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New contributions to Diatrypaceae from karst areas in China
expand article infoSihan Long, Lili Liu, Yinhui Pi, Youpeng Wu, Yan Lin, Xu Zhang, Qingde Long, Yingqian Kang, Jichuan Kang§, Nalin N. Wijayawardene|, Feng Wang, Xiangchun Shen, Qirui Li
‡ Guizhou Medical University, Guiyang, China
§ Guizhou University, Guiyang, China
| Qujing Normal University, Guiyang, China
¶ Guizhou Provincial Academician Workstation of Microbiology and Health, Guizhou Academy of Tobacco Science, Guiyang, China
Open Access

Abstract

In this study, fungal specimens of the family Diatrypaceae were collected from karst areas in Guizhou, Hainan and Yunnan Provinces, China. Morpho-molecular analyses confirmed that these new collections comprise a new genus Pseudodiatrype, three new species (Diatrype lancangensis, Diatrypella pseudooregonensis and Eutypa cerasi), a new combination (Diatrypella oregonensis), two new records (Allodiatrype thailandica and Diatrypella vulgaris) from China and two other known species (Neoeutypella baoshanensis and Paraeutypella citricola). The new taxa are introduced, based on multi-gene phylogenetic analyses (ITS, β-tubulin), as well as morphological analyses. The new genus Pseudodiatrype is characterised by its wart-like stromata with 5–20 ascomata immersed in one stroma and the endostroma composed of thin black outer and inner layers of large white cells with thin, powdery, yellowish cells. These characteristics separate this genus from two similar genera Allodiatrype and Diatrype. Based on morphological as well as phylogenetic analyses, Diatrype lancangensis is introduced as a new species of Diatrype. The stromata of Diatrype lancangensis are similar to those of D. subundulata and D. undulate, but the ascospores are larger. Based on phylogenetic analyses, Diatrype oregonensis is transferred to the genus Diatrypella as Diatrypella oregonensis while Diatrypella pseudooregonensis is introduced as a new species of Diatrypella with 8 spores in an ascus. In addition, multi-gene phylogenetic analyses show that Eutypa cerasi is closely related to E. lata, but the ascomata and asci of Eutypa cerasi are smaller. The polyphyletic nature of some genera of Diatrypaceae has led to confusion in the classification of the family, thus we discuss whether the number of ascospores per asci can still be used as a basis for classification.

Keywords

Five novel taxa, phylogeny, systematics, taxonomy, Xylariales

Introduction

Diatrypaceae is an important family of higher ascomycetes, belonging to Xylariales (Maharachchikumbura et al. 2016). In the latest compilation, Hyde et al. (2020a) revised the family Diatrypaceae and included several new genera (i.e. Allodiatrype Konta & K.D. Hyde, Halocryptovalsa Dayar. & K.D. Hyde and Neoeutypella M. Raza et al.). This was followed by Wijayawardene et al. (2020) in which 20 genera were accepted into Diatrypaceae. The Diatrypaceae is characterised by perithecial ascomata embedded in a poor or well-developed, brown or black-coloured stroma, long-stalked and 8-spored or numerous-spored asci and allantoid, unicellular ascospores (Glawe and Rogers 1984; Rappaz 1987; Mehrabi et al. 2015; de Almeida et al. 2016).

Members of Diatrypaceae occur on a wide range of hosts in terrestrial and marine environments worldwide, some of which are important plant pathogens (Moyo et al. 2018a; Mehrabi et al. 2019; Dayarathne et al. 2020; Konta et al. 2020). For many decades, canker diseases on grapevine have been attributed to the species of Diatrypaceae worldwide, for example in China Cryptovalsa Ces. & De Not., Cryptosphaeria Ces. & De Not, Diatrype Fr., Diatrypella (Ces. & De Not.) De Not., Eutypa Tul. & C. Tul. And Eutypella (Nitschke) Sacc., are responsible for canker diseases in grapevine (Trouillas et al. 2011; Gao et al. 2013; Moyo et al. 2018b). Besides cankers of grapevine, some species have been reported as the causal pathogentic agents of fruit trees and woody plants in Europe and the USA (Trouillas et al. 2011; Gao et al. 2013).

Thirteen species of Cryptosphaeria and Diatrype were introduced by Vasiljeva and Ma (2014) from north-eastern China, which includes two new species and four new records. China has the largest range of karst distribution in the world. The landform of karst can be found in almost all Provinces of China, with the most extensive distribution in Guizhou and Yunnan Provinces (Miao et al. 2007). Karst virgin forest is a relatively stable ecosystem with rich biological resources, highly primitive and maintaining stable biological diversity (Dong et al. 2002). The special karst and ecological environment is home to a rich diversity of diatrypaceous fungi.

In this study, we revisit species of Diatrypaceae collected from karst areas in Guizhou, Hainan and Yunnan Provinces of China. Based on morpho-molecular analyses, one new genus and three new species are introduced; in addition, a new combination and two new records from China are reported. Descriptions and illustrations of new taxa and new records are provided.

Materials and Methods

Fungi collection, isolation and identification

Samples of decaying wood were collected from October 2019 to November 2020 in forests and nature reserves of Guizhou, Hainan and Yunnan Provinces in China. The specimens were observed with a stereomicroscope while microscopic images of the samples were taken using a Nikon ECLIPSE Ni compound microscope, with a Canon EOS 700D digital camera. Measurements were taken with Tarosoft (R) Image Frame Work (v.0.9.7). More than 30 asci and ascospores were measured for each specimen examined. Photoplates were arranged and improved by using Adobe Photoshop CS6 software. Isolations of fungi were made by single spore isolation (Chomnunti et al. 2014) and germinated spores were transferred to potato dextrose agar (PDA) medium for purification. The specimens were deposited at the Herbarium of Cryptogams, Kunming Institute of Botany Academia Sinica (KUN-HKAS) and Herbarium of Guizhou Medical University (GMB). Strains of the new genus and new species are maintained in the Guizhou Medical University Collection Centre (GMBC).

Table 1.

Taxa used in the phylogenetic analyses and their corresponding GenBank accession numbers.

