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The Hidden Diversity of Diatrypaceous Fungi in China

Frontiers in Microbiology

May 2021
12:646262

DOI:10.3389/fmicb.2021.646262

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CC BY 4.0

Authors:

Hai-Yan Zhu

Institute of Microbiology, Chinese Academy of Sciences

Meng Pan

Peking University

Nalin Wijayawardene

Center for Yunnan Plateau Biological Resources Protection and Utilization, Qujing Normal University, Qujing City, Yunnan Province, P.R. China.

Ning Jiang

Chinese Academy of Forestry

Show all 8 authorsHide

In this study, we investigated the diversity of diatrypaceous fungi from six regions in China based on morpho-molecular analyses of combined ITS and tub2 gene regions. We accept 23 genera in Diatrypaceae with 18 genera involved in the phylogram, and the other five genera are lacking living materials with sequences data. Eleven species included in four genera (viz. Allocryptovalsa, Diatrype, Diatrypella, and Eutypella) have been isolated from seven host species, of which nine novel species (viz. Allocryptovalsa castaneae, A. castaneicola, Diatrype betulae, D. castaneicola, D. quercicola, Diatrypella betulae, Da. betulicola, Da. hubeiensis, and Da. shennongensis), a known species of Diatrypella favacea, and a new record of Eutypella citricola from the host genus Morus are included. Current results show the high diversity of Diatrypaceae which are wood-inhabiting fungi in China.

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| Phylogram of Diatrypaceae based on combined ITS and tub2 sequence data. The MP and ML bootstrap support values above 70% are shown at the first and second positions, respectively. Thickened branches represent posterior probabilities above 0.95 from the BI. Ex-type strains are in bold, type species are denoted with the superscript "TS" and the disputable type species are denoted with the superscript "TSQ." Strains from the current study are in blue.

…

| Holomorph of Allocryptovalsa castaneae (BJFU CF2020518, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of ascoma. (D) Longitudinal section through ascoma. (E) Conidiomata on the host. (F) Conidioma on the host. (G) Asci and ascospores. (H) Ascospores. (I,J) Conidia attatch to conidiogenous cells. (K) Conidia. Scale bars: (A,F) = 1 mm; (B−E) = 500 µm; (G−K) = 10 µm.

…

| Sexual morph of Allocryptovalsa castaneicola (BJFU CF2020519, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Longitudinal section through ascoma. (D) Transverse section of ascoma. (E,F) Ascus and ascospores. (G) Ascospores. Scale bars: (A) = 1 mm; (B-D) = 500 µm; (E-G) = 10 µm.

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| Asexual morph of Diatrype betulae (BJFU CF2020510, holotype). (A) Conidiomata on the host. (B) Conidioma on the host. (C,D) Transverse section of conidioma. (E) Longitudinal section through conidioma. (F) Conidiogenous cells. (G,H) Conidia. Scale bars: (A) = 1 mm; (B−E) = 500 µm; (F−H) = 10 µm.

…

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fmicb-12-646262 May 25, 2021 Time: 18:38 # 1

ORIGINAL RESEARCH

published: 31 May 2021

doi: 10.3389/fmicb.2021.646262

Edited by:

Quan Lu,

Chinese Academy of Forestry, China

Reviewed by:

Saowaluck Tibpromma,

Chinese Academy of Sciences, China

K. W. Thilini Chethana,

Mae Fah Luang University, Thailand

*Correspondence:

Xinlei Fan

xinleifan@bjfu.edu.cn

†These authors have contributed

equally to this work and share ﬁrst

authorship

Specialty section:

This article was submitted to

Microbe and Virus Interactions with

Plants,

a section of the journal

Frontiers in Microbiology

Received: 25 December 2020

Accepted: 09 March 2021

Published: 31 May 2021

Citation:

Zhu H, Pan M,

Wijayawardene NN, Jiang N, Ma R,

Dai D, Tian C and Fan X (2021) The

Hidden Diversity of Diatrypaceous

Fungi in China.

Front. Microbiol. 12:646262.

doi: 10.3389/fmicb.2021.646262

The Hidden Diversity of

Diatrypaceous Fungi in China

Haiyan Zhu1†, Meng Pan1†, Nalin N. Wijayawardene2, Ning Jiang1, Rong Ma3,

Dongqin Dai2, Chengming Tian1and Xinlei Fan1*

1The Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing, China,

2Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food

Engineering, Qujing Normal University, Yunnan, China, 3College of Forestry and Horticulture, Xinjiang Agricultural University,

Ürümqi, China

In this study, we investigated the diversity of diatrypaceous fungi from six regions in

China based on morpho-molecular analyses of combined ITS and tub2 gene regions.

We accept 23 genera in Diatrypaceae with 18 genera involved in the phylogram, and

the other ﬁve genera are lacking living materials with sequences data. Eleven species

included in four genera (viz.Allocryptovalsa,Diatrype,Diatrypella, and Eutypella) have

been isolated from seven host species, of which nine novel species (viz.Allocryptovalsa

castaneae,A. castaneicola,Diatrype betulae,D. castaneicola,D. quercicola,Diatrypella

betulae,Da. betulicola,Da. hubeiensis, and Da. shennongensis), a known species of

Diatrypella favacea, and a new record of Eutypella citricola from the host genus Morus

are included. Current results show the high diversity of Diatrypaceae which are wood-

inhabiting fungi in China.

