically rounded, with a massive, doughnut-shaped, refractive, J- reaction, apical ring. Ascospores 35–50 × 5.7–8.6 μm (x̄ = 45.7 × 7.0 μm, n = 40), overlapping uniseriate or 2-seriate, claviform typically with a thin pedicel, aseptate, hyaline, straight or slight curved, without appendage and septum-like bands, guttulate when maturity. Asexual morph: Undetermined.

Culture characters

Ascospores germinated on PDA within 12 hours at both ends. Colonies on PDA reaching 5 cm diameter after 7 days at 25 °C, white to grey with strong radiations outwards on forward side. Colonies became dark brown and black on the reverse after a long time of cultivation. The hyphae are septate, branched, smooth.


This study establishes a new genus and also provides further insights into the phylogeny of members associated with Linocarpaceae. Morphologically-based examinations of Claviformispora (as discussed above) clearly show that the morphological circumscriptions (familial concept) of species should be broadened and possibly indicate that this family is much more diverse than expected. Our collection can be clearly distinguished from other groups of similar fungi in Linocarpaceae with its interesting ascospore morphology. In addition, we also noted some peculiarities in the DNA sequences we analysed. A comparison of ITS sequences, based on BLAST reveals 34%, 26% and 30% base pair differences with L. cocois (MFLUCC 15-0812), N. arengae (MFLUCC 15-0323) and N. rachidis (MFLUCC 15-0814a), respectively. There are more than 9% and 5% sequence differences with the three taxa when the LSU and SSU rDNA sequences were compared respectively. Following the guidelines recommended by Jeewon and Hyde (2016), there are therefore sufficient grounds to establish a new species at the genus rank.

Species of Linocarpaceae have been found on Arecaceae, Poaceae, Euphorbiaceae, Zingiberaceae, Pandanaceae, Fagaceae, Fabaceae and Smilacaceae, including Arenga, Attalea, Calamus, Trachycarpus, Acrocomia, Archontophoenix, Cocos, Daemonorops, Licuala, Livistona, Plectocomia, Phoenix, Raphia, Sabal, Mauritia, Nypa, Elaeis, Pinanga, Eugeissona, Pennisetum, Gramineae, Stipa, unidentified bamboo, Hevea, Manihot, Alpinia, Pandanus, Quenrcus, Cajanus and Smilax (Hyde 1988, 1992a, b; Dulymamode et al. 1998; Hyde et al. 1998; Hyde and Alias 1999; Thongkantha et al. 2003; Cai et al. 2004; Bhilabutra et al. 2006; Vitoria et al. 2013; Konta et al. 2017; Senwanna et al. 2018). More than 50% of the species were recorded from hosts of the Arecaceae. Species in Linocarpaceae are mostly saprobic, except Linocarpon palmetto which was discovered as a pathogen of Sabal palmetto in Florida (Barr 1978). Four species in Linocarpaceae from Poaceae have been reported so far, including Neolinocarpon penniseti on Pennisetum purpureum (Bhilabutra et al. 2006), Linocarpon williamsii on Gramineae sp. (Hansford 1954), L. stipae on Stipa sp. (Hansford 1954) and L. bambusicola on unidentified bamboo submerged in a river (Cai et al. 2004).

Phyllostachys heteroclada, mainly a food source and use as a material in the weaving industry, is distributed along the Yellow River Valley and the southern Provinces in China. It is common in the mountainous areas of Sichuan Province with distribution up to 1,500 m above sea level (Yi 1997; Yi et al. 2008). There is a large area of pure forest in Yibin, Leshan and Ya’an Cities and sporadic distribution in other areas. According to preliminary statistics, bambusicolous fungi from seven orders (excluding fungi referred to as Sordariomycetes incertae sedis) have been recorded on P. heteroclada, including Hypocreales, Ostropales, Pleosporales, Phyllachorales, Pucciniales, Ustilaginales and Xylariales, of which Pleosporales is the largest one. Most bambusicolous fungi in China were recorded with inadequate morphological descriptions or molecular data. The early known fungi on P. heteroclada are documented as Aciculosporium take, Ellisembia pseudoseptata, Fusarium oxysporum, F. semitectum, Phyllachora gracilis, Ph. orbicular, Shiraia bambusicola, Stereostratum corticioides and Ustilago shiraiana (Zhou et al. 2001; Xu et al. 2006). In recent years, some new records and taxa, viz. Bambusicola subthailandica, B. sichuanensis, Neostagonosporella sichuanensis, Parakarstenia phyllostachydis, Phyllachora heterocladae, Podonectria sichuanensis, Arthrinium yunnanum and A. phyllostachium have been reported (Yang et al. 2019a, b, c, d, e, f). Here, we introduce a new genus in order Chaetosphaeriales, which is a contribution to fungal diversity on P. heteroclada.