Taxa Strain number GenBank Accession number Reference
ITS β-tubulin
Allocryptovalsa elaeidis MFLUCC 15-0707 MN308410 MN340296 Konta et al. (2020)
A. polysporaT MFLUCC 17-0364 MF959500 MG334556 Senwanna et al. (2017)
A. rabenhorstii WA08CB HQ692619 HQ692523 Trouillas et al. (2011)
Allodiatrype arengaeT MFLUCC 15-0713 MN308411 MN340297 Konta et al. (2020)
A. elaeidicola MFLUCC 15-0737a MN308415 MN340299 Konta et al. (2020)
A. elaeidis MFLUCC 15-0708a MN308412 MN340298 Konta et al. (2020)
A. thailandica MFLUCC 15-3662 KU315392 NA Li et al. (2016)
A. thailandica MFLUCC 15-0711 MN308414 NA Konta et al. (2020)
A. thailandica GMB0050 MW797108 MW814880 This study
Anthostoma decipiensT IPV-FW349 AM399021 AM920693 Unpublished.
A. decipiensT JL567 JN975370 JN975407 Luque et al. (2012)
Cryptosphaeria ligniota CBS 273.87 KT425233 KT425168 Acero et al. (2004)
C. pullmanensis ATCC 52655 KT425235 KT425170 Trouillas et al. (2015)
C. subcutanea DSUB100A KT425189 KT425124 Trouillas et al. (2015)
C. subcutanea CBS 240.87 KT425232 KT425167 Trouillas et al. (2015)
Cryptovalsa ampelina A001 GQ293901 GQ293972 Trouillas et al. (2010)
C. ampelina DRO101 GQ293902 GQ293982 Trouillas et al. (2010)
Diatrype bullata UCDDCh400 DQ006946 DQ007002 Rolshausen et al. (2006)
D. disciformisT GNA14 KR605644.1 KY352434.1 Senanayake et al. (2015)
D. disciformisT D21C, CBS 205.87 AJ302437 NA Acero et al. (2004)
D. enteroxantha HUEFS155114 KM396617 KT003700 de Almeida et al. (2016)
D. enteroxantha HUEFS155116 KM396618 KT022236 de Almeida et al. (2016)
D. lancangensis GMB0045 MW797113 MW814885 This study
D. lancangensis GMB0046 MW797114 MW814886 This study
D. lancangensis GMB0047 MW797116 MW814887 This study
D. palmicola MFLUCC 11-0020 KP744438 NA Liu et al. (2015)
D. palmicola MFLUCC 11-0018 KP744439 NA Liu et al. (2015)
D. spilomea D17C AJ302433 NA Acero et al. (2004)
D. stigma DCASH200 GQ293947 GQ294003 Trouillas et al. (2010)
D. undulata D20C, CBS 271.87 AJ302436 NA Acero et al. (2004)
Diatrypella atlantica HUEFS 136873 KM396614 KR259647 de Almeida et al. (2016)
D. banksiae CPC 29118 KY173402 NA Crous et al. (2013)
D. delonicis MFLUCC 15-1014 MH812994 MH847790 Hyde et al. (2019)
D. delonicis MFLU 16-1032 MH812995 MH847791 Hyde et al. (2019)
D. elaeidis MFLUCC 15-0279 MN308417 MN340300 Konta et al. (2020)
D. favacea Islotate 380 KU320616 NA de Almeida et al. (2016)
D. favacea DL26C AJ302440 NA Unpublished
D. frostii UFMGCB 1917 HQ377280 NA Vieira et al. (2011)
D. heveae MFLUCC 15-0274 MN308418 MN340301 Konta et al. (2020)
D. heveae MFLUCC 17-0368 MF959501 MG334557 Senwanna et al. (2017)
D. hubeiensis CFCC 52413 MW632937 NA Zhu et al. (2021)
D. iranensis KDQ18 KM245033 KY352429 Mehrabi et al. (2015)
D. macrospora KDQ15 KR605648 KY352430 Mehrabi et al. (2016)
D. oregonensis (Diatrype oregonensis) DPL200 GQ293940 GQ293999 Trouillas et al. (2010)
D. oregonensis (Diatrype oregonensis) CA117 GQ293934 GQ293996 Trouillas et al. (2010)
D. pseudooregonensis GMB0039 MW797115 MW814888 This study
D. pseudooregonensis GMB0040 MW797117 MW814889 This study
D. pseudooregonensis GMB0041 MW797118 MW814890 This study
D. pseudooregonensis GMB0042 MW797119 MW814891 This study
D. pseudooregonensis GMB0043 MW797120 MW814892 This study
D. pseudooregonensis GMB0044 MW797110 MW814882 This study
D. pulvinata H048 FR715523 FR715495 de Almeida et al. (2016)
D. pulvinata DL29C AJ302443 NA Unpublished
D. tectonae MFLUCC 12-0172a KY283084 NA Shang et al. (2017)
D. tectonae MFLUCC 12-0172b KY283085 KY421043 Shang et al. (2017)
D. verruciformisT UCROK1467 JX144793 JX174093 Lynch et al. (2013)
D. verruciformisT UCROK754 JX144783 JX174083 Lynch et al. (2013)
D. vulgaris HVFRA02 HQ692591 HQ692503 Trouillas et al. (2011)
D. vulgaris HVGRF03 HQ692590 HQ692502 Trouillas et al. (2011)
D. vulgaris GMB0051 MW797107 MW814879 This study
D. yunnanensis VT01 MN653008 MN887112 Zhu et al. (2021)
Eutypa armeniacae ATCC 28120 DQ006948 DQ006975 Rolshausen et al. (2006)
E. astroidea E49C, CBS 292.87 AJ302458 DQ006966 Rolshausen et al. (2006)
E. cerasi GMB0048 MW797104 MW814893 This study
E. cerasi GMB0049 MW797105 MW814877 This study
E. flavovirens E48C, CBS 272.87 AJ302457 DQ006959 Rolshausen et al. (2006)
E. laevata E40C CBS 291.87 AJ302449 NA Acero et al. (2004)
E. lataT CBS290.87 HM164736 HM164770 Trouillas and Gubler (2010)
E. lataT EP18 HQ692611 HQ692501 Trouillas et al. (2011)
E. lataT RGA01 HQ692614 HQ692497 Trouillas et al. (2011)
E. leioplaca CBS 248.87 DQ006922 DQ006974 Rolshausen et al. (2006)
E. leptoplaca CBS 287.87 DQ006924 DQ006961 Rolshausen et al. (2006)
E. maura CBS 219.87 DQ006926 DQ006967 Rolshausen et al. (2006)
E. microasca BAFC 51550 KF964566 KF964572 Grassi et al. (2014)
E. sparsa 3802 3b AY684220 AY684201 Trouillas and Gubler (2004)
E. tetragona CBS 284.87 DQ006923 DQ006960 Rolshausen et al. (2006)
Eutypella caricae EL51C AJ302460 NA Acero (2000)
E. cerviculataT M68 JF340269 NA Arhipova et al. (2012)
E. cerviculataT EL59C AJ302468 NA Acero et al. (2004)
E. leprosa EL54C AJ302463 NA Acero et al. (2004)
E. leprosa Isolate 60 KU320622 NA de Almeida et al. (2016)
E. microtheca BCMX01 KC405563 KC405560 Paolinelli-Alfonso et al. (2015)
E. parasitica CBS 210.39 DQ118966 NA Jurc et al. (2006)
E. semicircularis MP4669 JQ517314 NA Mehrabi et al. (2016)
Halocryptovalsa salicorniae MFLUCC 15-0185 MH304410 MH370274 Dayarathne et al. (2020)
Halodiatrype avicenniae MFLUCC 15-0953 KX573916 KX573931 Dayarathne et al. (2016)
H. salinicolaT MFLUCC 15-1277 KX573915 KX573932 Dayarathne et al. (2016)
Kretzschmaria deusta CBS 826.72 KU683767 KU684190 U’Ren et al. (2016)
Monosporascus cannonballus T CMM3646 JX971617 NA Unpublished
M. cannonballus T ATCC 26931 FJ430598 NA Unpublished
Neoeutypella baoshanensisT GMB0052 MW797106 MW814878 This study
N. baoshanensisT LC 12111 MH822887 MH822888 Hyde et al. (2019)
N. baoshanensisT EL51C, CBS 274.87 AJ302460 NA Acero et al. (2004)
N. baoshanensisT MFLUCC 16-1002 MT310662 NA Phukhamsakda et al. (2020)
N. baoshanensisT GL08362 JX241652 NA Gao et al.(2013)
Paraeutypella citricola HVVIT07 HQ692579 HQ692512 Trouillas et al. (2011)
Pa. citricola HVGRF01 HQ692589 HQ692521 Trouillas et al. (2011)
Pa. citricola GMB0053 MW797109 MW814881 This study
Pa. guizhouensisT KUMCC 20-0016 MW039349 MW239660 Dissanayake et al. (2021)
Pa. guizhouensisT KUMCC 20-0017 MW036141 MW239661 Dissanayake et al. (2021)
Pa. vitis UCD2291AR HQ288224 HQ288303 Úrbez-Torres et al. (2012)
Pa. vitis UCD2428TX FJ790851 GU294726 Úrbez-Torres et al. (2009)
Pedumispora rhizophoraeT BCC44877 KJ888853 NA Klaysuban et al. (2014)
Pe. rhizophoraeT BCC44878 KJ888854 NA Klaysuban et al. (2014)
Peroneutypa alsophila EL58C, CBS 250.87 AJ302467 NA Acero et al. (2004)
Pe. curvispora HUEFS 136877 KM396641 NA de Almeida et al. (2016)
Pe. diminutispora MFLUCC 17-2144 MG873479 NA Shang et al. (2018)
Pe. mackenziei MFLUCC 16-0072 KY283083 KY706363 Shang et al. (2017)
Pe. mangrovei PUFD526 MG844286 MH094409 Phookamsak et al. (2019)
Pseudodiatrype hainanensisT GMB0054 MW797111 MW814883 This study
Ps. hainanensisT GMB0055 MW797112 MW814884 This study
Quaternaria quaternata EL60C, CBS 278.87 AJ302469 NA Acero et al. (2004)
Q. quaternata GNF13 KR605645 NA Mehrabi et al. (2016)
Xylaria hypoxylon CBS 122620 AM993141 KX271279 Peršoh et al. (2009)