Keywords: Allocryptovalsa,Diatrype,Diatrypella,Eutypella, fungal diversity, phylogeny, taxonomy

INTRODUCTION

Diatrypaceae is an important family in Xylariales (Sordariomycetes, Ascomycota), containing many

taxa with a worldwide distribution (Glawe and Jacobs, 1987;Mayorquin et al., 2016;Senwanna

et al., 2017;Moyo et al., 2018a,b;Konta et al., 2020;Wijayawardene et al., 2020). Species of the

Diatrypaceae are frequently saprobic on the decaying wood of angiosperms (Tendulkar, 1970;

Acero et al., 2004;de Almeida et al., 2016). However, an endophyte of Picea abies, a gymnosperm

host, was identiﬁed as an asexual morph of Diatrypaceae (Libertella sp.) by Caroll et al. (1977). Few

endophytes such as Diatrypella frostii and Peroneutypa scoparia were reported later (de Errasti et al.,

2010;Vieira et al., 2011). Strikingly, several plant pathogens of Diatrypaceae were reported causing

canker, dieback, and grapevine trunk diseases, e.g., Cryptosphaeria populina was linked to canker

in Populus species (Glawe and Rogers, 1984); Cryptosphaeria pullmanensis caused canker disease in

Populus alba and Salix alba (Ma et al., 2016); Cryptovalsa ampelina caused grapevine trunk disease

on Vitis species (Luque et al., 2006); Eutypa lata was isolated from Prunus armeniaca and Vitis

species with canker and dieback symptoms (Lardner et al., 2005); Eutypa leptoplaca contributed to

the dieback of grapevines (Trouillas and Gubler, 2004;Catal et al., 2007); and Eutypella parasitica

caused canker in Acer species (Rappaz, 1987).

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Zhu et al. Diatrypaceous Fungi in China

Nitschke (1867) proposed the ﬁrst important study

of Diatrypaceae (as Diatrypeae) including ﬁve genera

(Supplementary Table 1). Kirk et al. (2001) accepted nine

genera. Later, the family was considered to accommodate

13 genera by Kirk et al. (2008) (Supplementary Table 1).

Diatrypasimilis,Monosporascus, and Pedumispora have been

added to the family in subsequent studies (Abdel-Wahab et al.,

2014;Klaysuban et al., 2014;Maharachchikumbura et al.,

2015). Wijayawardene et al. (2018) reported 17 genera in the

family (Supplementary Table 1). Subsequently, Dayarathne

et al. (2016),Senwanna et al. (2017),Phookamsak et al. (2019)

respectively introduced Allocryptovalsa, Halodiatrype, and

Neoeutypella, and accepted them as members of Diatrypaceae.

Wijayawardene et al. (2020) listed 20 genera in the family. Later,

Dayarathne et al. (2020a,b),Konta et al. (2020) respectively

added Allodiatrype,Halocryptosphaeria, and Halocryptovalsa to

this family.

The sexual morph members of Diatrypaceae are characterized

by perithecial ascomata usually with ostiolar necks, 8-spored

or polysporous asci with a very long pedicel and J-/J+apical

apparatus, and allantoid ascospores (Senanayake et al., 2015;

Dayarathne et al., 2016;de Almeida et al., 2016). Several

asexual genera included in coelomycetes or hyphomycetes

(viz.Cytosporina,Libertella, and Phaeoisaria) have been

linked to the family Diatrypaceae (Dayarathne et al., 2016;

de Almeida et al., 2016;Mehrabi et al., 2016;Shang et al.,

2017). However, the asexual morphs of many species were

still indistinguishable (Acero et al., 2004). Senanayake

et al. (2015) summarized that the asexual morph of this

family had acervular and astromatic conidiomata, branched

conidiophores and ﬁliform, allantoid or rarely straight

conidia with ﬂattened base and blunt apex. However, in

most cases, it is diﬃcult to diﬀerentiate diatrypaceous

species based on asexual morphs (Glawe and Rogers, 1986;

de Almeida et al., 2016).

Due to overlapping phenotypic characters in Diatrypaceae,

polyphasic approaches to solve the taxonomy of fungi were

very common in recent studies (Wijayawardene et al., 2016;

Norphanphoun et al., 2017;Fan et al., 2018, 2020;Lawrence et al.,

2018;Zhu et al., 2020). The ﬁrst molecular phylogenetic analysis

of Diatrypaceae based on ITS showed that Cryptosphaeria,

Diatrype,Diatrypella,Eutypa, and Eutypella were polyphyletic

(Acero et al., 2004). Recently, the identiﬁcation and classiﬁcation

of diatrypaceous taxa were performed by the multiple sequence

data (mostly ITS and tub2) and morphological characters

(Phookamsak et al., 2019;Dayarathne et al., 2020a;Hyde et al.,

2020b;Konta et al., 2020). Moreover, Konta et al. (2020)

introduced one new genus Allodiatrype and ﬁve new species

belonging to Allocryptovalsa,Allodiatrype, and Diatrypella from

palms (Arecaceae) based on this criterion.

During the investigation of forest pathogens in China,

86 diatrypaceous specimens associated with various disease

symptoms were collected from Beijing City, Xinjiang Uygur

Autonomous Region, and four other provinces in China viz.

Hubei, Hebei, Jiangsu, and Yunnan. The objectives were to

supplement a multi-gene DNA dataset of Diatrypaceae including

ITS and tub2, improve the phylogenetic systematics of this

family, and provide a theoretical basis for the identiﬁcation of

diseases and pathogens.

METHODS

Isolates

Symptomatic branches or twigs were collected from seven tree

hosts (Betula albosinensis,B. davurica,B. platyphylla,Castanea

mollissima,Juglans regia,Morus alba, and Quercus mongolica)

from Beijing City, Xinjiang Uygur Autonomous Region, and

four other provinces in China viz. Hubei, Hebei, Jiangsu, and

Yunnan. Eighty-six fresh specimens of Diatrypaceae were put

into envelopes with records of their altitude, collector, collecting

time, host, longitude, and latitude. A total of 21 representative

isolates were obtained by removing the ascospores or conidial

mass from fresh specimens on the surface of 1.8% potato dextrose

agar (PDA) and incubating at 25◦C for 24 h. Single germinating

spore was transferred onto a fresh PDA plate. Specimens and

isolates were deposited in the Beijing Forestry University (BJFU)

and the Beijing Museum of Natural History (BJM). Strains of

the new species are maintained in the China Forestry Culture

Collection Centre (CFCC).

