Research Article
Research Article
Two novel species and three new records of Torulaceae from Yunnan Province, China
expand article infoWen-Peng Wang, Hong-Wei Shen§, Dan-Feng Bao§, Yong-Zhong Lu|, Qiu-Xia Yang, Xi-Jun Su, Zong-Long Luo
‡ Dali University, Dali, China
§ Mae Fah Luang University, Chiang Rai, Thailand
| School of Food and Pharmaceutical Engineering, Guizhou Institute of Technology, Guiyang, China
Open Access


While investigating the diversity of lignicolous fungi in Yunnan Province, China, six fresh collections of Torulaceae were collected and identified based on morphological examination and phylogenetic analyses of combined LSU, ITS, SSU, tef1-α, and rpb2 sequence data. Two new species, viz. Neopodoconis yunnanensis and Torula suae, and three new records, viz. T. canangae (new freshwater habitat record), T. masonii (new host record), and T. sundara (new freshwater habitat record) are reported. Detailed descriptions, illustrations, and a phylogenetic tree to show the placement of these species are provided.

Key words

2 new species, lignicolous fungi, morphology, multigene phylogeny, Pleosporales


Torulaceae (Pleosporales) was introduced by Sturm (1829) to accommodate the type genus Torula, which is only known by its asexual morph. The family is characterized by septate, subcylindrical conidiophores with or without apical branches, doliiform to ellipsoid or clavate, smooth to verruculose, mono- to polyblastic conidiogenous cells, and subcylindrical or fusiform, smooth to verrucose conidia which form branched chains (Crous et al. 2014, 2015; Hyde et al. 2016; Li et al. 2016; Su et al. 2016, 2018). Crous et al. (2015) revised the classification of Torulaceae and accepted two genera, viz. Dendryphion and Torula in the family. Afterward, two freshwater genera, Neotorula and Rostriconidium, and a terrestrial genus, Sporidesmioides, were introduced to Torulaceae (Li et al. 2016; Su et al. 2016, 2018). Recently, two additional torula-like genera, Cylindrotorula and Rutola, have been added to the family (Crous et al. 2020; Boonmee et al. 2021). Qiu et al. (2022) combined Rostriconidium and Sporidesmioides into the Neopodoconis based on morphology and phylogeny. Currently, Torulaceae comprises six genera, viz. Cylindrotorula, Dendryphion, Neopodoconis, Neotorula, Rutola and Torula. Members of Torulaceae distributed worldwide, and most taxa are saprobes on dead or decaying wood in freshwater and terrestrial habitats (Crous et al. 2015; Su et al. 2016, 2018; Li et al. 2017; Pratibha and Prabhugaonkar 2017; Hyde et al. 2020; Shen et al. 2021; Boonmee et al. 2021).

Neopodoconis was introduced by Rifai (2008) to accommodate N. ampullacea (type species) and N. megasperma which were previously placed in Exosporium Link. The genus is characterized by macronematous, mononematous, unbranched, smooth-walled, septate conidiophores, integrated, elongated sympodially conidiogenous cells that are terminal and monotretic or polytretic, and acropleurogenous, obclavate, fusiform to pyriform, smooth walled or verrucous conidia with euseptate and a truncate dark scar at the base (Rifai 2008; Li et al. 2016; Su et al. 2018; Tibpromma et al. 2018; Shen et al. 2021; Qiu et al. 2022). Based on the morphological characteristics and phylogenetic results, Qiu et al. (2022) regarded Sporidesmioides and Rostriconidium as synonyms of Neopodoconis and accordingly transferred R. aquaticum, R. cangshanense, R. pandanicola and S. thailandica to Neopodoconis, as well as introducing five new species. Currently, eleven species are accepted in Neopodoconis, of which only N. aquaticum, N. cangshanense and N. pandanicola have been reported in freshwater habitats (Su et al. 2018; Shen et al. 2021; Qiu et al. 2022).

Torula is typified by T. herbarum (Pers.). It is characterized by terminal or lateral, monoblastic or polyblastic conidiogenous cells produced in branched chains, and subglobose, verrucose, septate, conidia (Crous et al. 2015; Crane and Miller 2016; Hyde et al. 2017, 2019; Su et al. 2018). Members of Torula are widely distributed in different habitats around the world. Index Fungorum (2023) lists more than 500 epithets, while Species Fungorum (2023) lists more than 200 records of Torula. However, most records have been transferred to other genera; thus, currently, only 63 Torula species are accepted in Species Fungorum. Morphological differences among Torula species are not significant, hence species identification in recent years has relied mainly on molecular sequence data. Most Torula species have been reported from terrestrial habitats, with only nine species reported from freshwater habitats, and four species viz. T. fici, T. gaodangensis, T. mackenziei and T. masonii have been found in both terrestrial and freshwater habitats (Su et al. 2018; Hyde et al. 2020; Boonmee et al. 2021; Tian et al. 2023).

During our investigation of lignicolous fungi in Yunnan Province, China, six fresh collections were isolated from decaying wood. Based on morphological characteristics and phylogenetic analyses of combined LSU, ITS, SSU, tef1-α, and rpb2, two new species viz. Neopodoconis yunnanensis and Torula suae, two new freshwater habitat records viz. T. canangae and T. sundara, and a new host record of T. masonii are reported.