DNA extraction, Polymerase Chain Reaction (PCR) and phylogenetic analyses

Genomic DNA was extracted from fungal mycelium following the manufacturer’s protocol of the BIOMIGA Fungal gDNA isolation Kit (BIOMIGA, Hangzhou City, Zhejiang Province, China). Extracts of DNA were stored at –20 °C.

PCR was carried out in a volume of 25 μl containing 9.5 μl of ddH2O, 12.5 μl of 2× Taq PCR Master Mix (2 × Taq Master Mix with dye, TIANGEN, China), 1 μl of DNA extracts and 1 μl of forward and reverse primers (10 μM each) in each reaction. Primers pairs, ITS4 and ITS5, fRPB2-7CR and fRPB2-5f, LROR and LR5, T1 and Bt2b, as well as Bt2a and Bt2b (Vilgalys and Hester 1990; White et al. 1990; Glass and Donaldson 1995; O’Donnell and Cigelnik 1997), were used to amplify internal transcribed spacer (ITS) sequences, RNA polymerase II second largest subunit (RPB2) sequences, large subunit ribosomal (LSU) sequences and β-tubulin (TUB2) sequences, respectively.

PCR profiles for the ITS and LSU are as follows: initially at 95 °C for 5 minutes, followed by 35 cycles of denaturation at 94 °C for 1 minute, annealing at 52 °C for 1 minute, elongation at 72 °C for 1.5 minutes and a final extension at 72 °C for 10 minutes. PCR profile for the RPB2 is as follows: initially at 95 °C for 5 minutes, followed by 35 cycles of denaturation at 95 °C for 1 minute, annealing at 54 °C for 2 minutes, elongation at 72 °C for 1.5 minutes and a final extension at 72 °C for 10 minutes (Konta et al. 2020). PCR profile for the TUB2 are as follows: initially at 95 °C for 5 minutes, followed by 35 cycles of denaturation at 94 °C for 1 minute, annealing at 52 °C for 1 minute, elongation at 72 °C for 1.5 minutes and a final extension at 72 °C for 10 minutes (de Almeida et al. 2016). PCR products were submitted to Sangon Biotech, Shanghai, China for purification and sequencing.

Phylogenetic analyses

Phylogenetic analyses were performed by searching homologous sequence data of the family Diatrypaceae in the GenBank database, selected from NCBI and recently published papers (Mehrabi et al. 2019; Dayarathne et al. 2020; Konta et al. 2020; Dissanayake et al. 2021; Zhu et al. 2021). After the preliminary identification results of the sequences, multiple sequence alignments (ITS and β-tubulin) were aligned using BioEdit v. 7.0 (Hall 1999). Alignments were converted from FASTA to PHYLIP format by using Alignment Transformation Environment online (https://sing.ei.uvigo.es/ALTER/, Glez-Peña et al. 2010). Maximum Likelihood (ML) analyses and Bayesian posterior probabilities (BYPP) were performed by using RAxML-HPC BlackBox (8.2.12) and MrBayes on XSEDE (3.2.7a) tools in the CIPRES Science Gateway platform, based on a combination of ITS and TUB2 sequence data (Miller et al. 2010). Both of the two methods use the GTR+I+G model of evolution (Nylander 2004). The Bootstrap supports of ML analyses were obtained by running 1,000 pseudo-replicates and BYPP using a simulation technique called Markov chain Monte Carlo (or MCMC) to approximate the posterior probabilities of trees. Six simultaneous Markov Chains were run for 3,000,000 generations and trees were sampled every 1,000th generation. Finally, the tree was visualised in FigTree v.1.4.4 (Rambaut 2012) and edited by using Adobe Photoshop CS6 software. The final alignment and phylogenetic trees were deposited in TreeBASE under the submission ID28176 (http://www.treebase.org/)

Result

Phylogenetic analyses

Based on RAxML and BYPP analyses, phylogenetic analyses were similar in overall tree topologies and did not differ significantly. The dataset consists of 105 taxa for representative strains of species in Diatrypaceae, including outgroup taxa with 1071 characters, including gaps (ITS: 1–486, β-tubulin: 486–1071). The RAxML analyses resulted in a best scoring likelihood tree selected with a final ML optimisation likelihood value of -15731.506304, which is shown in Fig. 1.

Figure 1. 

Phylogram generated from Maximum Likelihood (RAxML) analyses, based on ITS-β-tubulin matrix. ML bootstrap supports (≥ 70%) and Bayesian posterior probability (≥ 0.90) are indicated as ML/BYPP. The tree is rooted to Kretzschmaria deusta (CBS 826.72) and Xylaria hypoxylon (CBS 122620). Ex-type strains are in red. Newly generated strains are in black bold.

The phylogenetic tree, based on combining ITS and β-tubulin sequence data, is also shown in Fig. 1 and contains 17 clades within Diatrypaceae. Below, we list the placements of new taxa:

Clade 1: Diatrypella pseudooregonensis and Diatrypella oregonensis clustered with the species of Diatrypella in Clade 1 with high bootstrap support, Diatrypella pseudooregonensis is introduced as an 8-spored new species of Diatrypella and Diatrype oregonensis is renamed as Diatrypella oregonensis.

Clade 4: Pseudodiatrype formed a separate branch in a clade (Clade 4) basal to the genus Allodiatrype.

Clade 7: Diatrype lancangensis clusters with the species of Diatrypella and Diatrype in an unresolved clade. However, Diatrype and Diatrypella have previously shown confused classification which is difficult to distinguish, based on phylogenetic aspects alone. Therefore, we introduce Diatrype lancangensis as a new species of Diatrype, based on phylogenetic analyses and morphological differences (Table 2).

Table 2.

The dimensions of the present species and some related species of Diatrype and Allodiatrype.