Morphological Analysis

Species identiﬁcation was based on morphological features of

fruiting bodies and micromorphology supplemented by cultural

characteristics. Macro-morphological observations including

structure and size of stromata, ectostromatic disc, and ostioles

were determined using a Leica stereomicroscope (M205 FA)

(Leica Microsystems, Wetzlar, Germany). Micro-morphological

photographs were captured using a Nikon Eclipse 80i microscope

(Nikon Corporation, Tokyo, Japan), including conidiophores,

asci, and conidia/ascospores. Adobe Bridge CS v. 6 and Adobe

Photoshop CS v. 5 were used for manual editing. At least

10 conidiomata/ascomata, 10 asci, and 30 conidia/ascospores

were randomly selected for measurement to calculate the mean

width/length and respective standard deviations (SD). Cultural

characteristics of strains incubated in the dark at 25◦C were

recorded. Colony morphology was described using the color

charts of Rayner (1970). Nomenclatural novelties were deposited

in the MycoBank (1Crous et al., 2004).

DNA Extraction, PCR Ampliﬁcation, and

Sequencing

Fungal mycelium grown on the cellophane on PDA was scraped

for the extraction of genomic DNA following the modiﬁed CTAB

method (Doyle and Doyle, 1990). Two loci were ampliﬁed,

including the internal transcribed spacer (ITS) region and partial

beta-tubulin (tub2) using the primer pairs ITS1/ITS4 (White

et al., 1990) and T1/Bt2b (Glass and Donaldson, 1995;O’Donnell

and Cigelnik, 1997), respectively. The additional combination of

Bt2a and Bt2b (Glass and Donaldson, 1995) was used in case of

ampliﬁcation failure of the primer T1 and Bt2b. The polymerase

chain reaction (PCR) assay was conducted as described in

1www.mycobank.org

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Zhu et al. Diatrypaceous Fungi in China

Fan et al. (2020). PCR ampliﬁcation products were estimated

via electrophoresis in 2% agarose gels. DNA sequencing was

performed using an ABI PRISMR

3730XL DNA Analyzer with a

BigDye Terminater Kit v. 3.1 (Invitrogen, United States) at the

Shanghai Invitrogen Biological Technology Company Limited

(Beijing, China).

DNA Sequence Analysis

The initial identities of our strains sequenced were obtained

by morphological observations and nucleotide BLAST search.

To clarify the phylogenetic position, the alignment based on a

combined matrix using ITS and tub2 sequences was performed

to compare with other available species in Diatrypaceae.

Reference sequences were selected based on ex-type or ex-

epitype sequences available from relevant recently published

literature (de Almeida et al., 2016;Senwanna et al., 2017;

Shang et al., 2017, 2018;Hyde et al., 2019, 2020a;Phookamsak

et al., 2019;Dayarathne et al., 2020a,b;Konta et al., 2020;

Supplementary Table 2). Xylaria hypoxylon (CBS 122620) was

selected as the outgroup. For each gene, sequences were aligned

using MAFFT v. 7 (Katoh and Standley, 2013) and manually

improved where necessary using MEGA v. 6 (Tamura et al.,

2013). Ambiguously aligned sequences were excluded from

the analysis. Alignments were used to infer a preliminary

phylogenetic relationship for our sequences based on Maximum

Parsimony (MP) with PAUP v. 4.0b10 (Swoﬀord, 2003),

Maximum Likelihood (ML) with PhyML v. 3.0 (Guindon et al.,

2010), and Bayesian Inference (BI) analyses with MrBayes v. 3.1.2

(Ronquist and Huelsenbeck, 2003).

Maximum parsimony analysis was performed using a heuristic

search option of 1,000 random-addition sequences. The tree

bisection and reconnection (TBR) was selected as option to

the branch swapping algorithm (Swoﬀord, 2003). The branches

of zero length were collapsed, and all equally parsimonious

trees were saved. Clade stability was assessed with a bootstrap

analysis of 1,000 replicates (Hillis and Bull, 1993). Tree length

(TL), consistency index (CI), retention index (RI), and rescaled

consistency (RC) were calculated (Swoﬀord, 2003). ML analysis

including 1,000 bootstrap replicates (Hillis and Bull, 1993) was

conducted with a general time reversible (GTR) model of site

substitution, including gamma-distributed rate heterogeneity

and a proportion of invariant sites (Guindon et al., 2010). The

nucleotide model of evolution for each of the data partitions were

estimated by MrModeltest v. 2.3 (Posada and Crandall, 1998)

before the Bayesian analysis. BI analysis was performed using a

Markov Chain Monte Carlo (MCMC) algorithm with Bayesian

posterior probabilities (Rannala and Yang, 1996). Two MCMC

chains were run for 1,000,000 generations with a sampling

frequency at every 100th generation. The ﬁrst 25% of trees

were discarded as the burn-in phase of each analysis, and

the posterior probabilities (BPP) were calculated to assess the

remaining trees (Rannala and Yang, 1996). The branch support

from MP and ML analyses were evaluated with a bootstrapping

(BS) method of 1,000 replicates (Hillis and Bull, 1993). The

resulting trees were plotted in Figtree v. 1.4.4 and edited in Adobe

Illustrator CS6 v. 16.0.0. All sequences from this study were

deposited in GenBank (Supplementary Table 2). The multi-gene

sequence alignment ﬁles were submitted to TreeBASE (2accession

number: S27126).

RESULTS

Phylogenetic Analyses

The phylogenetic analysis combined ITS and tub2 contained

146 ingroup strains with 1,175 characters including gaps (713

for ITS and 462 for tub2), of which 471 were constant, 191

variable characters were parsimony-uninformative, and 513

characters were variable and parsimony-informative. The MP

analysis resulted 500 parsimonious trees, and the ﬁrst tree

(TL = 3,637, CI = 0.362, RI = 0.771, RC = 0.279) was presented

in Figure 1. For BI analyses, the best-ﬁt model of nucleotide

evolution was deduced on the AIC (ITS: GTR +I+G; tub2:

HKY +I+G). Tree topologies of ML and BI analyses did not

signiﬁcantly diﬀer from the MP. Topology of the phylogenetic

analyses were similar to the relevant recently published literature

(Senwanna et al., 2017;Shang et al., 2017, 2018;Hyde et al.,

2019, 2020a;Phookamsak et al., 2019;Dayarathne et al., 2020a;

Konta et al., 2020).