Materials and methods

Isolation and morphology

Specimens of decaying wood were collected from lotic habitats and riverbanks in Dali City and Wenshan City, Yunnan Province, China, and returned to the laboratory in plastic bags. Methods of morphological observation and isolation followed Luo et al. (2018) and Senanayake et al. (2020). Macromorphological characteristics of samples were observed using Optec SZ 760 compound stereomicroscope. Temporarily prepared microscope slides were placed under a Nikon ECLIPSE Ni-U compound stereomicroscope for observation and micro-morphological-photography. The morphology of colonies on native substrates was photographed with a Nikon SMZ1000 stereo zoom microscope. Single spore isolation was performed according to the following steps: the conidia suspension from specimens, absorbed with a sterilized pipette, was placed on potato dextrose agar (PDA) and incubated at room temperature overnight. Germinated conidia were transferred to new PDA (Beijing land bridge technology CO., LTD., China) plates and incubated in an incubator at room temperature (25 °C). The specimens were deposited in the Herbarium of Cryptogams Kunming Institute of Botany, Academia Sinica (KUN-HKAS), Kunming, China. Living cultures are deposited in the China General Microbiological Culture Collection Center (CGMCC), Beijing, China, and Kunming Institute of Botany Culture Collection Center, Kunming, China (KUNCC). The MycoBank numbers were registered in MycoBank database ( New species were established following the recommendations outlined by Jeewon and Hyde (2016) and Chethana et al. (2021).

DNA extraction, PCR amplification, and sequencing

DNA extraction, PCR amplification, sequencing and phylogenetic analyses followed Dissanayake et al. (2020) with the following modifications. Fungal mycelia (200–500 mg) were scraped from grown on PDA or malt extract agar (MEA) plates using a sterile scalpel, transferred to microcentrifuge tubes with sterilized needles, and then ground with liquid nitrogen or quartz sand to break the cells. DNA was extracted using the TreliefTM Plant Genomic DNA Kit (TSP101) according to the manufacturer’s instructions.

Five gene regions, LSU, ITS, SSU, tef1-α, and rpb2 were amplified using LR0R/LR5, ITS5/ITS4, NS1/NS4, 983F/EF1-2218R, and RPB2-5F/RPB2-7cR (Vilgalys and Hester 1990; White et al. 1990; Liu et al. 1999) primer pairs respectively. Primer sequences are available in the WASABI database on the AFTOL website ( The PCR mixture contained 12.5 μL of 2× Power Taq PCR Master Mix (a premix and ready to use solution, including 0.1 Units/μL Taq DNA Polymerase, 500 μm dNTP Mixture each (dATP, dCTP, dGTP, dTTP), 20 mm Tris-HCl pH 8.3, 100 Mm KCl, 3 mM MgCl2, stabilizer and enhancer), 1 μL of each primer including forwarding primer and reverse primer (10μM), 1 μL template DNA extract and 9.5 μL deionized water (Luo et al. 2018). The PCR cycling conditions of LSU, ITS, SSU and tef1-α were as follows: 94 °C for 3 mins, followed by 35 cycles of denaturation at 94 °C for 30s, annealing at 55 °C for 50s, elongation at 72 °C for 1 min, and a final extension at 72 °C for 10 mins. The PCR thermal cycle of rpb2 has a total of 40 cycles, and the conditions are as follows: initially denature at 95 °C for 5 mins, and then enter 40 cycles: denaturation at 95 °C for 1 min, annealing at 52 °C for 2 mins, extension at 72 °C for 90s, and finally at 72 °C for 10 mins. PCR products were then purified using minicolumns, purification resin, and buffer according to the manufacturer’s protocols (Amersham product code: 27–9602–01). The sequences were carried out at Beijing Tsingke Biotechnology Co., Ltd. (Beijing, P.R. China).

Phylogenetic analyses

Preliminary identification of genes obtained from fresh strains by GenBank database. The LSU, ITS, SSU, tef1-α and rpb2 used for phylogenetic analysis are selected based on the preliminary identification results and the related publications (Li et al. 2020; Phukhamsakda et al. 2020; Tian et al. 2023). The sequences were aligned using MAFFT online service: Multiple alignment program for amino acid or nucleotide sequences MAFFT version 7 (Katoh and Standley 2013:, and edited manually in BioEdit v. 7.0 (Hall 1999). The sequence dataset was combined using SquenceMatrix v.1.7.8 (Vaidya et al. 2011). The alignment formats were changed to PHYLIP and NEXUS formats by AliView and ALigment Transformation EnviRonment (ALTER) website (

Maximum likelihood (ML) analysis was using the RAxML-HPC2 on XSEDE (8.2.12) (Stamatakis 2006; Stamatakis et al. 2008) of CIPRES Science Gateway website (Miller et al. 2010: and the estimated proportion of invariant sites are (GTRGAMMA+I) model.

