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New contributions to Diatrypaceae
from karst areas in China
Sihan Long1,2, Lili Liu3, Yinhui Pi1, Youpeng Wu1, Yan Lin1, Xu Zhang1,
Qingde Long1, Yingqian Kang4, Jichuan Kang5, Nalin N. Wijayawardene1,6,
Feng Wang7, Xiangchun Shen1,2, Qirui Li1,2
1State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medical University, Gui-
yang 550004, China 2e High Ecacy Application of Natural Medicinal Resources Engineering Center of
Guizhou Province (e Key Laboratory of Optimal Utilization of Natural Medicine Resources), School of Phar-
maceutical Sciences, Guizhou Medical University, University Town, Guian New District, Guizhou 550025,
China 3 Immune Cells and Antibody Engineering Research Center of Guizhou Province/ Key Laboratory of
Biology and Medical Engineering, Guizhou Medical University, Guiyang 550004, China 4Key Laboratory
of Environmental Pollution Monitoring and Disease Control, Ministry of Education of Guizhou and Guizhou
Talent Base for Microbiology and Human Health, School of Basic Medical Sciences, Guizhou Medical Universi-
ty, Guiyang, China 5Engineering and Research Center for Southwest Bio-Pharmaceutical Resources of National
Education Ministry of China, Guizhou University, Guiyang, Guizhou 550025, China 6Center for Yunnan
Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering,
Qujing Normal University, Qujing, Yunnan 655011, China 7Guizhou Provincial Academician Workstation
of Microbiology and Health, Guizhou Academy of Tobacco Science, Guiyang, Guizhou, 550000, China
Corresponding author: Qirui Li (lqrnd2008@163.com)
Academic editor: Andrew Miller|Received 20 May 2021|Accepted 20 July 2021|Published 20 August 2021
Citation: Long S–H, Liu L–L, Pi Y–H, Wu Y–P, Lin Y, Zhang X, Long Q–D, Kang Y–Q, Kang J–C, Wijayawardene
NN, Wang F, Shen X–C, Li Q–R (2021) New contributions to Diatrypaceae from karst areas in China. MycoKeys 83:
1–37. https://doi.org/10.3897/mycokeys.83.68926
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 conrmed that these new collections
comprise a new genus Pseudodiatrype, three new species (Diatrype lancangensis, Diatrypella pseudoore-
gonensis 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). e new taxa are introduced, based on multi-gene phylogenetic analyses (ITS,
β-tubulin), as well as morphological analyses. e 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
MycoKeys 83: 1–37 (2021)
doi: 10.3897/mycokeys.83.68926
https://mycokeys.pensoft.net
Copyright Sihan Long et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
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RESEARCH ARTICLE
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Sihan Long et al. / MycoKeys 83: 1–37 (2021)
2
outer and inner layers of large white cells with thin, powdery, yellowish cells. ese characteristics separate
this genus from two similar genera Allodiatrype and Diatrype. Based on morphological as well as phyloge-
netic analyses, Diatrype lancangensis is introduced as a new species of Diatrype. e 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. e polyphyletic nature of some genera of Diatrypaceae has
led to confusion in the classication of the family, thus we discuss whether the number of ascospores per
asci can still be used as a basis for classication.
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) re-
vised 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.).
is was followed by Wijayawardene et al. (2020) in which 20 genera were accepted
into Diatrypaceae. e Diatrypaceae is characterised by perithecial ascomata embed-
ded 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 dec-
ades, 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 grape-
vine, 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).
irteen 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. e landform
of karst can be found in almost all Provinces of China, with the most extensive distri-
bution in Guizhou and Yunnan Provinces (Miao et al. 2007). Karst virgin forest is a
relatively stable ecosystem with rich biological resources, highly primitive and main-
taining stable biological diversity (Dong et al. 2002). e special karst and ecological
environment is home to a rich diversity of diatrypaceous fungi.