Based on phylogenetic analyses, the phylogram included 27

lineages, representing 21 known (Allocryptovalsa/Eutypella sensu

lato,Allodiatrype,Anthostoma,Cryptosphaeria 1, Cryptosphaeria

2, Cryptovalsa,Diatrypasimilis,Diatrype sensu stricto,Diatrypella

1, Diatrypella 2, Eutypa sensu lato,Eutypella sensu stricto,Eutypa

sensu stricto,Halocryptosphaeria,Halocryptovalsa,Halodiatrype,

Monosporascus,Neoeutypella,Pedumispora,Peroneutypa,

Quaternaria) and six incertae sedis clades. Eleven lineages are,

herein, described as nine new species and two known species

belonging to four genera in Diatrypaceae (Figure 1).

Some confused taxa were excluded in the current phylogram

after the primary analyses. Diatrype decorticata (ANM 1498),

Diatrype enteroxantha (HUEFS 155116), Diatrype macowaniana

(CBS 214.8), Diatrype oregonensis (DCA600), Diatrype polycocca

(CBS 213.87), Diatrype prominens (ATCC MYA-4410), Diatrype

whitmanensis (CDB011), Eutypella parasitica (CBS 210.39), and

Eutypella prunastri (CBS 277.87) are not the type strains,

which have single clade in phylogenetic tree or mixed

with in clade of other genera. And the sequence data of

Halocryptovalsa avicenniae (MAW 2017a) is inconsistent with

the position of genus.

Clade 06 (Diatrypella 2): This clade comprises nine newly

generated strains and other six Diatrypella strains with strong

statistical supports (MP/ML/BI = 96/99/1) in Figure 1.

Diatrypella hubeiensis (CFCC 52413) was the basal subclade

close to Da. yunnanensis.Diatrypella shennongensis (CFCC

52414 and CFCC 52415) formed a single clade with high

support (MP/ML/BI = 100/100/1). Diatrypella betulae (CFCC

52404, CFCC 52405 and CFCC 52406) clustered close to

Da. shennongensis.Diatrypella betulicola (CFCC 52411 and

CFCC 52412) also formed a distinct strongly supported clade

(MP/ML/BI = 95/96/1). The isolate CFCC 52409 clustered with

2www.treebase.org

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Zhu et al. Diatrypaceous Fungi in China

Diatrypella favacea (CBS 198.49, R191, and DL26C), which was

recognized as known species.

Clade 09 (Diatrype sensu stricto): The type species of Diatrype,

D. disciformis (CBS 205.87 and GB 5815), and other Diatrype

and Diatrypella species grouped with strong statistical supports

(MP/ML/BI = 100/97/1) in Figure 1. Our six new strains

clustered in this clade as three diﬀerent subclades viz. Diatrype

quercicola (CFCC 52418, CFCC 52419, and CFCC 52420),

D. castaneicola (CFCC 52425 and CFCC 52426) and D. betulae

(CFCC 52416). Diatrype betulae was the basal subclade close

to D. undulata.Diatrype castaneicola was the internal clade

with high support (MP/ML/BI = 99/100/1) close to D. stigma.

Diatrype quercicola formed a single clade with high support

(MP/ML/BI = 89/98/1) and grouped with D. virescens with

no support value.

Clade 12 (Allocryptovalsa/Eutypella sensu lato): This clade

comprises Allocryptovalsa and part Eutypella species clustered

with strong support values (MP/ML/BI = 87/94/1) in Figure 1.

Two isolates (CFCC 52433 and CFCC 52434) grouped together

with Eutypella citricola (HVVIT07 and HVGRF01) with strong

support (MP/ML/BI = 100/100/1). The isolates CFCC 52432

formed a separate branch separated from Eutypella citricola and

FIGURE 1 | Continued

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Zhu et al. Diatrypaceous Fungi in China

FIGURE 1 | Phylogram of Diatrypaceae based on combined ITS and tub2 sequence data. The MP and ML bootstrap support values above 70% are shown at the

ﬁrst and second positions, respectively. Thickened branches represent posterior probabilities above 0.95 from the BI. Ex-type strains are in bold, type species are

denoted with the superscript “TS” and the disputable type species are denoted with the superscript “TSQ.” Strains from the current study are in blue.

Eutypella vitis (MP/ML/BI = 95/96/1), which represented a new

species Allocryptovalsa castaneicola.Allocryptovalsa castaneae

(CFCC 52427, CFCC 52428, and CFCC 52429) also regarded

as a new species with the distinct strongly supported clade

(MP/ML/BI = 96/99/1), which was clustered with Allocryptovalsa

cryptovalsoidea,A. elaeidis,A. polyspora, and A. truncata

(MP/ML/BI = 99/100/1).

Taxonomy

Allocryptovalsa Senwanna, Phookamsak & K.D. Hyde,

Mycosphere 8(10): 1839 (2017).

Type:Allocryptovalsa polyspora Senwanna, Phookamsak &

K.D. Hyde, Mycosphere 8(10): 1840 (2017).

Known distribution: Australia, China, Germany, India,

Thailand, and United States (Saccardo, 1882;Trouillas et al.,

2011;Senwanna et al., 2017;Hyde et al., 2020b;Konta et al.,

2020; This study).

Notes:Allocryptovalsa typiﬁed with A. polyspora was originally

introduced to accommodate another two new combination

species (A. cryptovalsoidea and A. rabenhorstii), which was with

the character of immersed perithecia, polysporous asci, and

allantoid ascospores (Senwanna et al., 2017). Later, Hyde et al.

(2020a); Konta et al. (2020) reported A. elaeidis and A. truncata

isolated from Elaeis guineensis and decaying twig, respectively. In

this study, we introduce two additional species, Allocryptovalsa

castaneae and A. castaneicola, based on morphological coupled

with molecular data (Figure 1; clade 12).

Allocryptovalsa castaneae N. Jiang & X.L. Fan sp. nov. Figure 2.

MycoBank MB 837777.