Bayesian analyses were performed in MrBayes 3.2.6 (Ronquist et al. 2012) and the best-fit model (LSU, ITS, SSU, tef1-α, and rpb2 are all GTR+I+G) of sequences evolution was estimated via MrModeltest 2.2 (Guindon and Gascuel 2003; Nylander 2004; Darriba et al. 2012). The Markov Chain Monte Carlo (MCMC) sampling approach was used to calculate posterior probabilities (PP) (Rannala and Yang 1996). Bayesian analyses of six simultaneous Markov chains were run for 100,000,00 generations with trees sampled every 1000 generations. Phylogenetic trees were visualized using FigTree v. 1.4.0 (Rambaut 2012:, edited in Microsoft Office PowerPoint. The new sequences were submitted in GenBank and the strain information used in this paper is provided in Table 1.

Table 1.

Names, culture accession numbers, and corresponding GenBank accession numbers used for the phylogenetic analyses.

Species Source GenBank accession number Reference
LSU ITS SSU tef1-α rpb2
Arthopyrenia salicis CBS 368.94 AY538339 KF443410 AY538333 KF443404 KF443397 Ahmed et al. (2014)
Cycasicola goaensis MFLUCC 17–0754 MG829001 MG828885 NG_061287 MG829198 Devadatha et al. (2018)
C. goaensis MFLU 17–0581 NG_059057 NR_157510 Devadatha et al. (2018)
C. leucaenae MFLUCC 17–0914 MK347942 MK347726 MK347833 Jayasiri et al. (2019)
Cylindrotorula indica NFCCI 4836 MT339442 MT339444 MT321492 MT321490 Boonmee et al. (2021)
C. indica NFCCI 4837 MT339443 MT339445 MT321493 MT321491 Boonmee et al. (2021)
Dendryphion aquaticum MFLUCC 15–0257 KU500573 KU500566 KU500580 Su et al. (2016)
D. comosum CBS 208.69 MH871026 MH859293 Vu et al. (2019)
D. europaeum CPC 23231 KJ869202 KJ869145 Crous et al. (2014)
D. fluminicola DLUCC 0849 MG208140 MG208161 MG207991 MG207972 Su et al. (2018)
D. fluminicola MFLUCC 17–1689 MG208141 NR_157490 MG207992 Su et al. (2018)
D. hydei KUMCC 18–0009 MH253927 MN061343 MH253929 MH253931 Li et al. (2020)
D. nanum HKAS 84010 KU500575 KU500568 KU500582 Su et al. (2016)
MFLUCC 16–0987 MG208135 MG208156 MG207986 MG207967 Su et al. (2018)
D. submersum MFLUCC 15–0271 KU500572 KU500565 KU500579 Su et al. (2016)
Mauritiana rhizophorae BCC 28866 GU371824 GU371832 GU371817 GU371796 Poonyth et al. (2000)
M. rhizophorae BCC 28867 GU371825 GU371833 GU371818 GU371797 Poonyth et al. (2000)
Neooccultibambusa thailandensis MFLUCC 16–0274 MH260308 MH275074 MH260348 MH412780 MH412758 Tibpromma et al. (2018)
Neopodoconis aquaticum MFLUCC 16–1113 MG208143 MG208164 MG207994 MG207974 Su et al. (2018)
N. aquaticum KUMCC 15–0297 MG208144 MG208165 MG207995 MG207975 Su et al. (2018)
N. cangshanense MFLUCC 20–0147 MW010281 MW010285 MW012636 Shen et al. (2021)
N. jiangxiensis HJAUP C0947 ON693846 ON693847 Qiu et al. (2022)
N. meilingensis HJAUP C0905 ON693849 ON693843 Qiu et al. (2022)
N. obclavata HJAUP C0829 ON693848 ON693844 Qiu et al. (2022)
N. pandanicola KUMCC 17–0176 MH260318 MH275084 MH260358 MH412781 MH412759 Tibpromma et al. (2018)
N. saprophyticus HJAUP C0830 ON693851 ON705129 Qiu et al. (2022)
N. sinensis HJAUP C0909 ON693845 ON693850 Qiu et al. (2022)
N. thailandica MFLUCC 13–0840 NG_059703 MN061347 KX437759 KX437766 KX437761 Li et al. (2016)
N. thailandica KUMCC 16–0012 KX437758 MN061348 KX437760 KX437767 KX437762 Li et al. (2016)
N. yunnanensis KUNCC 22–10737 OP359410 OP359401 OP369295 OP471613 OP476726 This study
Neoroussoella bambusae MFLUCC 11–0124 KJ474839 KJ474827 KJ474848 KJ474856 Liu et al. (2014)
N. entadae MFLUCC 17–0920 NR_163325 NG_065773 MK434898 Liu et al. (2014)
N. leucaenae MFLUCC 17–0927 NG_070073 NR_165226 NG_065774 MK360066 MK434896 Liu et al. (2014)
Neotorula aquatica MFLUCC 15–0342 KU500576 KU500569 KU500583 Su et al. (2016)
N. submersa HKAS 92660 NG_059727 NR_154247 Hyde et al. (2016)
Occultibambusa bambusae MFLUCC 13–0855 KU863112 KU940123 KU940193 KU940170 Dai et al. (2016)
Pseudocoleodictyospora tectonae MFLUCC 12–0385 KU764709 NR_154338 NG_061232 KU712491 Doilom et al. (2016)