We all thank Dr. Shaun Pennycook for his help with the nomenclature of the novel species. Xiu-Lan Xu thanks the Chengdu Science and Technology Bureau for funding her research. Ying-Gao Liu and Chun-Lin Yang acknowledge the “Study on identification of pathogenic fungi and prevention techniques for witches’ broom of Phyllostachys violascens” (fund from the Bureau of Economic Information and Science & Technology of Yucheng District in Ya’an City) for supporting this work. Dhanushka Wanasinghe would like to thank CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (number 2019PC0008) and the National Science Foundation of China and the Chinese Academy of Sciences (grant no.: 41761144055). R Jeewon thanks University of Mauritius for support.


  • Bahl J (2006) Molecular evolution of three morphologically similar families in the Xylariomycetidae (Apiosporaceae, Clypeosphaeriaceae, Hyponectriaceae). PhD Thesis, The University of Hong Kong, China.
  • Barr ME (1978) The Diaporthales in North America: with emphasis on Gnomonia and its segregates. Mycologia Memoir 7: 1–232.
  • Bhilabutra W, Lumyong S, Jeewon R, McKenzie EHC, Hyde KD (2006) Neolinocarpon penniseti sp. nov. on the grass Pennisetum purpureum (Poaceae). Cryptogamie, Mycologie 27(4): 305–310.
  • Chomnunti P, Hongsanan S, Hudson BA, Tian Q, Peršoh D, Dhami MK, Alias AS, Xu J, Liu X, Stadler M, Hyde KD (2014) The sooty moulds. Fungal Diversity 66(1): 1–36.
  • Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
  • Hansford CG (1954) Australian Fungi. II. New species and revisions. Proceedings of the Linnean Society of New South Wales 79: 97–141.
  • Hongsanan S, Maharachchikumbura SSN, Hyde KD, Samarakoon MC, Jeewon R, Zhao Q, Al-sadi AM, Bahkali AH (2017) An updated phylogeny of Sordariomycetes based on phylogenetic and molecular clock evidence. Fungal Diversity 84(1): 25–41.
  • Huhndorf SM, Miller AN (2011) A molecular re-appraisal of taxa in the Sordariomycetidae and a new species of Rimaconus from New Zealand. Studies in Mycology 68: 203–210.
  • Hyde KD (1992b) Fungi from palms. I. The genus Linocarpon, a revision. Sydowia 44: 32–54.
  • Hyde KD (1988) The genus Linocarpon from the mangrove palm Nypa fruticans. Transactions of the Mycological Society of Japan 29: 339–350.
  • Hyde KD, Alias SA (1999) Linocarpon angustatum sp. nov., and Neolinocarpon nypicola sp. nov. from petioles of Nypa fruticans, and a list of fungi from aerial parts of this host. Mycoscience 40(2): 145–149.
  • Hyde KD, Norphanphoun C, Maharachchikumbura SSN, Bhat DJ, Jones EBG, Bundhun D, Chen YJ, Bao DF, Boonmee S, Calabon MS, Chaiwan N, Chethana KWT, Dai DQ, Dayarathne MC, Devadatha B, Dissanayake AJ, Dissanayake LS, Doilom M, Dong W, Fan XL, Goonaselara ID, Hongsanan S, Huang SK, Jayawardena RS, Jeewon R, Karunarathna A, Konta S, Kumar V, Lin CG, Liu JK, Liu NG, Luangsa-ard J, Lumyong S, Luo ZL, Marasinghe DS, McMenzie EHC, Niego AGT, Niranjan M, Perera RH, Phukhamsakda C, Rathnayaka AR, Samarakoon MC, Samarakoon SMBC, Sarma VV, Senanayake IC, Shang QJ, Stadler M, Tibpromma S, Wanasinghe DN, Wei DP, Wijayawardene NN, Xiao YP, Yang J, Zeng XY, Zhang SN, Xiang MM (2020) Refined families of Sordariomycetes. Mycosphere 11(1): 305–1059.