Typiﬁcation: CHINA. Hebei Province, Qinhuangdao City,

Qinglong County, 119◦11052.2500 E 40◦22052.1300 N, 246 m

msl., from branches of Castanea mollissima, 16 Oct. 2017,

C.M. Tian & N. Jiang, holotype BJFU CF2020518, ex-type

culture CFCC 52428. Hebei Province, Qinhuangdao City,

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Zhu et al. Diatrypaceous Fungi in China

FIGURE 2 | Holomorph of Allocryptovalsa castaneae (BJFU CF2020518, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of

ascoma. (D) Longitudinal section through ascoma. (E) Conidiomata on the host. (F) Conidioma on the host. (G) Asci and ascospores. (H) Ascospores. (I,J) Conidia

attatch to conidiogenous cells. (K) Conidia. Scale bars: (A,F) =1mm;(B−E) = 500 µm; (G−K) =10µm.

Qinglong County, 119◦11052.2500 E 40◦22052.1300 N, 246 m msl.,

from branches of Castanea mollissima, 16 Oct. 2017, C.M. Tian &

N. Jiang, isotype BJM 240506, ex-isotype culture CFCC 52429.

Etymology: Named after the host genus from which it was

collected, Castanea.

Diagnosis: Phylogenetically sister to

Allocryptovalsa rabenhorstii, diﬀers by the smaller size of

ascospores (8–11 ×2.5–3.5 vs. 13.5–15 ×4–5 µm).

Descriptions:Necrotrophic on branches of Castanea

mollissima. Sexual morph: Stromata solitary to gregarious,

immersed in the bark, erumpent through the surface of bark,

with 3–5 perithecia arranged irregularly (0.3–)0.5–0.8 mm

(av. = 0.6 ±0.2 mm, n= 10) in diam. Ectostromatic disc

orange, unconspicuous, circular to oblong, with 3–5 ostioles

arranged irregularly per disc. Ostioles numerous, brown to

black, at the same level as the disc, scattered (85–)120–130 µm

(av. = 122.4 ±14.0 µm, n= 10) in diam. Perithecia outer

surface lacking powdery entostroma, black, ﬂask-shaped to

spherical, with discrete perithecial necks (320–)360–400(−420)

µm (av. = 379.7 ±19.7 µm, n= 10) in diam. Asci clavate to

elongate obovoid, polysporous, thin-walled, short pedicellate,

apically rounded (52–)60–83(–92) ×(11–)12–17(–25) µm

(av. = 71.5 ±11.4 ×14.4 ±2µm, n= 30). Ascospores

elongate-allantoid, thin-walled, pale yellowish to pale brown at

maturity, slightly curved, aseptate, 8–11(–13) ×2.5–3.5 (–4)

µm (av. = 10.1 ±0.8 ×3.1 ±0.4 µm, n= 30). Asexual morph:

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Coelomycetous. Conidiomata pycnidial, immersed in the bark,

scattered, erumpent through the surface of bark. Ectostromatic

disc ﬂat or concave, orange, surrounded by bark ﬂaps, circular to

ovoid, with 8–10 ostioles arranged circularly on per disc (200–

)260–320(–370) µm (av. = 280.8 ±43.3 µm, n= 10) in diam.

Ostioles black, at the same level as the disc surface (45–)60–70

µm (av. = 64.1 ±9.8 µm, n= 10) in diam. Conidiogenous

cells holoblastic conidiogenesis, approximately cylindrical,

hyaline, integrated, arising from pseudoparenchymatous

cells, unicellular, with wide base producing conidia

at the apex (15–)19–30(–31) ×(1–)1.5–2(–2.5) µm

(av. = 24.5 ±4.9 ×1.7 ±0.2 µm, n= 30).

Conidia hyaline, elongate-allantoid, not curved,

smooth, aseptate (4–)5–7(–8) ×(1–) 1.5–2(–2.5) µm

(av. = 5.9 ±0.9 ×1.8 ±0.2 µm, n= 30).

Culture characteristics: Cultures are initially white with

irregular margin, becoming dark green at the margin and

stopping growing with 7 cm in diam. after 2 weeks, comprising

dense, irregular, ﬂat mycelium.

Known host and distribution: Known on Castanea mollissima

and Juglans regia in China.

Additional collection examined: CHINA. Yunnan

Province, Chuxiong Yi Autonomous Prefecture, Dayao

County, 101◦20015.700 E 25◦44047.1900 N, 2,002 m msl.,

from branches of Juglans regia, August 07, 2015, N.

Zhao, paratype BJFU CF2020516, ex-paratype culture

CFCC 52427.

Notes: Three new strains isolated from branches of Castanea

mollissima and Juglans regia, show high support value

(MP/ML/BI = 99/100/1) with the closely clustered isolates

in Allocryptovalsa (Figure 1; Clade 12: Allocryptovalsa/Eutypella

sensu lato). Moreover, this species has diﬀerent morphological

characters. Morphological comparison of members of

Allocryptovalsa is provided in Supplementary Table 3.

Other species of this genus were not reported with asexual

morph. Therefore, Allocryptovalsa castaneae showed

coelomycetous asexual morph from host in China for the

ﬁrst time. In addition, Allocryptovalsa castaneae diﬀers from

A. castaneicola (from Castanea mollissima), A. cryptovalsoidea

(from Ficus carica), A. elaeidis (from Elaeis guineensis),

A. polyspora (from Hevea brasiliensis), and A. rabenhorstii

(from Vitis vinifera and Sambuscus nigra) in host association

(Saccardo, 1882;Trouillas et al., 2011;Senwanna et al., 2017;

Konta et al., 2020).

Allocryptovalsa castaneicola N. Jiang & X.L. Fan sp. nov.

Figure 3.

MycoBank MB 837787.

Typiﬁcation: CHINA. Hebei Province, Chengde

City, Kuancheng Manchu Nationality Autonomous

County, 118◦2705400 E 40◦3803700 N, 450 m msl., from

branches of Castanea mollissima, 14 Oct. 2017, C.M.