New contributions to Diatrypaceae 3
In this study, we revisit species of Diatrypaceae collected from karst areas in
Guizhou, Hainan and Yunnan Provinces of China. Based on morpho-molecular analy-
ses, one new genus and three new species are introduced; in addition, a new combina-
tion 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. e
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 purication. e specimens were deposited at the Herbarium of Cryptogams, Kun-
ming 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).
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 Don-
aldson 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 proles 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 nal extension at 72 °C for 10 min-
utes. PCR prole 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,
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
4
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. polysporaTMFLUCC 17-0364 MF959500 MG334556 Senwanna et al. (2017)
A. rabenhorstii WA08CB HQ692619 HQ692523 Trouillas et al. (2011)
Allodiatrype arengaeTMFLUCC 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 is study
Anthostoma decipiensTIPV-FW349 AM399021 AM920693 Unpublished.
A. decipiensTJL567 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. disciformisTGNA14 KR605644.1 KY352434.1 Senanayake et al. (2015)
D. disciformisTD21C, 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 is study
D. lancangensis GMB0046 MW797114 MW814886 is study
D. lancangensis GMB0047 MW797116 MW814887 is 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 is study
D. pseudooregonensis GMB0040 MW797117 MW814889 is study
D. pseudooregonensis GMB0041 MW797118 MW814890 is study
D. pseudooregonensis GMB0042 MW797119 MW814891 is study
D. pseudooregonensis GMB0043 MW797120 MW814892 is study
D. pseudooregonensis GMB0044 MW797110 MW814882 is 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. verruciformisTUCROK1467 JX144793 JX174093 Lynch et al. (2013)
New contributions to Diatrypaceae 5
Taxa Strain number GenBank Accession number Reference
ITS β-tubulin
D. verruciformisTUCROK754 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 is 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 is study
E. cerasi GMB0049 MW797105 MW814877 is study
E. avovirens E48C, CBS 272.87 AJ302457 DQ006959 Rolshausen et al. (2006)
E. laevata E40C CBS 291.87 AJ302449 NA Acero et al. (2004)
E. lataTCBS290.87 HM164736 HM164770 Trouillas and Gubler (2010)
E. lataTEP18 HQ692611 HQ692501 Trouillas et al. (2011)
E. lataTRGA01 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. cerviculataTM68 JF340269 NA Arhipova et al. (2012)
E. cerviculataTEL59C 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. salinicolaTMFLUCC 15-1277 KX573915 KX573932 Dayarathne et al. (2016)
Kretzschmaria deusta CBS 826.72 KU683767 KU684190 U’Ren et al. (2016)
Monosporascus cannonballusTCMM3646 JX971617 NA Unpublished
M. cannonballusTATCC 26931 FJ430598 NA Unpublished
Neoeutypella baoshanensisTGMB0052 MW797106 MW814878 is study
N. baoshanensisTLC 12111 MH822887 MH822888 Hyde et al. (2019)
N. baoshanensisTEL51C, CBS 274.87 AJ302460 NA Acero et al. (2004)
N. baoshanensisTMFLUCC 16-1002 MT310662 NA Phukhamsakda et al. (2020)
N. baoshanensisTGL08362 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 is study
Pa. guizhouensisTKUMCC 20-0016 MW039349 MW239660 Dissanayake et al. (2021)
Pa. guizhouensisTKUMCC 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 rhizophoraeTBCC44877 KJ888853 NA Klaysuban et al. (2014)
Pe. rhizophoraeTBCC44878 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 hainanensisTGMB0054 MW797111 MW814883 is study
Ps. hainanensisTGMB0055 MW797112 MW814884 is 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)
T: Types species of the genus; NA: No sequence is available in GenBank; Newly generated sequences are indicated in bold.