P. tectonae MFLUCC 12–0387 KU764704 KU712444 KU712462 KU712492 Doilom et al. (2016)
Roussoella hysterioides HH 26988 AB524622 AB524481 AB539102 AB539102 Liu et al. (2014)
R. pustulans KT 1709 AB524623 AB524482 AB539116 AB539103 Liu et al. (2014)
Roussoellopsis macrospor MFLUCC 12–0005 KJ474847 KJ739604 KJ739608 KJ474855 KJ474862 Liu et al. (2014)
R. tosaensis KT 1659 AB524625 AB524484 AB539117 AB539104 Liu et al. (2014)
Rutola graminis CPC 33267 MN317295 MN313814 Crous et al. (2020)
R. graminis CPC 33695 MN317296 MN313815 Crous et al. (2020)
Subglobosporium tectonae MFLUCC 12–0390 KU764702 KU712446 KU712463 KU712495 Doilom et al. (2016))
S. tectona MFLUCC 12–0393 KU764703 NR_154426 NG_061233 KU712485 Doilom et al. (2016))
Thyridariella mahakoshae NFCCI 4215 MG020438 MG020435 MG020441 MG023140 MG020446 Devadatha et al. (2018)
T. mangrovei NFCCI 4213 MG020437 MG020434 MG020440 MG020443 MG020445 Devadatha et al. (2018)
Torula acaciae CPC 29737 NG_059764 NR_155944 KY173594 Crous et al. (2016)
T. aquatica DLUCC 0550 MG208145 MG208166 MG207996 MG207976 Su et al. (2018)
T. aquatica MFLUCC 16–1115 MG208146 MG208167 MG207977 Su et al. (2018)
T. breviconidiophora KUMCC 18–0130 MK071672 MK071670 MK071697 MK077673 Hyde et al. (2019)
T. camporesii KUMCC 19–0112 MN507402 MN507400 MN507401 MN507403 MN507404 Hyde et al. (2020)
T. canangae MFLUCC 21-0169 OL830816 OL966950 Silva et al. (2022)
T. canangae KUNCC 22–12432 OP359414 OP359405 OP369299 OP471617 OP476729 This study
T. chiangmaiensis KUMCC 16–0039 KY197856 MN061342 KY197863 KY197876 Li et al. (2017)
T. chinensis UESTCC 22.0085 OQ128004 OQ127986 OQ127995 Tian et al. (2023)
T. chromolaenae KUMCC 16–0036 KY197860 MN061345 KY197867 KY197880 KY197873 Li et al. (2017)
T. fici CBS 595.96 KF443385 KF443408 KF443387 KF443402 KF443395 Crous et al. (2015)
T. fici KUMCC 16–0038 KY197859 MN061341 KY197866 KY197879 KY197872 Li et al. (2017)
T. gaodangensis MFLUCC 17–0234 NG_059827 MF034135 NG_063641 Hyde et al. (2020)
T. goaensis NFCCL 4040 NG_060016 NR_159045 Pratibha and Prabhugaonkar (2017)
T. herbarum CPC 24414 KR873288 KR873260 Crous et al. (2015)
T. hollandica CBS 220.69 NG_064274 NR_132893 KF443389 KF443401 KF443393 Crous et al. (2015)
T. hydei KUMCC 16–0037 MH253926 MN061346 MH253928 MH253930 Li et al. (2020)
T. lancangjiangensis MFLUCC 21–0098 MW879526 MW723059 MW774582 MW729785 MW729780 Boonmee et al. (2021)
T. lancangjiangensis HKAS 112709 MZ538563 MZ538529 MZ567104 Boonmee et al. (2021)
T. longiconidiophora UESTCC 22.0088 OQ128001 OQ127983 OQ127992 Tian et al. (2023)
T. longiconidiophora UESTCC 22.0125 OQ128002 OQ127984 OQ127993 Tian et al. (2023)
T. mackenziei HKAS 112705 MW879525 MW723058 MW774581 MW729784 MW729779 Boonmee et al. (2021)
T. mackenziei MFLUCC 13–0839 KY197861 MN061344 KY197868 KY197881 KY197874 Li et al. (2017)
T. masonii CBS 245.57 NG_058185 NR_145193 Crous et al. (2015)
T. masonii DLUCC 0588 MG208152 MG208173 MG208000 MG207982 Su et al. (2018)
KUMCC 16–0033 KY197857 MN061339 KY197864 KY197877 KY197870 Li et al. (2017)
UESTCC 22.0089 OQ128000 OQ127982 OQ127991 Tian et al. (2023)
KUNCC 22–12429 OP359411 OP359402 OP369296 OP471614 OP476727 This study
T. pluriseptata MFLUCC 14–0437 KY197855 MN061338 KY197862 KY197875 KY197869 Li et al. (2017)
T. polyseptata KUMCC 18–0131 MK071673 MK071671 MK071698 MK077674 Hyde et al. (2019)
T. sichuanensis UESTCC 22.0087 OQ127999 OQ127981 OQ127990 Tian et al. (2023)
T. sp. CBS 246.57 KR873290 KF443411 Crous et al. (2015)
T. suae CGMCC 3.24259 OP359415 OP359406 OP369300 OP471618 OP476730 This study
T. submersa UESTCC 22.0086 OQ128003 OQ127985 OQ127994 Tian et al. (2023)
T. sundara MFLUCC 21–0067 OM287866 OM276824 Jayawardena et al. (2022)
KUNCC 22–12430 OP359412 OP359403 OP369297 OP471615 This study
KUNCC 22–13431 OP359413 OP359404 OP369298 OP471616 OP476728 This study
T. thailandica GZCC 20-0011 MN907428 MN907426 MN907427 Silva et al. (2022)