Tian & N. Jiang, holotype BJFU CF2020519, ex-type

culture CFCC 52432. Hebei Province, Chengde City,

Kuancheng Manchu Nationality Autonomous County,

118◦2705400 E 40◦3803700 N, 450 m msl., from branches of

Castanea mollissima, 14 Oct. 2017, C.M. Tian & N. Jiang,

isotype BJM 240515.

Etymology: Named after the host genus from which it was

collected, Castanea.

Diagnosis:Allocryptovalsa castaneicola diﬀers from other

Allocryptovalsa species by its polysporous asci and larger size of

ascospores (22–25 ×5–6 µm).

Descriptions:Necrotrophic on branches of Castanea

mollissima. Sexual morph: Stromata scattered to gregarious,

immersed in the bark, erumpent through the surface of bark,

with 8–10 perithecia arranged irregularly (1.5–)1.7–2.0 mm

(av. = 1.8 ±0.2 mm, n= 10) in diam. Ectostromatic disc brown,

circular to oblong, with more than 10 ostioles arranged circularly

per disc, 0.7–0.9(–1.0) mm (av. = 0.8 ±0.1 mm, n= 10) in

diam. Ostioles numerous, gregarious, umbilicate, 4-sulcate dark

brown to black, at the same level as the disc, 104–120(–140) µm

(av. = 114.9 ±11.3 µm, n= 10) in diam. Perithecia outer surface

coated with yellow, powdery entostromablack, ﬂask-shaped,

perithecial necks erumpent in groups (200–)250–320(–380) µm

(av. = 282.7 ±49.8 µm, n= 10) in diam. Asci clavate to elongate

obovoid, polysporous, thin-walled, long pedicellate, apically

ﬂat, 194–202 ×15–21 µm (av. = 198.4 ±3.3 ×18.9 ±0.5 µm,

n= 10). Ascospores elongate-allantoid, thin-walled, pale yellowish

to pale brown at maturity, slightly curved, aseptate, smooth,

22–25 ×5–6 µm (av. = 23.8 ±1.1 ×5.4 ±0.3 µm, n= 30).

Asexual morph: not observed.

Culture characteristics: Colonies are initially white, uniform,

becoming dark after 2 weeks.

Known host and distribution: Known only on Castanea

mollissima in Hebei Province, China.

Notes: The new species displays some features of morphology

typical of the recent genus Allocryptovalsa (well-developed

ascostromata producing polysporous asci and allantoid

ascospores) (Senwanna et al., 2017), although it appears

closer placed in Eutypella sensu lato (Figure 1; Clade 12).

Morphologically, Allocryptovalsa castaneicola diﬀers from the

closest species Eutypella australiensis by larger size of asci

(194–202 ×15–21 vs. 40–50 ×7–8.5 µm) and ascospores

(22–25 ×5–6 vs. 8–10 ×3µm) (Trouillas et al., 2010a).

Allocryptovalsa castaneicola was also distinguished from other

Eutypella species resembles by having polysporous asci rather

than the 8-spored asci (Trouillas et al., 2010b). Thus, we

introduce it here as a new species in genus Allocryptovalsa.

Moreover, Allocryptovalsa castaneicola can diﬀer from

Cryptovalsa species by having a yellow rather than white powdery

entostroma appeared on the ascomatal outer surface (Dayarathne

et al., 2020b). Also, phylogenetic analyses show aﬃnities of this

fungus with strains from Eutypella spp. Therefore, the assignment

of the strains to the genus Eutypella sensu lato (Figure 1; Clade

12) may require future reconsideration.

Diatrype Fr., Summa veg. Scand., Sectio Post.

(Stockholm): 384, 1849.

Type:Diatrype disciformis (Hoﬀm.) Fr., Summa veg. Scand.,

Sectio Post. (Stockholm): 385 (1849).

Known distribution: Asia, Europe, North America, Oceania,

and South Africa (Doidge, 1950;Munk, 1957;Conners, 1967;

Rappaz, 1987;Mulenko et al., 2008;Trouillas et al., 2010a,b).

Notes: The genus Diatrype was established by Fries (1849) with

Diatrype disciformis as the generic type, which have often been

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FIGURE 3 | Sexual morph of Allocryptovalsa castaneicola (BJFU CF2020519, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Longitudinal section

through ascoma. (D) Transverse section of ascoma. (E,F) Ascus and ascospores. (G) Ascospores. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–G) =10µm.

regarded as saprobes on decaying wood and have a strong ability

to resist harsh conditions (Senanayake et al., 2015). Due to the

taxonomic confusion, Diatrype may require a thorough revision

together with the entire family in the future.

Diatrype betulae H.Y. Zhu & X.L. Fan sp. nov. Figure 4.

MycoBank MB 837784.

Typiﬁcation: CHINA. Beijing City, Mentougou District,

Mount Dongling, Xiaolongmen Forestry Centre, 115◦26051.2700

E 39◦58019.6200 N, 1,302 m msl., from branches of Betula

davurica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan, holotype

BJFU CF2020510, ex-type culture CFCC 52416. Beijing City,

Mentougou District, Mount Dongling, Xiaolongmen Forestry

Centre, 115◦26051.2700 E 39◦58019.6200 N, 1,302 m msl., from

branches of Betula davurica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan,

isotype BJM 240512.

Etymology: Named after the host genus from which it was

collected, Betula.

Diagnosis: Phylogenetically, Diatrype betulae formed

a separate clade.

Descriptions:Necrotrophic on branches of Betula davurica.

Sexual morph: not observed. Asexual morph: Coelomycetous.

Conidiomata pycnidial, immersed in the bark, scattered,

erumpent slightly through the surface of bark, with multiple

locules and orange colloid conidial drops exuding from

the ostioles. Locules numerous, buﬀ, circular to ovoid, 1.0–

1.4 mm (av. = 1.2 ±0.2 mm, n= 10) in diam. Conidiogenous

cells approximately cylindrical, mostly straight, discrete

or integrated, arising from pseudoparenchymatous cells,

hyaline, unicellular, with wide base producing conidia

at the apex, holoblastic conidiogenesis (12–)14–20 ×1–

2µm (av. = 16.6 ±3.5 ×1.2 ±0.2 µm, n= 30). Conidia

hyaline, ﬁliform, smooth or rough, aseptate, 10–13 ×1–2 µm

(av. = 11.7 ±1.2 ×1.5 ±0.1 µm, n= 30).