Species name Stromata Asci Ascospores Reference
Length (mm) Wide (mm) Length (μm) Wide (μm) Length (μm) Wide (μm)
Allodiatrype arengae 0.69–0.94 0.37–0.93 54–109 6–10 7–10 2–3 Konta et al. (2020)
A. elaeidicola 1.2–2.8 0.9–1.66 60–91 4–7 8–10 1.5–3 Konta et al. (2020)
A. elaeidis 0.47–0.86 0.44–0.71 56–95 9–11 8–10 1.5–3 Konta et al. (2020)
A. thailandica NA 1–2 55–80 5–7 3.8–6.9 1–1.4 Li et al. (2016)
Diatrype acericola 1–2 1–1.5 23–27 5–7 7.5–9 0.9–1.1 Vasilyeva and Ma (2014)
D. albopruinosa 0.5–1 diam. 0.5–1 diam 40–60 10–15 12–15 3.5–4 Vasilyeva and Ma (2014)
D. bullata 2–7 diam. 2–7 diam 25–30 5–7 7.5–9 Very thin Vasilyeva and Ma (2014)
D. disciformis NA NA 75–115 NA 5–9 1.5–2 Senanayake et al. (2015)
D. enteroxantha NA 1–3.5 18–28.5 5–9 7–10 1.5–2.5 de Almeida et al. (2016)
D. hypoxyloides NA NA 20–25 4–6 4–6 Very thin Vasilyeva and Ma (2014)
D. lancangensis NA NA 90.5–160.5 7–15 11–18.5 2–4 This study
D. lijiangensis 1 diam. 1 diam 50–90 6–9 6–8 1–2 Thiyagaraja et al. (2019)
D. macounii 1–1.8 diam. 1–1.8 diam 25–30 4–6 4–6 0.7–1 Vasilyeva and Ma (2014)
D. stigma NA NA 25–30 5–7 6–8 1.5–2 Vasilyeva and Ma (2014)
D. subundulata NA NA 35–40 5–7 7–9 1.7–1.9 Vasilyeva and Ma (2014)
D. undulata NA NA 25–30 3.5–4.5 5–7 0.9–1.3 Vasilyeva and Ma (2014)
D. whitmanensis NA NA 50–82 8–15 7.5–10 1–1.5 Trouilla et al. 2010
Pseudodiatrype hainanensis 2–3.6 1.6–3 110–155.5 6–10 8.5–13 1.5–2.5 This study

Clade 8: Eutypa cerasi forms a distinct lineage which is sister to Eutypa lata (EP18, RGA01) (Fig. 1).

Taxonomy

Diatrype Fr.

Notes

The genus Diatrype was introduced by Fries (1849). The genus is characterised by stromata widely effuse or verrucose, flat or slightly convex, with discoid or sulcate ostioles at the surface, 8-spored and long-stalked asci and hyaline or brownish, allantoid ascospores. In this study, we introduce a new species of Diatrype from China.

Diatrype lancangensis S.H. Long & Q. R. Li, sp. nov.

MycoBank No: 839655
Fig. 2

Holotype

GMB0045.

Etymology

Refers to the name of the location, where the type specimen was collected.

Description

Saprobic on decaying branches of an unidentified plant. Sexual morph: Stromata immersed in bark, aggregated, irregular in shape, widely effused, flat, margin diffuse, surface dark brown to black, with punctiform ostioles scattered at surface, with tissues soft, white between perithecia. Entostroma dark with embedded perithecia in one layer. Perithecium semi-immersed in stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous or longicollous 283.5–343.5 μm high, 207–290 μm broad (av. = 315.5 × 248.0 μm, n = 10), ovoid, obovoid to oblong, monostichous, aterrimus. Ostiole opening separately, papillate or apapillate, central. Peridium 30–50 μm thick, dark brown to hyaline with textura angularis cell layers. Asci 90.5–160.5 × 7.0–15.0 μm (av. = 129.5 × 10.5 μm n = 30) 8-spored clavate, unitunicate, with rounded apex, apical rings inamyloid. Ascospores 11–18.5 × 2–4 μm (av. = 14.9 × 2.8 μm, n = 30), irregularly arranged, allantoid, slightly curved, brown to dark brown, smooth, aseptate, usually with oil droplets. Asexual morph: undetermined.

Figure 2. 

Diatrype lancangensis (GMB0045, holotype) A stromata on host substrate B, C stromata on host D transverse sections through ascostroma E vertical section through ascostroma F culture on PDA G ostiolar canal H peridium I–K ascospores L–N asci. Scale bars: 10 μm (G–N).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became luteous, dense but, thinning towards edge, margin rough, white from above, reverse white at margin, pale yellow to luteous at centre, no pigmentation produced on PDA medium, no conidia observed on PDA or on OA media.

Specimens examined

China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'17.44"N, 99°35'10.05"E) on branches of an unidentified plant, 4 October 2019. Altitude: 2549 m., Y.H. Pi & Qiong Zhang, LC172 (GMB0045, holotype, KUN-HKAS 112664, isotype, ex-type living culture GMBC0045).

Additional specimens examined

China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'17.44"N, 99°35'10.05"E) on branches of an unidentified plant, 4 October 2019. Altitude: 2549 m., Y.H. Pi and Qiong Zhang, LC173 (GMB0046, KUN-HKAS 112665, living culture GMBC0046); CHINA, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'15.48"N, 99°35'24.08"E) on branches of an unidentified plant, 5 October 2019. Altitude: 2623 m., Y.H. Pi and Qiong Zhang, LC262 (GMB0047, KUN-HKAS 112672, living culture GMBC0047).

Additional sequences

GMB0045 (LSU: MW797057, RPB2: MW81490); GMB00046 (LSU: MW797058); GMB0047 (LSU: MW797060, RPB2: MW814903)

Note

Our new strain, GMBC0045 falls into the unresolved clade (Clade 7) which comprises five Diatrypella and one Diatrype species (Fig. 1), this clade is consistent with the study of Konta et al. (2020). The taxonomic confusion of Diatrypaceae has led to difficulties in separating the genera. We consider that the new species belongs to the genus Diatrype, based on the stromata features mentioned above which closely resemble descriptions of Diatrype subundulata Lar. N. Vassiljeva & Hai X. Ma and Diatrype undulata (Pers.) Fr. (Vasilyeva et al. 2014). However, the ascospores of these species are larger than the ascospores of D. subundulata and D. undulata (Table 2). Phylogenetic analyses also showed that D. lancangensis falls on a separate branch that clustered with species of Diatrypella and Diatrype (Fig. 1). Hence, by combining morphological characteristics and phylogenetic analyses, it seems appropriate to categorise this species as Diatrype.

In the phylogenetic analyses, it can be seen that Clade 7 can be defined as a new genus, but it is difficult to find the common morphological similarities among these species. More specimens and sequence or chemical composition analysis are needed in the future to determine whether Clade 7 can be a new genus. The characteristics of the stromata of Diatrypella spp. in clade 7 are solitary and scattered, which is distinctly different from widely effuse, flat and slightly convex stromata of Diatrype lancangensis and Diatrype palmicola (Liu et al. 2015; Hyde et al. 2020b; Zhu et al. 2021). And in the recent study, Zhu et al. (2021) proposed that the species of Diatrypella in Clade 7 were isolated from Betula spp., it may have host specificity. Because of the above two reasons, we think it is better to classify our strains into Diatrype.

Pseudodiatrype S.H. Long & Q.R. Li, gen. nov.

MycoBank No: 839658

Etymology

Refers to this genus resembling Diatrype in morphology, but it is phylogenetically distinct.

Type species

Pseudodiatrype hainanensis S. H. Long & Q.R. Li sp. nov.

Description

Saprobic on decaying branches of an unidentified plant. Sexual morph: Stromata scattered or aggregated on host, wart-like, pustulate, visible as black, rounded to irregular in shape on host surface, erumpent through host bark, 5–20 ascomata immersed in one stroma. Endostroma consists of outer layer of black, small, dense, thin parenchymal cells and inner layer of white, large, loose parenchymal cells, thin, pale yellow, powdery near margin of the black cells. Ostiole opening through host bark and appearing as black spots, separately, papillate or apapillate, central. Perithecium immersed in stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous or longicollous. Peridium is composed of an outer layer of dark brown to black, thin-walled cells, arranged in textura angularis, the inner layer of hyaline thin-walled cells of textura angularis. Asci 8-spored, unitunicate, clavate, long-stalked, apically rounded, apical rings inamyloid. Ascospores irregularly arranged, allantoid, slightly or moderately curved, smooth, subhyaline, aseptate, usually with two oil droplets. Asexual morph: undetermined.