Phylogenetic analyses

The phylogenetic analyses comprised LSU (1–829 bp), ITS (830–1322 bp), SSU (1323–2176 bp), tef1-α (2177–2996 bp) and rpb2 (2997–3981 bp) gene regions with 3981 characters, with Occultibambusa bambusae (MFLUCC 13–0855) and Neooccultibusa thailandensis (MFLUCC 16–0274) as the outgroup taxa (Fig. 1). Bayesian (PP) and Maximum likelihood (ML) analyses of the combined dataset resulted in phylogenetic reconstructions with largely similar topologies, and the best scoring RAxML tree is shown in Fig. 1. The final ML optimization likelihood value of -34180.028053. The aligned matrix had 1645 distinct alignment patterns, with 31.25% completely undetermined characters or gaps. Base frequency and rate are as follows: A = 0.246934, C = 0.258869, G = 0.272596, T = 0.221601; rate AC = 1.558046, AG = 3.589878, AT = 1.463717, CG = 1.029375, CT = 7.027540, GT = 1.000000; gamma distribution shape: α = 0.190356.

Figure 1. 

The maximum likelihood (ML) tree based on combined LSU, ITS, SSU, tef1-α, and rpb2 sequence data. Bootstrap support values with an ML greater than 75% and Bayesian posterior probabilities (PP) greater than 0.95 are given above the nodes, shown as “ML/PP”. The tree is rooted with Occultibambusa bambusae (MFLUCC 13–0855) and Neooccultibusa thailandensis (MFLUCC 16–0274). New species and collections are indicated in red; while the type strains are in bold black.

Phylogenetic analyses have revealed that our six new isolates are nested in Torulaceae. Five new strains were grouped within Torula, while one was clustered within Neopodoconis. Neopodoconis yunnanensis (KUNCC 22–10737) clustered with N. aquaticum, N. cangshanense, N. saprophyticus and N. pandanicola with 100% ML/1.00 PP support. The new isolate Torula canangae (KUNCC 22–12432) was clustered with the ex-type strain of T. canangae (MFLUCC 21–0169) with 100% ML/1.00 PP support. Torula masonii (KUNCC 22–12429) clustered with the ex-type strain of T. masonii (CBS 245.57) with 82% ML support. Torula suae (KUNCC 22–12433) was clustered sister to T. goaensis (NFCCL 4040) with a low bootstrap support. Torula sundara (KUNCC 22–12430, KUNCC 22–12431) was clustered with T. sundara (MFLUCC 21–0076) and T. longiconidiophora (UESTCC 22.0088) with 94% ML/0.99 PP statistical support.


Neopodoconis yunnanensis W.P. Wang, H.W. Shen & Z.L. Luo, sp. nov.

MycoBank No: 848412
Fig. 2


Referring to the collection location, Yunnan Province of China.


KUN-HKAS 121702.


Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph: Colonies grow on the surface of the substrate, black, hairy, and distinct branches can be seen. Mycelium immersed in the substrate, composed of pale brown, septate, unbranched hyphae. Conidiophores 174–648 × 8.2–17.5 μm ( = 311 × 12 μm, n = 30), macronematous, mononematous, concentrated, erect, dark brown to black, smooth-walled, septate, unbranched, straight or slightly flexuous, pale pigment at apex. Conidiogenous cells 18–34 × 9–14 μm ( = 27 × 12 μm, n = 30), monoblastic or polyblastic, integrated, terminal, cylindrical, smooth, dark brown to black. Conidia 100–155 × 23–38 μm ( = 128 × 28 μm, n = 20), solitary, smooth, dry, pyriform to fusiform, dark brown to black, light brown at the apex, granular inclusions, rostrate, guttulate, 5–7-septate, dark bands at the septa, slightly cicatrized at narrow, black truncate scar at base and pale pigment cell above the scar, wide in the middle.

Figure 2. 

Neopodoconis yunnanensis (KUN-HKAS 121702, holotype) A, B fungal structures on the substratum C conidiophore with conidia on the stroma D conidiophore with conidia E–G conidiogenous cells H–M conidia N germinating conidium. Scale bars: 150 μm (C, D), 20 μm (E–G), 50 μm (H–N).

Material examined

China, Yunnan Province, Dali City, Cangshan Mountain, Mocanxi Stream (25°64′82.95″N, 100°15′80.33″E), on submerged decaying wood, 11 April 2020, Zheng-Quan Zhang, S2690 (KUN-HKAS 121702, holotype), ex-type culture (KUNCC 22–10737).