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
6
elongation at 72 °C for 1.5 minutes and a nal extension at 72 °C for 10 minutes
(Konta et al. 2020). PCR prole 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 nal extension at 72°C
for 10 minutes (de Almeida et al. 2016). PCR products were submitted to Sangon
Biotech, Shanghai, China for purication 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 pub-
lished papers (Mehrabi et al. 2019; Dayarathne et al. 2020; Konta et al. 2020; Dissa-
nayake et al. 2021; Zhu et al. 2021). After the preliminary identication results of the
sequences, multiple sequence alignments (ITS and β-tubulin) were aligned using Bi-
oEdit v. 7.0 (Hall 1999). Alignments were converted from FASTA to PHYLIP format
by using Alignment Transformation Environment online (https://sing.ei.uvigo.es/AL-
TER/, Glez-Peña et al. 2010). Maximum Likelihood (ML) analyses and Bayesian pos-
terior 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). e Boot-
strap 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 us-
ing Adobe Photoshop CS6 software. e nal 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 dier signicantly. e 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). e RAxML analyses
resulted in a best scoring likelihood tree selected with a nal ML optimisation likeli-
hood value of -15731.506304, which is shown in Fig. 1.
e 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:
New contributions to Diatrypaceae 7
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. e 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.
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
8
Figure 1. Continued.
Clade 1: Diatrypella pseudooregonensis and Diatrypella oregonensis clustered with the
species of Diatrypella in Clade 1 with high bootstrap support, Diatrypella pseu-
dooregonensis 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 classication which is dicult to distinguish, based on phylogenetic
aspects alone. erefore, we introduce Diatrype lancangensis as a new species of
Diatrype, based on phylogenetic analyses and morphological dierences (Table 2).
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.
New contributions to Diatrypaceae 9
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 unidentied plant. Sexual
morph: Stromata immersed in bark, aggregated, irregular in shape, widely eused,
at, margin diuse, 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 283.5–343.5µm high,
207–290 µm broad (av. = 315.5 × 248.0 µm, n = 10), ovoid, obovoid to oblong, mon-
ostichous, aterrimus. Ostiole opening separately, papillate or apapillate, central. Peridi-
um 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.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies 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. C, Yunnan Province, Baoshan City, Lancang River
Nature Reserve (25°1'17.44"N, 99°35'10.05"E) on branches of an unidentied 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. C, Yunnan Province, Baoshan City, Lan-
cang River Nature Reserve (25°1'17.44"N, 99°35'10.05"E) on branches of an uniden-
tied 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 unidentied plant, 5 October 2019. Altitude: 2623m., 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 ve Diatrypella and one Diatrype species (Fig. 1), this clade is consistent with
the study of Konta et al. (2020). e taxonomic confusion of Diatrypaceae has led to
diculties 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 undu-
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
10
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
Gostiolar canal H peridium I–K ascospores L–N asci. Scale bars: 10 µm (G–N).
New contributions to Diatrypaceae 11
lata (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 dened as a new
genus, but it is dicult to nd 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. e characteristics of
the stromata of Diatrypella spp. in clade 7 are solitary and scattered, which is distinctly
dierent from widely euse, at 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 specicity. 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 phylo-
genetically distinct.
Type species. Pseudodiatrype hainanensis S. H. Long & Q.R. Li sp. nov.
Description. Saprobic on decaying branches of an unidentied 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. Peri-
thecium 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, api-
cally rounded, apical rings inamyloid. Ascospores irregularly arranged, allantoid, slightly
or moderately curved, smooth, subhyaline, aseptate, usually with two oil droplets.
Asexual morph: undetermined.
Note. e genus Pseudodiatrype is introduced to accommodate the new collec-
tion made from Hainan Province of China and typied 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
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
12
distributed over large areas, sometimes covering the surface of the host (Vasilyeva and
Ma 2014; Konta et al. 2020). Pseudodiatrype diers 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 dierent from the
endostroma of Pseudodiatrype having black outer and inner cells surrounded by pow-
dery, pale yellow cells. In addition, the sizes of stroma and ascospores are dierent 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 with-
in 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 unidentied 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 gre-
garious. 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 parenchy-
mal cells with powdery, thin, yellowish tissue. Ostiole opening separately, papillate or
apapillate, central. Perithecium immersed in the stroma, globose to subglobose, gla-
brous, with cylindrical neck, brevicollous or longicollous, 193–347 µm high, 138–
206µm 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.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies on PDA, white when young, became pale brown, dense, but thinning towards
edge, uy to slightly uy, white from above, pale brown from below, no pigmenta-
tion produced on PDA medium, no conidia observed on PDAor on OA media.