Culture characteristics: Cultures are white, uniform, dense,

slow growing, reaching 4 cm after 2 weeks, not produced

pigmentation on PDA media.

Known host and distribution: Known only on Betula davurica

in Beijing City, China.

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FIGURE 4 | Asexual morph of Diatrype betulae (BJFU CF2020510, holotype). (A) Conidiomata on the host. (B) Conidioma on the host. (C,D) Transverse section of

conidioma. (E) Longitudinal section through conidioma. (F) Conidiogenous cells. (G,H) Conidia. Scale bars: (A) =1mm;(B−E) = 500 µm; (F−H) =10µm.

Notes:Diatrype betulae was isolated from branches of Betula

davurica in Beijing, China. One strain of Diatrype betulae

(CFCC 52416) clusters as a single lineage (Figure 1). Diatrype

albopruinosa,D. undulata, and D. stigma were also reported from

Betula sp. (Tiﬀany and Gilman, 1965;Chlebicki, 2005;Mulenko

et al., 2008;Goos, 2010). However, Diatrype betulae can be

easily distinguished from the other three. Diatrype albopruinosa

and D. undulata lack the asexual morph and sequences of

D. albopruinosa are unavailable. Diatrype stigma was known

on various hosts with worldwide distribution, but it can

diﬀer from D. betulae by smaller conidia (4.5–7.5 ×1–2 vs.

10–13 ×1–2 µm) (Rappaz, 1987). Diatrype betulae is also

phylogenetically closely related to D. bullata,D. castaneicola,

D. disciformis,D. iranensis,D. macrospora,D. quercicola,

D. quercina,D. spilomea, and D. virescens.Diatrype betulae

can diﬀer from D. bullata that is a common species isolated

from willows in the northern hemisphere in host association

(Vasilyeva and Ma, 2014). Diatrype iranensis,D. macrospora,

D. quercicola, and D. quercina were only reported from

Quercus sp. (Croxall, 1950;Mehrabi et al., 2015, 2016). Diatrype

betulae can easily be distinguished from them by its plant

host (Betula sp.) and smaller conidia (18–38 ×0.6–0.8 µm

in D. iranensis and 20–40 ×0.6–0.8 µm in D. macrospora)

(Mehrabi et al., 2015, 2016). In morphology, D. betulae can

be diﬀerentiated from D. disciformis and D. virescens by

having asexual morph. Moreover, D. betulae diﬀers from other

closest species D. castaneicola and D. spilomea by the size of

conidia (10–13 ×1–2 vs. 4–6 ×1–1.5, 13–18.5 ×1–1.2 µm)

(Rappaz, 1987).

Diatrype castaneicola N. Jiang & X.L. Fan sp. nov. Figure 5.

MycoBank MB 837785.

Typiﬁcation: CHINA. Hebei Province, Qinhuangdao City,

Qinglong County, 119◦11052.2500 E 40◦22052.1300 N, 246 m

msl., from branches of Castanea mollissima, 16 Oct. 2017,

C.M. Tian & N. Jiang, holotype BJFU CF2020515, ex-type

culture CFCC 52425. Hebei Province, Qinhuangdao City,

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FIGURE 5 | Asexual morph of Diatrype castaneicola (BJFU CF2020515, holotype). (A) Conidiomata on the host. (B) Conidioma on the host. (C) Transverse section

of conidioma. (D) Longitudinal section through conidioma. (E) Conidiogenous cells. (F,G) Conidia. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–G) =10µm.

Qinglong County, 119◦11052.2500 E 40◦22052.1300 N, 246 m msl.,

from branches of Castanea mollissima, 16 Oct. 2017, C.M. Tian &

N. Jiang, isotype BJM 240513, ex-isotype culture CFCC 52426.

Etymology: Named after the host genus from which it was

collected, Castanea.

Diagnosis: Phylogenetically, Diatrype castaneicola formed

a separate clade.

Descriptions:Necrotrophic on branches of Castanea

mollissima. Sexual morph: not observed. Asexual morph:

Coelomycetous. Conidiomata pycnidial, immersed in the bark,

scattered, erumpent slightly through the surface of bark, with

multiple locules (0.7–)0.8–1.2(–1.6) mm (av. = 1.0 ±0.2 mm,

n= 10). Ectostromatic disc brown, unconspicuous, circular

to ovoid. Ostiole unconspicuous, gray to black, at the

same the level as the disc surface, covered by ectostroma

tissue. Locules numerous, circular to ovoid, 1.0–1.4 mm

(av. = 1.2 ±0.2 mm, n= 10) in diam. Conidiogenous

cells approximately cylindrical, mostly straight, discrete

or integrated, arising from pseudoparenchymatous cells,

hyaline, unicellular, with wide base producing conidia at

the apex, holoblastic conidiogenesis (15–)18–26(–33) ×1–

1.5 µm (av. = 22.5 ±3.5 ×1.2 ±0.2 µm, n= 30). Conidia

hyaline, elongate-allantoid, slightly curved, smooth, aseptate,

multiguttulate, often containing guttules per cell, 4–6 ×1–1.5 µm

(av. = 5.3 ±0.6 ×1.3 ±0.2 µm, n= 30).

Culture characteristics: Colonies are white, dense, not

produced pigmentation on PDA media. Pycnidia distributed

irregularly on colony surface with yellow cream conidial drops

exuding from the ostioles.

Known host and distribution: Known only on Castanea

mollissima in Hebei Province, China.

Notes:Diatrype castaneicola was isolated from branches

of Castanea mollissima in Hebei Province, China. Our new

isolates (CFCC 52425 and CFCC 52426) grouped in Diatrype

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FIGURE 6 | Sexual morph of Diatrype quercicola (BJFU CF2020512, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of

ascoma. (D) Longitudinal section through ascoma. (E,F) Ascus and ascospores. (G) Ascospores. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–G) =10µm.

sensu stricto as a separate clade with high statistical support

(MP/ML/BI = 99/100/1) (Figure 1). Diatrype castaneicola

diﬀers from the closely related one, D. stigma, by its smaller

conidia (4–6 ×1–1.5 vs. 4.5–7.5 ×1–2 mm) (Rappaz, 1987).