Neopodoconis yunnanensis fits well with the generic concept of Neopodoconis in having erect, septate conidiophores, terminal, cylindrical conidiogenous cells, and rostrate, septate conidia with a subhyaline base. Phylogenetic analyses showed that Neopodoconis yunnanensis constitutes a strongly supported (100% ML/1.00 PP) independent lineage that is basal to four Neopodoconis species viz. N. aquaticum, N. cangshanense, N. saprophyticus and N. pandanicola. Neopodoconis yunnanensis (KUN-HKAS 121702) differs from N. aquaticum in having shorter but broader conidia (100–155 × 23–38 vs. 134–180 × 22–26 μm). Additionally, it has smooth conidiophores, lacks constriction at septum, larger conidia compared to N. pandanicola (100–155 × 23–38 μm vs. 55–110 × 18–26 μm), different from N. cangshanense in terms of larger size (100–155 × 23–38 vs. 94–109 × 11–24 μm), and lacks hyaline sheath in the apex of conidia. We, therefore, describe the newly obtained taxon as a new species based on both morphology and multigene phylogeny (Su et al. 2018; Tibpromma et al. 2018; Shen et al. 2021).

Torula canangae N.I. de Silva,S, Lumyong & K.D. Hyde. Mycosphere 13(1): 955–1076 (2022)

MycoBank No: 559523
Fig. 3


Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse on the natural substrate, neat, hairy, brown. Mycelium immersed to superficial, hyaline, septate, branched hyphae. Conidiophores indistinct. Conidiogenous cells 6–10(–13) × 5–7 (–13) μm ( = 8 × 7 μm, n = 15), holoblastic, mono-to polyblastic, integrated, terminal, doliiform to spherical, brown to dark brown. Conidia (28–) 78–113 (–142) × 6–9 μm ( = 82 × 7 μm, n = 20), acrogenous, dry, brown to dark brown, subhyaline at terminal cell, constricted at septa, verrucose, easily separating, 5–29-septate, cell size is uniform, chiefly subcylindrical.

Figure 3. 

Torula canangae (KUNCC 22-12432) A fungal structure on the substratum B–D conidiogenous cells E conidiogenous cells with conidia F–K conidia L germinating conidium M, N colonies on PDA from surface and reverse. Scale bars: 10 μm (B–D), 15 μm (E), 30 μm (F–H), 50 μm (I–K), 100 μm (L).

Culture characters

Conidia germinating on PDA within 12 h, and germ tubes produced at the side. Mycelium superficial, branched, septate, hyaline, smooth. After two weeks of incubation at room temperature, colony appears distinctly rounded, the central hyphae are longer, white, velvety, and the edges are white to brown and the hyphae are shorter.

Material examined

China, Yunnan Province, Wenshan, Bamei Town (24°31′96.49″N, 105°03′84.35″E), on submerged decaying wood, 7 February 2022, Wen-Peng Wang S3492 (KUN-HKAS 124619), living culture, KUNCC 22–12432 = CGMCC 3.24258.


Silva et al. (2022) first introduced Torula canangae, which was collected from terrestrial habitats on dead twigs of Cananga odorata in Thailand. In this study, phylogenetic analyses showed that our collection clustered with the ex-type strain of T. canangae (MFLUCC 21–0169) with 100% ML/1.00 PP support (Fig. 1). Our collection has similar morphological features to T. canangae, such as indistinct conidiophores, ellipsoid to coronal, terminal conidiogenous cells, and mainly subcylindrical conidia (Silva et al. 2022). Thus, we identify our isolate as T. canangae based on both morphology and multigene phylogeny, and it is a new record for freshwater habitat in China.

Torula masonii Crous. IMA Fungus 6(1): 195 (2015)

MycoBank No: 812806
Fig. 4


Saprobic on dead Artemisia carvifolia stems. Sexual morph Undetermined. Asexual morph: Colonies effuse on the natural substrate, scattered, hairy, dark brown to black. Mycelium mostly immersed. Conidiophores 16–28 (–45) × 3–4 μm ( = 26 × 4 μm, n = 10), macronematous mononematous, subcylindrical, erect, septate, smooth, straight or slightly flexuous, brown to dark brown, the uppermost side of a transverse compartment is concave inward. Conidiogenous cells 8–10 × 5–7 μm ( = 9 × 6 μm, n =15), holoblastic, polyblastic, doliiform to ellipsoid, dark brown, smooth. Conidia (16–) 25–48 (–70) × 6–10 μm ( = 35 × 8 μm, n = 50), phragmosporous, in branched chains, acrogenous, dry, brown to dark brown, subhyaline at terminal cell, and central cells are significantly larger than both end cells, nearly ellipsoid, constricted at septa, verrucose, easily separating, 2–16-septate, cells subglobose.

Figure 4. 

Torula masonii (KUNCC 22-12429) A fungal structure on the substratum C–F conidiophore with conidiogenous cells B, G–O conidia P germinating conidium Q, R colonies on PDA from surface and reverse. Scale bars: 20 μm (B), 10 μm (C–N), 25 μm (O, P)

Culture characteristics

Conidia germinating on PDA within 12 h. and germ tubes produced at the side. Mycelium superficial, branched, septate, hyaline, smooth. After two weeks of incubation at room temperature, colony edges are irregularly ellipsoid, center is white with gray fuzzy protrusions on the sides, and a translucent gelatinous substance at the outermost periphery. Hyphae flocculent, velvety.

Material examined

China, Yunnan Province, Dali, Cangshan mountain (25°64′82.95″N, 100°15′80.33″E), on dead Artemisia carvifolia stems, 16 October 2021, Wen-Peng Wang H630 (KUN-HKAS 124616), living culture, KUNCC 22–12429 = CGMCC 3.23734.