  • Hyde KD, Taylor JE, Fröhlich J (1998) Fungi from palms. XXXIV. The genus Neolinocarpon with five new species and one new combination. Fungal Diversity 1: 115–131.
  • Hyde KD, Tennakoon DS, Jeewon R, Bhat DJ, Maharachchikumbura SSN, Rossi W, Leonardi M, Lee HB, Mum HY, Houbraken J, Nguyen TTT, Jeon SJ, Frisvad JC, Wanasinghe DN, Lücking R, Aptroot A, Cáceres MES, Karunarathna SC, Hongsanan S, Phookamsak R, de Silva NI, Thambugala KM, Jayawardena RS, Senanayake IC, Boonmee S, Chen J, Luo ZL, Phukhamsakda C, Pereira OL, Abreu VP, Rosado AWC, Bart B, Randrianiohany E, Hofstetter V, Gibertoni TB, da Silva Soares AM, Plautz Jr HL, Sotão HMP, Xavier WKS, Bezerra JDP, de Oliveira TGL, de Souza-Motta CM, Magalhães OMC, Bundhun D, Harishchandra D, Manawasinghe IS, Dong W, Zhang SN, Bao DF, Samarakoon MC, Pem D, Karunarathna A, Lin CG, Yang J, Perera RH, Kumar V, Huang SK, Dayarathne MC, Ekanayaka AH, Jayasiri SC, Xiao YP, Konta S, Niskanen T, Liimatainen K, Dai YC, Ji XH, Tian XM, Mešić A, Singh SK, Phutthacharoen K, Cai L, Sorvongxay T, Thiyagaraja V, Norphanphoun C, Chaiwan N, Lu YZ, Jiang HB, Zhang JF, Abeywickrama PD, Aluthmuhandiram JVS, Brahmanage RS, Zeng M, Chethana T, Wei DP, Réblová M, Fournier J, Nekvindová J, do Nascimento Barbosa R, dos Santos JEF, de Oliveira NT, Li GJ, Ertz D, Shang QJ, Phillips AJL, Kuo CH, Camporesi E, Bulgakov TS, Lumyong S, Jones EBG, Chomnunti P, Gentekaki E, Bungartz F, Zeng XY, Fryar S, Tkalčec Z, Liang JM, Li GS, Wen TC, Singh PN, Gafforov Y, Promputtha I, Yasanthika E, Goonasekara ID, Zhao RL, Zhao Q, Kirk PM, Liu JK, Yan JY, Mortimer PE, Xu JC, Doilom M (2019) Fungal diversity notes 1036–1150: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 96: 1–242.
  • 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 SC, Liu JK, Luangsa-ard JJ, Lumbsch HT, Maharachchikumbura SSN, McKenzie EHC, Moncalvo JM, Ghobad-Nejhad M, Nilsson H, Pang KL, Pereira O, 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.
  • Jeewon R, Hyde KD (2016) Establishing species boundaries and new taxa among fungi: recommendations to resolve taxonomic ambiguities. Mycosphere 7(11): 1669–1677.
  • Jeewon R, Liew ECY, Hyde KD (2003) Molecular systematics of the Amphisphaeriaceae based on cladistic analyses of partial LSU rDNA gene sequences. Mycological Research 107(12): 1392–1402.
  • Jeewon R, Liew ECY, Simpson JA, Hodgkiss IJ, Hyde KD (2003) Phylogenetic significance of morphological characters in the taxonomy of Pestalotiopsis species. Molecular Phylogenetics and Evolution 27(3): 372–383.
  • Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20: 1160–1166.