Note

The genus Pseudodiatrype is introduced to accommodate the new collection made from Hainan Province of China and typified by Pseudodiatrype hainanensis. Pseudodiatrype is monotypic and, morphologically, resembles Diatrype and Allodiatrype Konta & K.D. Hyde. However, Pseudodiatrype can be distinguished from Diatrype by its 5–20 ascomata immersed in a stroma, while the stroma of species of Diatrype is distributed over large areas, sometimes covering the surface of the host (Vasilyeva and Ma 2014; Konta et al. 2020). Pseudodiatrype differs from Alloiatrype by having its 5–20 ascomata immersed in a stroma, whereas the stroma of Allodiatrype has only 1–10 ascomata. Moreover, the endostroma of Allodiatrype is composed of dark brown outer layer cells and yellow inner layer cells (Konta et al. 2020), which are different from the endostroma of Pseudodiatrype having black outer and inner cells surrounded by powdery, pale yellow cells. In addition, the sizes of stroma and ascospores are different from species of Diatrype and Allodiatrype (Table 2). In the phylogenetic analyses, species of Pseudodiatrype appeared in a separate branch which is distinct from other genera within Diatrypaceae (Fig. 1), thus, justifying the erection of the new genus Pseudodiatrype.

Pseudodiatrype hainanensis S. H. Long & Q.R. Li, sp. nov.

MycoBank No: 839659
Fig. 3

Holotype

GMB0054.

Etymology

Refers to the location of collections, Hainan Province.

Description

Saprobic on decaying branches of an unidentified plant. Sexual morph: Stromata wart-like, pustulate, 2–3.6 mm long and 1.6–3 mm broad (av. = 3.2 × 1.9 mm, n = 30), about 2 mm thick, 5–20 in single stroma, visible as black, rounded to irregular in shape on the host surface, erumpent through host bark, solitary to gregarious. Endostroma composed of an outer layer of dark brown to black, small, tightly packed, thin parenchymatous cells and an inner layer of white, large, loose parenchymal cells with powdery, thin, yellowish tissue. Ostiole opening separately, papillate or apapillate, central. Perithecium immersed in the stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous or longicollous, 193–347 μm high, 138–206 μm diam. (av. = 278 × 156 μm, n = 10). Peridium 30–50 μm thick, dark brown to hyaline with textura angularis cell layers. Asci 110–155.5 × 6–10 μm (av. = 132 × 8 μm, n = 30), 8-spored, unitunicate, clavate, long-stalked, apically rounded with inamyloid rings. Ascospores 8.5–13 × 1.5–2.5 μm (av. = 10.5 × 2 μm, n = 30), irregularly arranged, allantoid, slightly or moderately curved, smooth, subhyaline, aseptate, usually with two oil droplets. Asexual morph: undetermined.

Figure 3. 

Pseudodiatrype hainanensis (GMB0054, holotype) A stromata on host substrate B, C stromata on host D transverse section through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H ostiolar canal I peridium J–M ascospores N–P asci. Scale bars: 40 μm (G); 10 μm (H–P).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became pale brown, dense, but thinning towards edge, fluffy to slightly fluffy, white from above, pale brown from below, no pigmentation produced on PDA medium, no conidia observed on PDAor on OA media.

Specimens examined

China, Hainan Province, Wuzhishan City, Wuzhishan Nature Reserve (18°54'21.81"N, 109°40'54.12"E) on branches of unidentified plant, 14 November 2020. Altitude: 775 m. Y.H. Pi & Q.R. Li, WZS59 (GMB0054, holotype, KUN-HKAS 112700, isotype, ex-type living culture GMBC0054).

Additional specimen examined

China, Hainan Province, Wuzhishan City, Wuzhishan Nature Reserve (18°54'21.81"N, 109°40'54.12"E) on branches of an unidentified plant, 14 November 2020. Altitude: 775 m, Y.H. Pi & Q.R. Li, WZS66 (GMB0055, living culture GMBC0055)

Additional sequences

GMB0054 (LSU: MW797055, RPB2: MW814900); GMB0055 (LSU: MW797056, RPB2 MW814901).

Note

A peculiar feature of Pseudodiatrype hainanensis is the composition of endostroma. There are black outer layer cells, white inner layer cells and powdery, yellowish cells that are smaller than the white cells at the edge of the endostroma near the black cells in endostroma.

Diatrypella (Ces. & De Not.) De Not.

Notes

The genus Diatrypella was introduced by Cesati & De Notaris (1863) and was typified with Diatrypella verruciformis (Ehrh.) Nitschke. This genus was characterized by pustule-like stromata erumpent through the host surface, polysporous asci and allantoid ascospores and libertella-like asexual morphs (Senanayake et al. 2015; Hyde et al. 2017; Shang et al. 2017). In this study, we introduce a new species, a new combination and a new record of Diatrypella vulgaris from Guizhou Province for China.

Diatrypella pseudooregonensis S.H. Long & Q.R. Li, sp. nov.

MycoBank No: 839656
Fig. 4

Holotype

GMB0041

Etymology

Refers to its similar species of Diatrype oregonensis.

Description

Saprobic on decaying branches of unidentified plant. Sexual morph: Stromata pustulate, with groups of 3–16 perithecia, rugose, visible as black, erumpent, scattered, surrounded by a thin, black line in host tissue, solitary to gregarious, 1–3 mm long and 0.5–2 mm broad (av. = 2 × 1.5 mm, n = 30), about 1 mm thick. Endostroma white to light yellow. Ostiole opening separately, papillate or apapillate, central. Perithecium immersed in stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous or longicollous 218.5–465 μm high, 112–257 μm diam. (av. = 306 × 164 μm, n = 10), globose to subglobose, glabrous, ostioles individual. Peridium: 30–50 μm thick, dark brown to hyaline with textura angularis cell layers. Asci 95–149 × 6.5–11.5 μm (av. = 120 × 10.5 μm, n = 30), 8-spored, unitunicate, clavate or cylindrical, long-stalked, apically rounded, apical rings inamyloid. Ascospores 11–16 × 1.5–3.5 μm (av. = 14 × 2.5 μm, n = 30), irregularly arranged, allantoid, slightly or moderately curved, subhyaline to slightly brown, smooth, aseptate, usually with two oil droplets. Asexual morph: undetermined.

Figure 4. 

Diatrypella pseudooregonensis (GMB0041, holotype) A stromata on host substrate B, C stromata on host substrate D transverse section through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H ostiolar canal I, J asci K–N ascospores. Scale bars: 20 μm (G); 10 μm (H–N).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became pale brown, dense, but thinning towards the edge, margin rough, white from above, white at margin and light brown at centre from below, no pigmentation produced on PDA medium, no conidia observed on PDA or on OA media.

Specimens examined

China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'19.88"N, 99°35'30.68"E) on branches of an unidentified plant, 5 October 2019. Altitude: 2677 m, Y.H. Pi & Qiong Zhang, LC323 (GMB0041, holotype, KUN-HKAS 112646, isotype, ex-type living culture GMBC0041)

Additional specimens examined

China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'13.51"N, 99°35'25.59"E) on branches of an unidentified plant, 6 October 2019. Altitude: 2630 m, Y.H. Pi & Qiong Zhang, LC384 (GMB0043, KUN-HKAS 112681, living culture GMBC0043); China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'15.00"N, 99°35'39.73"E) on branches of an unidentified plant, 5 October 2019. Altitude: 2698 m, Y.H. Pi & Qiong Zhang, LC312 (GMB0040, KUN-HKAS 112674, living culture GMBC0040); China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°35'19.09"N, 99°35'19.09"E) on branches of an unidentified plant, 5 October 2019. Altitude: 2569 m, Y.H. Pi & Qiong Zhang, LC193 (GMB0039, KUN-HKAS 112667, living culture GMBC0039); China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°1'9.11"N, 99°35'24.80"E) on branches of an unidentified plant, 5 October 2019. Altitude: 2649 m, Y.H. Pi & Qiong Zhang, LC335 (GMB0042, KUN-HKAS 112647, living culture GMBC0042); China, Guizhou Province, Anshun City, Pingba District (26°25'9.65"N, 106°24'24.48"E) on branches of an unidentified plant, 1 August 2020. Altitude: 1250 m, Y.H.Pi, PB51 (GMB0044, KUN-HKAS 112693, living culture GMBC0044).