Torula masonii collected on Brassica sp. in the UK was introduced by Crous et al. (2015). Since then, it was reported from freshwater and terrestrial habitats in China and Italy (Li et al. 2017; Su et al. 2018; Tian et al. 2023). In this study, our new collection was obtained from dead stems of Artemisia carvifolia (Asteraceae) in Yunnan, China. Phylogenetic analyses showed that our new isolate clustered with the ex-type strain of T. masonii (CBS 245.57) with good bootstrap support (82% ML, Fig. 1). Morphologically, our new isolate is similar to T. masonii in having macronematous, mononematous, subcylindrical conidiophores, polyblastic conidiogenous cells and dry, constricted at septa, verrucose, easily separating conidia that are formed branched chains. Thus, we identify this isolate as T. masonii, which was first reported on Artemisia carvifolia (Asteraceae).

Torula suae W.P. Wang, H.W. Shen & Z.L. Luo, sp. nov.

MycoBank No: 848410
Fig. 5


“suae” (Lat) in memory of the Chinese mycologist Prof. Hong-Yan Su, who kindly helped the authors in many ways and sadly passed away on 3 May 2022.


KUN-HKAS 124620.


Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse on the natural substrate, neat, hairy, brown to dark brown. Mycelium immersed to superficial, composed of hyaline, becoming brown closer to fertile region, septate, branched hyphae. Conidiophores 17–54 × 3–4 μm ( = 32 × 3 μm, n = 10), macronematous to semi- macronematous, erect, straight, or slightly flexuous, without apical branches, light brown to brown, ellipsoid to subcylindrical, smooth, septate. Conidiogenous cells 6–8 × 5–7 μm ( = 7 × 6 μm, n = 20), mono- to polyblastic, integrated, terminal or intercalary, doliiform to subglobose, brown to dark brown. Conidia (16–) 31–115 (–160) × 6–9 μm ( = 69 × 7 μm, n = 35), in branched chains, acrogenous, phragmoconidia, golden at apex, brown to dark brown, 2–29-septate, constricted at the septa, verrucose, easily separating, guttulate, chiefly subcylindrical, globlose to subglobose of each cell.

Figure 5. 

Torula suae (KUN-HKAS 124620, holotype) A fungal structure on the substratum C, D Conidiophores with conidiogenous cells B, E–I conidia J germinating conidium K, L colonies on PDA from surface and reverse. Scale bars: 60 μm (B), 10 μm (C–F), 20 μm (G, H), 30 μm (I, J).

Culture characteristics

Conidia germinating on PDA within 12 h, and germ tubes produced at the side. Mycelium superficial, branched, septate, hyaline, smooth. After two weeks of incubation at room temperature, colony appears distinctly rounded, the central hyphae are longer, white, velvety, and the edges are brown and the hyphae are shorter.

Material examined

China, Yunnan Province, Wenshan City, Bamei Town (24°31′96.49″N, 105°03′84.35″E), on submerged decaying wood, 7 February 2022, Wen-Peng Wang S–3509 (KUN-HKAS 124620, holotype), ex-type living culture, KUNCC 22–12433 = CGMCC 3.24259.


According to the BLASTn results, the closest matches for our new species were Torula suae (KUNCC 22–12433) (NR 159045, 98.59% similarity in ITS) and T. goaensis (NFCCL 4040) (NG 060016, 99.60% similarity in LSU). Comparison of ITS and LSU nucleotide bases indicated that T. suae differs from T. goaensis in 7/494 (ITS) and 5/1257 (LSU). Phylogenetic analyses showed that T. suae clustered with T. goaensis with low support. Morphologically, T. suae is similar to T. goaensis in having conidiophores without apical branches and doliiform to subglobose conidiogenous cells (Pratibha and Prabhugaonkar 2017). However, T. suae can be distinguished from T. goaensis by the conidial type; T. goaensis has phragmoconidia and scolecoconidia, whereas T. suae only produces phragmoconidia, and T. suae has more conidial septa (2–29 vs. 7–20).

Torula sundara (Subram.) Y.R. Sun, Yong Wang & K.D. Hyde. Fungal Diversity 117:1–272. (2022)

MycoBank No: 559464
Fig. 6


Saprobic on submerged decaying wood. Sexual morph: Undeter­mined. Asexual morph: Colonies on the natural substrate, effuse, scattered, hairy, yellow to black, dry. Mycelium mostly immersed, hyaline, septate, branched hyphae. Conidiophores 20–53 × 3–4 μm ( = 36 × 4 μm, n = 15), micronematous to semi-macronematous, mononematous, subcylindrical, erect, septate, smooth, straight, or slightly flexuous, brown to dark brown, branched. Conidiogenous cells 6–9 × 5–7 μm ( = 7 × 6 μm, n = 20), holoblastic, mono-to polyblastic, integrated, terminal, doliiform to ellipsoid, brown to dark brown. Conidia two types, short conidia and long conidia. Short conidia 18–58 × 5–11 μm ( = 42 × 9 μm, n =30), acrogenous, phragmosporous, in branched chains, dry, brown to dark brown, subhyaline at terminal cell, constricted at septa, verrucose, subglobose cells and central cells are larger than the ends cells, 3–15-septate. Long conidia 165–368 × 4–7 μm ( =226 × 6 μm, n =10), acrogenous, phragmosporous, dry, straight to slightly flexuous, light brown to brown, subhyaline at terminal cell, constricted at septa, verrucose, easily separating, fusiform to ellipsoidal cells and uniform in size, 20–30-septate.