  • Konta S, Hongsanan S, Liu JK, Eungwanichayapant PD, Jeewon R, Hyde KD, Maharachchikumbura SSN, Boonmee S (2017) Leptosporella (Leptosporellaceae fam. nov.) and Linocarpon and Neolinocarpon (Linocarpaceae fam. nov.) are accommodated in Chaetosphaeriales. Mycosphere 8(10): 1943–1974.
  • Lin CG, McKenzie EHC, Liu JK, Jones EBG, Hyde KD (2019) Hyaline-spored chaetosphaeriaceous hyphomycetes from Thailand and China, with a review of the family Chaetosphaeriaceae. Mycosphere 10(1): 655–700.
  • Lu YZ, Liu KJ, Hyde KD, Bhat DJ, Xiao YP, Tian Q, Wen TC, Boonmee S, Kang JC (2016) Brunneodinemasporium jonesii and Tainosphaeria jonesii spp. nov. (Chaetosphaeriaceae, Chaetosphaeriales) from southern China. Mycosphere 7(9): 1322–1331.
  • Lumbsch HT, Huhndorf SM (2010) Myconet (Vol. 14). Part One. Outline of Ascomycota–2009. Part Two. Notes on Ascomycete Systematics. Nos. 4751–5113. Fieldina Life and Earth Sciences 1: 1–64.
  • Miller MA, Pfeiffer WT, Schwartz T (2010) Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Tree. Gateway Computing Environments Workshop (GCE), 8 pp.
  • Poonyth AD, Hyde KD, Wong SW, Peerally A (2000) Ultrastructure of asci and ascospore appendages in Linocarpon appendiculatum and L. nypae. Botanica Marina 43(3): 213–221.
  • Senwanna C, Phookamsak R, Bahkali AH, Elgorban AM, Cheewangkoon R, Hyde KD (2018) Neolinocarpon phayaoense sp. nov. (Linocarpaceae) from Thailand. Phytotaxa 362(1): 077–086.
  • Sydow H, Sydow P (1917) Beitrag zur Kenntniss der Pilzflora der Philippinen-Inseln. Annales Mycologici 15: 165–268.
  • Thongkantha S, Lumyong S, Lumyong P, Whitton SR, McKenzie EHC, Hyde KD (2003) Microfungi on the Pandanaceae: Linocarpon lammiae sp. nov., L. siamensis sp. nov. and L. suthepensis sp. nov. are described with a key to Linocarpon species from the Pandanaceae. Mycologia 95(2): 360–367.
  • 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.
  • Vitoria NS, Cavalcanti MAQ, Santos CDD, Pereira J, Bezerra JL (2013) Neolinocarpon attaleae sp nov on Attalea funifera (Arecaceae) from Brazil. Mycotaxon 123(1): 141–145.
  • Wei MJ, Zhang H, Dong W, Boonmee S, Zhang D (2018) Introducing Dictyochaeta qauatica sp. nov. and two new species of Chloridium (Chaetosphaeriaceae, Sordariomycetes) from aquatic habitat. Phytotaxa 362(2): 187–199.
  • Wijayawardene N, Hyde KD, Al-Ani LKT, Tedersoo L, Haelewaters D, Rajeshkumar KC, Zhao RL, Aptroot A, Leontyev DV, Saxena RK, Tokarev YS, Dai DQ, Letcher PM, Stephenson SL, Ertz D, Lumbsch HT, Kukwa M, Issi IV, Madrid H, Phillips AJL, Selbmann L, Pfliegler WP, Horváth E, Bensch K, Kirk P, Kolaříková Z, Raja HA, Radek R, Papp V, Dima B, Ma J, Malosso E, Takamatsu S, Rambold G, Gannibal PB, Triebel D, Gautam AK, Avasthi S, Suetrong S, Timdal E, Fryar SC, Delgado G, Réblová M, Doilom M, Dolatabadi S, Pawłowska J, Humber RA, Kodsueb R, Sánchez-Castro I, Goto BT, Silva DKA, de Souza FA, Oehl F, da Silva GA, Silva IR, Błaszkowski J, Jobim K, Maia LC, Barbosa FR, Fiuza PO, Divakar PK, Shenov BD, Castañeda-Ruiz