Additional sequences. GMB0041 (LSU: MW797062, RPB2: MW814906); GMB0043 (LSU: MW797064, RPB2: MW814907); GMB0040 (LSU: MW797061, RPB2: MW814905); GMB0039 (LSU: MW797059, RPB2: MW814904); GMB0042 (LSU: MW797063); GMLB0044 (LSU: MW979054, RPB2: MW814899).

Note. Morphologically, Diatrype has 8 ascospores in a single ascus, while Diatrypella has more than eight ascospores in each ascus (Senanayake et al. 2015). However, previous research (e.g. Acero et al. 2004 and Trouillas et al. 2011) suggested that both Diatrypella and Diatrype are polyphyletic within the family. In the phylogenetic analyses, Diatrypella pseudooregonensis grouped closely to the D. verruciformis and thus, we consider this new species to belong in the genus Diatrypella, because it is doubtful whether the number of ascospores per asci is useful as a basis for generic classification.

Diatrypella vulgaris Trouillas, W.M. Pitt & Gubler, Fungal Diversity 49: 212 (2011)

MycoBank No: 519404
Fig. 5

Description

Saprobic on decaying branches of an unidentified plant. Sexual morph: Stromata scattered on the host, 0.8–1.5 mm long and 0.8–2 mm broad (av. = 1.2 × 1.3 mm, n = 30) pustulate, visible as black, rounded to irregular in shape on host surface, semi-immersed, erumpent through host bark, with 2–8 ascomata immersed in one stroma. Endostroma consists of outer dark brown, small, dense, thin parenchymal cells and an inner layer of white, large, loose parenchymal cells. Ostiole opening separately, papillate or apapillate, central 710.7–787.2 μm high, 270.2–422 μm diam. (av. = 742 × 363 μm, n = 10). Perithecium immersed in stroma, round to oblong, with cylindrical neck, brevicollous or longicollous. Peridium composed of outer layer of dark brown to black, thin-walled cells, arranged in textura angularis, inner layer of hyaline thin-walled cells of textura angularis. Asci 111.4–152.9 × 10.6–17.5 μm (av. = 124.5 × 15.5 μm, n = 30), polysporous, clavate, long-stalked, apically rounded. Ascospores 8–11 × 1–2 μm (av. = 8.9 × 1.7 μm, n = 30), overlapping, crowded, allantoid, slightly or moderately curved, smooth, subhyaline, yellowish in mass, aseptate, usually with two oil droplets. Asexual morph: undetermined.

Figure 5. 

Diatrypella vulgaris (GMB0051, new record for China) A stromata on host substrate; B, C close-up of stroma D transverse sections through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H, I ostiolar canal J, K asci L–O ascospores. Scale bars: 20 μm (G); 10 μm (H–I).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became pale brown, dense, but thinning towards edge, medium dense, white from above, reverse side white at margin, flesh to pale brown at centre, no pigmentation produced on PDA medium, no conidia observed on PDA or on OA media.

Specimens examined

China, Guizhou Province, Guiyang City, Gaopo Township (26°29'72.02"N, 106°29'55.57"E), on branches of unidentified plant, 30 October 2020. Altitude: 1589 m, S.H. Long, GP02 (GMB0051, KUN-HKAS 112697, living culture GMBC0051).

Additional sequences

GMB0051 (LSU: MW797051, RPB2: MW814897).

Note

The comparison of ITS sequences in NCBI showed that this isolate is 100% similar to the strain of Diatrypella vulgaris (HVGRF03), isolated from holotype specimens. Morphologically, GMB0051 shows the same features as Diatrypella vulgaris. The stromata of these specimens are similar, but ascospores of GMB0051 are thinner than those of the HVGRF03 (8–10 × 2–2.5 μm) and, when compared with the ascospores of strain MFLUCC 17-0128 (4.5–7.5 × 1–2 μm), they are shorter than GMB0051 (Trouillas et al. 2011; Hyde et al. 2017). Here, we use the ITS sequence similarity between the new collection and the type strain of Diatrypella vulgaris as the identification tool. Diatrypella vulgaris has been reported in Austria and Thailand (Trouillas et al. 2011, Hyde et al. 2017). This is the first report of Diatrypella vulgaris from China.

Diatrypella oregonensis (Wehm.) S.H. Long & Q.R. Li, comb. nov.

MycoBank No: 839728

Eutypella oregonensis Wehm. Pap. Mich. Acad. Sci. 11: 163 (1930)

Diatrype oregonensis (Wehm.) Rappaz, Mycol. helv. 2(3): 420 (1987)

Description

See Trouillas et al. (2010).

Note

The strains of Diatrype oregonensis (DPL200, CA117) generated from Trouillas et al. (2010) grouped in Diatrypella s. str. Diatrype oregonensis was erected in 1930 as Eutypella oregonensis (Kauffman 1930). No available sequences from type material were found. After re-examination of holotype specimen of Diatrype oregonensis, Trouillas et al. (2010) introduced two strains of Diatrype oregonensis (DPL200 and CA117). Although neither of these strains are ex-type, they are, the most authoritative strains. Here, we tentatively transfer Diatrype oregonensis to Diatrypella as Diatrypella oregonensis, based on the phylogenetic analyses (Fig. 1). Diatrypella oregonensis is similar to D. pseudooregonensis in having 8-spored asci (Rappaz 1987; Trouillas et al. 2011). Nevertheless, we consider that the number of ascospores as a basis for distinguishing Diatrypella from Diatrype is not useful.

Allodiatrype Konta & K.D. Hyde Mycosphere 11(1): 247 (2020)

Notes

The genus Allodiatrype was introduced by Konta et al. (2020), which was characterised by regular or irregular-shaped stromata, erumpent through host surface, asci with 8 spores and aseptate, allantoid ascospores. In this study, we introduce a new record of Allodiatrype thailandica (R.H. Perera et al.) Konta & K.D. Hyde collected from Yunnan Province in China.

Allodiatrype thailandica (R.H. Perera et al.) Konta & K.D. Hyde, Mycosphere 11(1): 253 (2020)

MycoBank No: 556932
Fig. 6

Diatrype thailandica R.H. Pereraet al., Fungal Diversity 78: 1–237, [105] (2016)

Description

Saprobic on decaying branches of unidentified plant. Sexual morph: Stromata wart-like, pustulate, 0.5–1.8 mm long and 0.8–2.2 mm broad (av. = 1.2 × 1.3 mm, n = 30), about 1 mm thick, 1–18 in a single stroma, visible as black, rounded to irregular in shape on the host surface, erumpent through host bark, solitary to gregarious. Endostroma composed of an outer layer of dark brown to black, small, tightly packed, thin parenchymatous cells and an inner layer of white to yellow, large, loose parenchymal cells. Ostiole opening separately, papillate or apapillate, central. Perithecium immersed in stroma, globose to subglobose, glabrous, with cylindrical short neck, 377–447 μm high, 191–264 μm diam. (av. = 406 × 221 μm, n = 10). Peridium hyaline to dark brown with textura angularis cell layers. Asci 80–113.5 × 6.9–10 μm (av. = 109.3 × 8.5 μm, n = 30), 8-spored, unitunicate, clavate, long-stalked, upper part inflated, apically rounded to truncate, apical rings inamyloid. Ascospores 6–11 × 2–2.5 μm (av. = 8.9× 2.3 μm, n = 30), irregularly arranged, allantoid, slightly curved, smooth, subhyaline, aseptate, usually with two oil droplets. Asexual morph: undetermined.