Figure 6. 

Torula sundara (KUNCC 22-12431) A fungal structure on the substratum B, C mycelium, conidiophore and conidia D–F conidiophore, conidiogenous cells with conidia G–I short conidia J–L long conidia M germinating conidium N, O colonies on PDA from surface and reverse. Scale bars: 40 μm (B, G), 15 μm (C), 20 μm (D-F, H, I, M), 50 μm (J–L).

Culture characteristics

Conidia germinating on PDA within 12 h, and germ tubes produced at the side. Mycelium superficial, branched, septate, hyaline, smooth. After two weeks of incubation at room temperature, colony appears distinctly rounded; there is a spherical protrusion in the center with a circle of brown stripes around it. Hyphae flocculent, velvety.

Material examined

China, Yunnan Province, Wenshan, Bamei Town (24°31′96.49″N, 105°03′84.35″E), on submerged decaying wood, 7 February 2022, Wen-Peng Wang, S3256 (KUN-HKAS 124617), living culture, KUNCC 22–12430 = CGMCC 3.23735; submerged decaying wood, 7 February 2022, Wen-Peng Wang, S3269 (KUN-HKAS 124618), living culture, KUNCC 22–12431.


Torula sundara collected from terrestrial habitats on bamboo culms in Chiang Mai Province, Thailand was introduced by Jayawardena et al. (2022). In this study, phylogenetic analyses showed that our two new strains clustered with the strain of T. sundara (MFLUCC 21–0067) with 94% ML/0.99 PP support (Fig. 1). The most obvious feature of T. sundara is that there are two types of conidia, and long conidia are more than 100 μm long (Jayawardena et al. 2022), and our collections fit well with the description of T. sundara (MFLUCC 21–0067). Therefore, we identify our isolate as T. sundara, which was collected from a freshwater habitat for the first time.


Qiu et al. (2022) compared the morphological features of Rostriconidium and Sporidesmioides with the closely related genus Neopodoconis, and proposed Sporidesmioides and Rostriconidium as synonyms of Neopodoconis. In our phylogenetic analyses, Neopodoconis was grouped into two distinct clades. Clade 1 consists of N. aquatica (R. aquaticum), N. cangshanense (R. cangshanense), N. pandanicola (R. pandanicola), N. saprophyticus and N. yunnanensis, forming a monophyletic clade basal to Torulaceae; clade 2 comprises N. jiangxiensis, N. meilingensis, N. obclavate, N. sinensis and N. thailandica (Sporidesmioides thailandica), and this clade is located outside Torulaceae. The morphology of these two clades is quite similar and cannot be distinguished easily based on morphological characteristics, but phylogenetically, they are grouped into two different clades within different families. Both clades are only known by the asexual morphs, and the sexual morphs have not been found yet. Hence, further fresh collections are required to better understand the morphology of these two clades.

An interesting finding of our study is that the phylogenetic analyses yielded similar topologies to Qiu et al. (2022), except for N. meilingensis. In our phylogenetic results based on the combination of five gene loci, N. meilingensis clustered with the species in clade 2, while the phylogenetic results based on SSU and LSU by Qiu et al. (2022) showed that it was closely related to clade 1. In addition, the five new species of Qiu et al. (2022) did not provide more gene sequences, and some gene fragments were too short (about 300 bp), which is insufficient to support a natural taxonomic status of taxa (e.g., N. meilingensis). Therefore, we call on taxonomists to provide enough genetic loci for newly introduced species to facilitate subsequent more comprehensive phylogenetic studies.

Furthermore, the new genus Pseudohelminthosporium (Neomassarinaceae, Pleosporales), was recently proposed with the type species P. clematidis which has fusiform or obclavate>, rostrate, euseptate, verrucose, with a thick, black and protruding scars at the base of the conidia. These characteristics fit well with the description of Exosporium ampullaceum (Ellis 1961; Phukhamsakda et al. 2020), therefore, Koukol et al. (2021) proposed Pseudohelminthosporium as a synonym of Neopodoconis. Currently, Neopodoconis is a polyphyletic genus, and three clades are located in different families within Pleosporales.


Zheng-Quan Zhang, Long-Li Li, Sha Luan, and Liang Zhang are thanked for their help with sample collection, DNA extraction, and PCR amplification.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


We thank the National Natural Science Foundation of China (Project ID: 32060005), and the Yunnan Fundamental Research Project (202201AW070001) for financial support.

Author contributions

Data curation: DFB. Formal analysis: HWS. Funding acquisition: ZLL. Investigation: QXY, WPWW. Project administration: ZLL, XJS. Resources: QXY. Software: HWS. Supervision: XJS. Writing - original draft: WPWW. Writing - review and editing: YZL, DFB.

Author ORCIDs

Hong-Wei Shen

Yong-Zhong Lu

Zong-Long Luo

Data availability

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


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