Specimens examined. C, Hainan Province, Wuzhishan City, Wuzhishan Na-
ture Reserve (18°54'21.81"N, 109°40'54.12"E) on branches of unidentied 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. C, Hainan Province, Wuzhishan City,
Wuzhishan Nature Reserve (18°54'21.81"N, 109°40'54.12"E) on branches of an uni-
New contributions to Diatrypaceae 13
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:
40µm (G); 10 µm (H–P).
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
14
dentied 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 endos-
troma. ere 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. e genus Diatrypella was introduced by Cesati & De Notaris (1863) and was
typied with Diatrypella verruciformis (Ehrh.) Nitschke. is genus was characterized
by pustule-like stromata erumpent through the host surface, polysporous asci and al-
lantoid 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 combina-
tion 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 unidentied 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,
1–3mm 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 cy-
lindrical neck, brevicollous or longicollous 218.5–465 µm high, 112–257 µm diam.
(av. = 306×164µm, 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.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies 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
New contributions to Diatrypaceae 15
Figure 4. Diatrypella pseudooregonensis (GMB0041, holotype) A stromata on host substrate B, Cstro-
mata on host substrate D transverse section through ascostroma E vertical section through ascostroma
Fculture 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).
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
16
below, no pigmentation produced on PDA medium, no conidia observed on PDA or
on OA media.
Specimens examined. C, Yunnan Province, Baoshan City, Lancang River
Nature Reserve (25°1'19.88"N, 99°35'30.68"E) on branches of an unidentied 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. C, Yunnan Province, Baoshan City, Lan-
cang River Nature Reserve (25°1'13.51"N, 99°35'25.59"E) on branches of an uni-
dentied plant, 6 October 2019. Altitude: 2630 m, Y.H. Pi & Qiong Zhang, LC384
(GMB0043, KUN-HKAS 112681, living culture GMBC0043); C, Yunnan Prov-
ince, Baoshan City, Lancang River Nature Reserve (25°1'15.00"N, 99°35'39.73"E) on
branches of an unidentied plant, 5 October 2019. Altitude: 2698 m, Y.H. Pi & Qiong
Zhang, LC312 (GMB0040, KUN-HKAS 112674, living culture GMBC0040); C-
, Yunnan Province, Baoshan City, Lancang River Nature Reserve (25°35'19.09"N,
99°35'19.09"E) on branches of an unidentied plant, 5 October 2019. Altitude:
2569m, Y.H. Pi & Qiong Zhang, LC193 (GMB0039, KUN-HKAS 112667, living
culture GMBC0039); C, Yunnan Province, Baoshan City, Lancang River Nature
Reserve (25°1'9.11"N, 99°35'24.80"E) on branches of an unidentied plant, 5 Oc-
tober 2019. Altitude: 2649 m, Y.H. Pi & Qiong Zhang, LC335 (GMB0042, KUN-
HKAS 112647, living culture GMBC0042); C, Guizhou Province, Anshun City,
Pingba District (26°25'9.65"N, 106°24'24.48"E) on branches of an unidentied 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 Dia-
trypella has more than eight ascospores in each ascus (Senanayake et al. 2015). How-
ever, 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 classication.
Diatrypella vulgaris Trouillas, W.M. Pitt & Gubler, Fungal Diversity 49: 212
(2011)
MycoBank No: 519404
Fig. 5
Description. Saprobic on decaying branches of an unidentied plant. Sexual morph:
Stromata scattered on the host, 0.8–1.5 mm long and 0.8–2 mm broad (av. = 1.2 ×
New contributions to Diatrypaceae 17
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).