Figure 6. 

Allodiatrype thailandica (GMB0050, new record for China) A stromata on host substrate B, C close-up of stromata D transverse section through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H ostiolar canal I–K ascospores L–N asci. Scale bars: 20 μm (G); 10 μm (H–N).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became pale yellow, irregular in shape, medium dense, flat or effuse, slightly raised, with edge fimbriate, fluffy to fairly fluffy, white from above, reverse side white at margin, pale brown at centre, no pigmentation produced on PDA medium, no conidia observed on PDA or on OA media.

Specimens examined

China, Yunnan Province, Baoshan City, Lancang River Nature Reserve (24°57'25.35"N, 99°44'22.82"E), on branches of unidentified plant, 2 October 2019. Altitude: 1317 m, Y.H. Pi & Qiong. Zhang, LC103 (GMB0050, KUN-HKAS 112660, living culture GMBC0050).

Additional sequences

GMB0050 (LSU: MW797052).

Note

The ITS sequence data were subjected to BLAST in NCBI and the results showed that it is 100% similar to Allodiatrype thailandica. Additionally, based on morphological and phylogenetic analyses, this strain was identified as the A. thailandica. The stromata are similar, but the ascospores of GMB0050 are longer and wider than the ascospores of strain MFLUCC 15-3662 (3.8–6.9 × 1–1.4 μm) isolated from the holotype specimen, but it is similar to the strain MFLU 17-0735 (6.5–10.7 × 1.6–2.7 μm) (Perera et al. 2020). Here, we use the ITS sequence similarity between the new collection and the type strain of Allodiatrype thailandica as basis for identification. A. thailandica has been reported in Thailand in 2016 as Diatrype thailandica and recognised as A. thailandica by Konta et al. (2020). This is the first report of Allodiatrype thailandica from China.

Neoeutypella M. Raza, Q.J. Shang, Phookamsak & L. Cai, Fungal Diversity 95: 167 (2019)

Note

The genus Neoeutypella was introduced by Phookamsak et al. (2019) and is characterised by carbonaceous stromata immersed or semi-immersed on the host, 8-spored asci and hyaline or pale reddish-brown to brown ascospores. In this study, we introduce a new collection of N. baoshanensis, isolated from Guizhou Province in China.

Neoeutypella baoshanensis M. Raza, Q.J. Shang, Phookamsak & L. Cai, Fungal Diversity 95: 168 (2019)

MycoBank No: 555372
Fig. 7

Description

see Phookamsak et al. (2019).

Specimens examined

China, Guizhou Province, Guiyang City, Gaopo Township (26°29'72.37"N, 106°29'59.33"E), on branches of unidentified plant, 30 November 2020. Altitude: 1589 m, S.H. Long, GP01 (GMB0052, KUN-HKAS 112696, living culture GMBC0052).

Additional sequences

GMB0052 (LSU: MW797050, RPB2: MW814896).

Note

The morphological characteristics of this specimen are consistent with those of N. baoshanensis a species described by Phookamsak et al. (2019). Based on phylogenetic and morphological analyses, we consider that this specimen is Neoeutypella baoshanensis. Neoeutypella baoshanensis was described as the type species of Neoeutypella on dead wood of Pinus armandii Franch. from Yunnan Province in China (Phookamsak et al. 2019). This is the first record of N. baoshanensis from Guizhou Province, China.

Figure 7. 

Neoeutypella baoshanensis (GMB0052) A stromata on host substrate B close-up of stromata C transverse section through ascostroma D vertical section through ascostroma E pigments in KOH F culture on PDA G section through the ascostroma H ostiolar canal I, J ascospores K–M asci. Scale bars: 20 μm (G); 10 μm (H–M).

Eutypa Tul. & C. Tul.

Notes

Tulasne & Tulasne (1863) introduced the genus Eutypa with Eutypa lata as the type species. This genus includes several phytopathogens, such as E. lata (Pers.) Tul. & C. Tul. and E. leptoplaca (Durieu & Mont.) Rappaz (Moyo et al. 2017). The morphological characteristics of this genus are black, rounded to irregular-shaped stromata on the host surface, erumpent through host epidermis, solitary to gregarious, entostromatic region, consisting of white pseudoparenchymatous cells and thin black pseudoparenchymatous tissue around the white entostroma, 8-spored, spindle-shaped asci and hyaline, oblong to allantoid ascospores (Rappaz 1987; Moyo et al. 2017). We introduce a new species of Eutypa collected from Guizhou Province in China.

Eutypa cerasi S.H. Long & Q.R. Li, sp. nov.

MycoBank No: 839657
Fig. 8

Holotype

GMB0048.

Etymology

Refers to its host, Prunus cerasus.

Description

Saprobic on decaying branches of Prunus cerasus. Sexual morph: Stromata immersed in bark, covering surface of host, irregular in shape, widely effused, flat, margin diffuse, surface dark brown to black, with punctiform ostioles scattered at surface. Endostroma consists of an outer layer of black, small, dense, thin parenchymal cells and an inner layer of white, large, loose parenchymal cells. Perithecium semi-immersed in stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous 203–304 μm high, 346–477 μm diam. (av. = 408 × 250 μm, n = 10), ovoid, obovoid to oblong. Ostiole opening separately, papillate or apapillate, central. Peridium 30–50 μm thick, dark brown to hyaline with textura angularis cell layers. Asci 83.2–120 × 5.1–8.2 μm (av. = 104.4 × 6.3 μm n = 30) 8–spored clavate, unitunicate, rounded to truncate apex, apical rings inamyloid. Ascospores 7.3–9.9 × 1.4–2 μm (av. = 8.5 × 1.7 μm, n = 30), overlapping, allantoid, slightly curved, subhyaline, smooth, aseptate, usually with oil droplets. Asexual morph: undetermined.

Figure 8. 

Eutypa cerasi (GMB0048, holotype) A stromata on host substrate B, C close-up of stroma D transverse section through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H peridium I–K ascospores L–N asci. Scale bars: 20 μm (G); 10 μm (H–N).

Culture characteristics

Ascospores germinating on PDA within 24 hours. Colonies on PDA, white when young, became pale yellow, irregular in shape, medium dense, flat or effuse, white from above, reverse white at margin, pale yellow at centre, no pigmentation produced on PDA medium, no conidia observed on PDA or on OA media.

Specimens examined

China, Guizhou Province, Guiyang City, Aha Lake National Wetland Park (26°32'50.21"N, 106°40'15.78"E), on branches of Prunus cerasus, 12 August 2020. Altitude: 1089 m, S.H. Long, AH4 (GMB0048, holotype, KUN-HKAS 112685, isotype, ex-type living culture GMBC0048).

Additional specimens examined

China, Guizhou Province, Guiyang City, Aha Lake National Wetland Park (26°32'47.79"N, 106°40'21.09"E), on branches of Cerasus sp., 12 August 2020. Altitude: 1089 m, S.H. Long, AH40 (GMB0049, KUN-HKAS 112683, living culture GMBC0049).

Additional sequences

GMB0048 (LSU: MW797048, RPB2: MW814894); GMB0049 (LSU: MW797049, RPB2: MW814895).