RF, Somrithipol S, Karunarathna SC, Tibpromma S, Mortimer PE, Wanasinghe DN, Phookamsak R, Xu JC, Wang Y, Tian FH, Alvarado P, Li DW, Kušan I, Matočec N, Maharachchikumbura SSN, Papizadeh M, Heredia G, Wartchow F, Bakhshi M, Boehm E, Youssef N, Hustad VP, Lawrey JD, de A Santiago ALCM, Tian Q, Houbraken J, Hongsanan S, Tanaka K, Dissanayake AJ, Monteiro JS, Grossart HP, Suija A, Weerakoon G, Etayo J, Tsurykau A, Kuhnert E, Vázquez V, Mungai P, Damm U, Li QR, Zhang H, Boonmee S, Lu YZ, Becerra AG, Kendrick B, Brearley FQ, Motiejūnaitė J, Sharma B, Khare R, Gaikwad S, Wijesundara DSA, Tang LZ, He MQ, Flakus A, Rodriguez-Flakus P, Zhurbenko MP, McKenzie EHC, Stadler M, Bhat DJ, Liu JK, Raza M, Jeewon R, Nassonova ES, Prieto M, Ja yalal RGU, Yurkov A, Schnittler M, Shchepin ON, Novozhilov YK, Liu P, Cavender JC, Kang YQ, Mohammad S, Zhang LF, Xu RF, Li YM, Dayarathne MC, Ekanayaka AH, Wen TC, Deng CY, Lateef AA, Pereira OL, Navathe S, Hawksworth DL, Fan XL, Dissanayake LS, Erdoğdu M (2020) Outline of Fungi and fungi-like taxa. Mycosphere 11(1): 1060–1456.
  • Yang CL, Baral HO, Xu XL, Liu YG (2019a) Parakarstenia phyllostachydis, a new genus and species of non-lichenized Odontotremataceae (Ostropales, Ascomycota). Mycological Progress 18(6): 833–845.
  • Yang CL, Xu XL, Dong W, Wanasinghe DN, Liu YG, Hyde KD (2019b) Introducing Arthrinium phyllostachium sp. nov. (Apiosporaceae, Xylariales) on Phyllostachys heteroclada from Sichuan Province, China. Phytotaxa 406(2): 91–110.
  • Yang CL, Xu XL, Liu YG, Hyde KD, McKenzie EHC (2019d) A new species of Phyllachora (Phyllachoraceae, Phyllachorales) on Phyllostachys heteroclada from Sichuan, China. Phytotaxa 392(3): 186–196.
  • Yang CL, Xu XL, Wanasinghe DN, Jeewon R, Phookamsak R, Liu YG, Liu LJ, Hyde KD (2019f) Neostagonosporella sichuanensis gen. et sp. nov. (Phaeosphaeriaceae, Pleosporales) on Phyllostachys heteroclada (Poaceae) from Sichuan Province, China. MycoKeys 46: 119–150.
  • Yi TP (1997) Bamboos Flora of Sichuan. China Forestry Publishing House, 358 pp.
  • Yi TP, Shi JY, Ma LS, Wang HT, Yang L (2008) Iconographia Bambusoidearum Sinicarum. Science Press, 766 pp.
  • Zhou DQ, Hyde KD, Wu XL (2001) New records of Ellisembia, Penzigomyces, Sporidesmium and Repetophragma species on Bamboo from China. Acta Botanica Yunnanica 23(1): 45–51.

Supplementary materials

Supplementary material 1 

Figure S1

Xiu-Lan Xu, Chun-Lin Yang, Rajesh Jeewon, Dhanushka N. Wanasinghe, Ying-Gao Liu, Qian-Gang Xiao

Data type: multimedia

This dataset is made available under the Open Database License ( 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.
Download file (155.43 kb)
Supplementary material 2 

Figure S2

Xiu-Lan Xu, Chun-Lin Yang, Rajesh Jeewon, Dhanushka N. Wanasinghe, Ying-Gao Liu, Qian-Gang Xiao

Data type: multimedia

This dataset is made available under the Open Database License ( 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.
Download file (182.88 kb)