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
18
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 parenchy-
mal 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. As-
cospores 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.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies on PDA, white when young, became pale brown, dense, but thinning towards
edge, medium dense, white from above, reverse side white at margin, esh to pale
brown at centre, no pigmentation produced on PDA medium, no conidia observed on
PDA or on OA media.
Specimens examined. C, Guizhou Province, Guiyang City, Gaopo Town-
ship (26°29'72.02"N, 106°29'55.57"E), on branches of unidentied plant, 30 Octo-
ber 2020. Altitude: 1589 m, S.H. Long, GP02 (GMB0051, KUN-HKAS 112697,
living culture GMBC0051).
Additional sequences. GMB0051 (LSU: MW797051, RPB2: MW814897).
Note. e comparison of ITS sequences in NCBI showed that this isolate is 100%
similar to the strain of Diatrypella vulgaris (HVGRF03), isolated from holotype speci-
mens. Morphologically, GMB0051 shows the same features as Diatrypella vulgaris. e
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 identica-
tion tool. Diatrypella vulgaris has been reported in Austria and ailand (Trouillas et
al. 2011, Hyde et al. 2017). is is the rst 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. e strains of Diatrype oregonensis (DPL200, CA117) generated from
Trouillas et al. (2010) grouped in Diatrypella s. str. Diatrype oregonensis was erected
New contributions to Diatrypaceae 19
in 1930 as Eutypella oregonensis (Kauman 1930). No available sequences from type
material were found. After re-examination of holotype specimen of Diatrype oregonen-
sis, 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 simi-
lar 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. e genus Allodiatrype was introduced by Konta et al. (2020), which was char-
acterised 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 unidentied plant. Sexual morph: Stro-
mata wart-like, pustulate, 0.5–1.8 mm long and 0.8–2.2 mm broad (av. = 1.2×1.3mm,
n = 30), about 1 mm thick, 1–18 in a single stroma, visible as black, rounded to irregu-
lar in shape on the host surface, erumpent through host bark, solitary to gregarious. En-
dostroma 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 inated, apically rounded
to truncate, apical rings inamyloid. Ascospores 6–11 × 2–2.5µm (av. = 8.9× 2.3 µm,
n = 30), irregularly arranged, allantoid, slightly curved, smooth, subhyaline, aseptate,
usually with two oil droplets. Asexual morph: undetermined.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies on PDA, white when young, became pale yellow, irregular in shape, medium
dense, at or euse, slightly raised, with edge mbriate, uy to fairly uy, white
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
20
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).
New contributions to Diatrypaceae 21
from above, reverse side white at margin, pale brown at centre, no pigmentation pro-
duced on PDA medium, no conidia observed on PDA or on OA media.
Specimens examined. C, Yunnan Province, Baoshan City, Lancang River
Nature Reserve (24°57'25.35"N, 99°44'22.82"E), on branches of unidentied 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. e ITS sequence data were subjected to BLAST in NCBI and the results
showed that it is 100% similar to Allodiatrype thailandica. Additionally, based on mor-
phological and phylogenetic analyses, this strain was identied as the A. thailandica.
e 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–
2.7µm) (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 identication.
A.thailandica has been reported in ailand in 2016 as Diatrype thailandica and rec-
ognised as A. thailandica by Konta et al. (2020). is is the rst report of Allodiatrype
thailandica from China.
Neoeutypella M. Raza, Q.J. Shang, Phookamsak & L. Cai, Fungal Diversity 95:
167 (2019)
Note. e genus Neoeutypella was introduced by Phookamsak et al. (2019) and is char-
acterised 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. C, Guizhou Province, Guiyang City, Gaopo Town-
ship (26°29'72.37"N, 106°29'59.33"E), on branches of unidentied plant, 30 Novem-
ber 2020. Altitude: 1589 m, S.H. Long, GP01 (GMB0052, KUN-HKAS 112696,
living culture GMBC0052).
Additional sequences. GMB0052 (LSU: MW797050, RPB2: MW814896).
Note. e 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 Neoeutypellabaoshanensis.