Notes

Eutypa lata is an important pathogen that has a wide range of hosts. However, the classification of E. lata is confusing because there are many variants in previous studies; now all are classified as E. lata (Index Fungorum 2020). Morphologically, the new collection GMB0048 has similar stromata with Eutypa lata, but the ascomata of the new collection are smaller than the ascomata (400 μm diam.) of the original description of E. lata (Tulasne & Tulasne, 1863). The ascomata and asci of the new collection are smaller than the ascomata (400–600 μm diam.) and asci (110–180 × 5–7 μm) of the description of E. lata (Rappaz 1987). Additionally, in the phylogenetic analyses, E. cerasi is located on a branch that forms a sister clade with EP18 and RGA01 and CBS 290.87 basal to E. cerasi. Therefore, combining phylogenetic and morphological analyses, we introduce Eutypa cerasi as a new species of Eutypa.

Paraeutypella L.S. Dissan., J.C. Kang, Wijayaw. & K.D. Hyde.

Notes

Paraeutypella was introduced by Dissanayake et al. (2021) to accommodate Paraeutypella guizhouensis and the genus currently comprises three species. The genus is characterised by poorly developed stromata erumpent through the bark, grouped and irregularly shaped, sometimes confluent, dark brown to black, spindle-shaped, 8-spored asci and allantoid, overlapping, subhyaline ascospores (Trouillas et al. 2011; de Almeida et al. 2016; Dissanayake et al. 2021). In this study, we illustrate Paraeutypella citricola collected from Guizhou Province in China.

Paraeutypella citricola (Speg.). L.S. Dissan., Wijayaw., J.C. Kang & K.D. Hyde, in Dissanayake, Wijayawardene, Dayarathne, Samarakoon & Dai, Biodiversity Data Journal 9: e63864, 14 (2021)

MycoBank No: 228646
Fig. 9

Eutypella citricola Speg., Anal. Mus. nac. Hist. nat. B. Aires 6: 245 (1898)

Description

For description, see Dissanayake et al. (2021)

Specimens examined

China, Guizhou Province, Guiyang City: Aha Lake National Wetland Park (26°20'37.28"N, 108°21'4.34"E), on branches of unidentified plant, 30 August 2020. Altitude: 802 m, S.H. Long, LGS147 (GMB0053, KUN-HKAS 112704, living culture GMBC0053).

Additional sequences

GMB0053 (LSU: 797053, RPB2: MW814898).

Notes

The ITS sequence data were compared by using NCBI and the result showed that it is 100% similar to the ex-type strain (HVVIT07) of P. citricola. The morphological features of the new collection are consistent with those described by Dissanayake et al. (2021). This collection is identified as a P. citricolca, based on morphological and molecular data.

Figure 9. 

Paraeutypella citricola (GMB0053) A stromata on host substrate B, C stromata on host D transverse section through ascostroma E vertical section through ascostroma F culture on PDA G section through the ascostroma H ostiolar canal I peridium J–K ascospores L–O asci. Scale bars: 40 μm (G); 10 μm (H–O).

Discussion

In this study, one new genus, three new species, two new records from China, a novel combination and two known species were reported from karst areas of China. We used molecular data to delimit the species of Diatrypaceae. The new genus Pseudodiatrype is morphologically similar to Allodiatrype and Diatrype, but distinct in the size of stromata, number of ascomata and colour of endostroma; it also formed a distinct branch in the phylogenetic analyses (Fig. 1). Diatrype oregonensis was transferred to Diatrypella oregonensis based on the phylogenetic analyses. Based on phylogenetic analyses, Diatrypella pseudooregonensis was introduced as an 8-spored species of Diatrypella.

Our phylogenetic analyses, based on ITS and β-tubulin, agree with the previous studies (Acero et al. 2004; Trouillas et al. 2011; Mehrabi et al. 2015, 2016; de Almeida et al. 2016; Shang et al. 2017; Dissanayake et al. 2021; Zhu et al. 2021). However, several genera are not monophyletic;for example, Cryptosphaeria, Diatrype, Diatrypella, and Eutypa. The identification of species of Diatrypaceae has been a problem due to the polyphyletic generic concepts based on the features of the stromata in early research (Fries 1823). Recently, new approaches have been proposed for classifying Diatrypaceae. Acero et al. (2004) proposed to classify them by ITS sequence-based phylogenetic analyses, while Carmarán et al. (2006) suggested that the identification should be based on the morphology of the asci. However, due to the lack of type specimens, the lack of β-tubulin sequence and polyphyletic origins have resulted in molecular data that correlate poorly with morphological criteria used to delineate genera and species within the Diatrypaceae (Acero et al. 2004). Moreover, Acero et al. (2004) has mentioned that Diatrypella quercina should be placed in the genus Diatrype despite its polysporous asci since the molecular data placed Diatrypella quercina in the branch of the genus Diatrype.

Diatrype and Diatrypella have morphologically similar verruculose stromata and allantoid ascospores and the polysporous or 8-spored ascus serve as a basis for distinguishing the two genera. However, in phylogenetic analyses, species of these two genera overlap. In this study, we used the phylogenetic analyses as the main basis for classification following Vasilyeva and Stephenson (2005) and Liu et al. (2015). Clade 1 contains Diatrypella verruciformis which is the type species of Diatrypella, of which Diatrypella pseudooregonensis, Diatrypella oregonensis have 8-spored, and other species in clade 1 have polyspored ascus. Clade 12 contains the Diatrype type species Diatrype disciformis, of which Diatrype iranensis and Diatrype macrospora have polyspored ascus, and other species in clade 12 have 8-spored ascus. Hence, we concluded that the number of ascospores in each ascus cannot be used as a criterion for distinguishing Diatrypella from Diatrype.

The phylogenetic tree shows that the classification of Diatrypaceae is confusing. Members of Diatrypella (D. favacea, D. hubeiensis, D. pulvinata and D. yunnanensis) cluster with Diatrype palmicola and Diatrype lancangensis.Maybe this clade should be identified as a new genus. We will discuss its classification status after more strains, more gene sequences and new taxonomic features are collected. Some species of Diatrypella (D. iranensis and D. macrospora) which have polysporous ascus are placed between species of Diatrype, and they are transferred to Diatrype iranensis and Diatrype macrospora by Zhu et al. (Zhu et al. 2021). Diatrype enteroxantha is often derived from the sister clade of Allodiatrype rather than the Diatrype clade. Additionally, Eutypa microasca (BAFC51550) clusters with Peroneutypa species (Clade 17). The above-mentioned confusion also showed in the original publication and other recent studies (Grassi et al. 2014; Mehrabi et al. 2016; Shang et al. 2018; Hyde et al. 2019; Phookamsak et al. 2019; Konta et al. 2020). Therefore, addressing the taxonomic confusion of this family requires a re-examination of older taxa, based on morphological studies, epitypification and multi-gene phylogenetic analyses (Ariyawansa et al. 2014).

Acknowledgements

This research was supported by the National Natural Science Foundation of China (32000009 and 31960005); the Fund of the Science and Technology Foundation of Guizhou Province ([2020]1Y059); Guiyang Science and Technology Planning Project No. (2017)30-19; Guizhou Province Ordinary Colleges and Universities Youth Science and Technology Talent Growth Project [2021]154. Nalin N. Wijayawardene would like to thank the National Natural Science Foundation of China (No. NSFC 31950410558), the State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medicial University (No. FAMP201906K) and High-Level Talent Recruitment Plan of Yunnan Provinces (“Young Talents” Program and “High-End Foreign Experts” Program); the Fund of High-Level Innovation Talents [No. 2015-4029], the Base of International Scientific and Technological Cooperation of Guizhou Province [No. [2017]5802].

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