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
22
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).
New contributions to Diatrypaceae 23
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). is is the rst record of N. baoshanensis from Guizhou Province,China.
Eutypa Tul. & C. Tul.
Notes. Tulasne & Tulasne (1863) introduced the genus Eutypa with Eutypa lata as
the type species. is genus includes several phytopathogens, such as E. lata (Pers.)
Tul. & C. Tul. and E. leptoplaca (Durieu & Mont.) Rappaz (Moyo et al. 2017). e
morphological characteristics of this genus are black, rounded to irregular-shaped stro-
mata 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 eused,
at, margin diuse, 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-im-
mersed in stroma, globose to subglobose, glabrous, with cylindrical neck, brevicollous
203–304 µm high, 346–477 µm diam. (av. = 408×250µm, n = 10), ovoid, obovoid
to oblong. Ostiole opening separately, papillate or apapillate, central. Peridium 30–
50µm thick, dark brown to hyaline with textura angularis cell layers. Asci 83.2–120×
5.1–8.2µm (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 ×
1.7µm, n = 30), overlapping, allantoid, slightly curved, subhyaline, smooth, aseptate,
usually with oil droplets. Asexual morph: undetermined.
Culture characteristics. Ascospores germinating on PDA within 24 hours. Colo-
nies on PDA, white when young, became pale yellow, irregular in shape, medium
dense, at or euse, 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.
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
24
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 sec-
tion through the ascostroma H peridium I–K ascospores L–N asci. Scale bars: 20 µm (G); 10 µm (H–N).
New contributions to Diatrypaceae 25
Specimens examined. C, Guizhou Province, Guiyang City, Aha Lake Na-
tional 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. C, 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 classication of E. lata is confusing because there are many variants
in previous studies; now all are classied 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). e
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. erefore,
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. e genus
is characterised by poorly developed stromata erumpent through the bark, grouped
and irregularly shaped, sometimes conuent, 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 Paraeu-
typella 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)
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
26
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 Gsection through the
ascostroma H ostiolar canal I peridium J–K ascospores L–O asci. Scale bars: 40µm (G); 10µm (H–O).
New contributions to Diatrypaceae 27
Description. For description, see Dissanayake et al. (2021)
Specimens examined. C, Guizhou Province, Guiyang City: Aha Lake Na-
tional Wetland Park (26°20'37.28"N, 108°21'4.34"E), on branches of unidentied
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. e 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. e
morphological features of the new collection are consistent with those described by
Dissanayake et al. (2021). is collection is identied as a P. citricolca, based on mor-
phological and molecular data.
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. e new genus Pseudodiatrype
is morphologically similar to Allodiatrype and Diatrype, but distinct in the size of stro-
mata, 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, Dia-
trypella pseudooregonensis was introduced as an 8-spored species of Diatrypella.
Table 2. e 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 is study
D. lijiangensis 1 diam. 1 diam 50–90 6–9 6–8 1–2 iyagaraja 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 is study
Newly identied taxa are indicated in bold, NA: No description available.
Sihan Long et al. / MycoKeys 83: 1–37 (2021)
28
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. e identication 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 identication
should be based on the morphology of the asci. However, due to the lack of type speci-
mens, the lack of β-tubulin sequence and polyphyletic origins have resulted in molecu-
lar 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 dis-
tinguishing 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
classication 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 as-
cus, 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.
e phylogenetic tree shows that the classication 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
identied as a new genus. We will discuss its classication 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 spe-
cies 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). e 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). erefore, addressing the taxonomic confusion of this family
requires a re-examination of older taxa, based on morphological studies, epitypication
and multi-gene phylogenetic analyses (Ariyawansa et al. 2014).
New contributions to Diatrypaceae 29
Acknowledgements
is 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 Pro-
ject 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 For-
eign Experts” Program); the Fund of High-Level Innovation Talents [No. 2015-4029],
the Base of International Scientic and Technological Cooperation of Guizhou Prov-
ince [No. [2017]5802].
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