Currently, Diatrype castaneicola is reported with only the

asexual morph, thus more studies are essential to report the

sexual morph.

Diatrype quercicola H.Y. Zhu & X.L. Fan sp. nov. Figure 6.

MycoBank MB 837786.

Typiﬁcation: CHINA. Beijing City, Mentougou District,

Mount Dongling, Xiaolongmen Forestry, 115◦26051.2700 E

39◦58019.6200 N, 1,267 m msl., from branches of Quercus

mongolica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan, holotype

BJFU CF2020512, ex-type culture CFCC 52418. Beijing

City, Mentougou District, Mount Dongling, Xiaolongmen

Forestry, 115◦26051.2700 E 39◦58019.6200N, 1,267 m msl.,

from branches of Quercus mongolica, 21 Aug. 2017,

H.Y. Zhu & X.L. Fan, isotype BJM 240514, ex-isotype

culture CFCC 52419.

Etymology: Named after the host genus from which it was

collected, Quercus.

Diagnosis: Phylogenetically, Diatrype quercicola formed a

separate clade. However its asci are polysporous and diﬀer from

the common 8-ascospores asci in Diatrype.

Descriptions:Necrotrophic on branches of Quercus

mongolica. Sexual morph: Stromata solitary, immersed in

the bark, erumpent through the surface of bark, with more

than 10 perithecia arranged irregularly (2.3–)2.5–2.9 mm

(av. = 2.7 ±0.2 mm, n= 10) in diam. Ectostromatic disc

brown, circular to oblong, with more than 10 ostioles arranged

regularly per disc (1.5–)1.7–2.4 mm (av. = 2.0 ±0.3 mm,

n= 10) in diam. Ostioles dark brown to black, at the

same level as the disc, scattered, 210–275(–335) µm

(av. = 253.3 ±22.1 µm, n= 10) in diam. Perithecia outer

surface coated with yellow, powdery entostromablack,

ﬂask-shaped, with discrete perithecial necks (475–)515–

620(–665) µm (av. = 566.6 ±53.2 µm, n= 10) in diam. Asci

clavate to elongate obovoid, polysporous, thin-walled, long

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pedicellate, apically rounded, 172–183 ×(16–)20–43 µm

(av. = 178 ±3.3 ×31.7 ±10.9 µm, n= 10). Ascospores

elongate-allantoid, thin-walled, pale yellowish to pale brown

at maturity, slightly curved, aseptate, multiguttulate, often

containing 1–3 symmetrical guttules per cell, 17–27 ×4–

6µm (av. = 22.6 ±2.6 ×5.4 ±0.6 µm, n= 30). Asexual

morph: not observed.

Culture characteristics: Colonies are white, irregular, reaching

9 cm after 7 days, not produced pigmentation on PDA media.

Known host and distribution: Known only on Quercus

mongolica in Beijing City, China.

Additional collection examined: CHINA. Beijing City,

Mentougou District, Mount Dongling, Xiaolongmen Forestry,

115◦26051.2700 E 39◦58019.6200N, 1,267 m msl., from branches of

Quercus mongolica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan, BJFU

CF2020513, living culture CFCC 52420.

Notes:Diatrype quercicola is a unique Diatrype species

isolated from Quercus mongolica in China. Asci of this

species are polysporous and diﬀer from the common 8-

ascospores asci of Diatrype, including the closely related

taxa, D. virescens. Moreover, D. quercicola can diﬀer from

D. virescens by larger asci and ascospres (172–183 ×20–

43 vs. 35–40 ×4–6 µm; 17–27 ×4–6 vs. 12–14 ×2.5–

3µm) (Vasilyeva and Stephenson, 2005). Nevertheless, based

on phylogeny analyses, this taxon appears best placed in

Diatrype. Therefore, the assignment of this species to Diatrype

may require reconsideration due to the taxonomic confusion

around Diatrypaceae.

Diatrype albopruinosa,D. standleyi, and D. stigmaoides were

also isolated from Quercus sp. (Tiﬀany and Gilman, 1965;

Rappaz, 1987;Mendez-Mayboca et al., 2008;Vasilyeva and

Stephenson, 2009). Diatrype albopruinosa and D. standleyi diﬀer

from D. quercicola by its 8-ascospores asci (Vasilyeva and Ma,

2014). Diatrype stigmaoides diﬀers from D. quercicola by its

hyaline ascospores (Vasilyeva and Stephenson, 2009).

Diatrypella (Ces. & De Not.) Nitschke, Pyr. Germ: 69, 1867.

Type:Diatrypella verruciformis (Ehrh.) Nitschke, Pyrenomyc.

Germ. 1: 76, 1867 [Current name: Diatrypella favacea (Fr.)].

Known distribution: Asia, Europe, North America, Oceania,

South Africa, and South America (Unamuno, 1941;Doidge, 1950;

Rao, 1966;Conners, 1967;Hanlin, 1992;Crous et al., 2016).

Notes:Diatrypella was introduced by Nitschke (1867) to

accommodate Diatrype sect. Diatrypella Ces. & De Not. (1863).

The type species is disputable that Croxall (1950) relegated

Da. verruciformis (as D. verrucaeformis) to synonymy with

Da. favacea, whereas Munk (1957) recognized both species as

diﬀerent fungi appear to have been frequently included under

Da. verruciformis.Glawe and Rogers (1984) believed Diatrypella

was well distinguished genus as its well-delveloped stromata

and the single host aﬃliation (Da. verruciformis on Alnus

and Da. favacea on Betula). Further studies are needed to

clarify them. Vasilyeva and Stephenson (2005) pointed out that

Diatrypella morphologically resembled Cryptovalsa. Diatrypella

and Cryptovalsa were mentioned as the polysporous complement

of Diatrype and Eutypa (Vasilyeva and Stephenson, 2005).

Nevertheless, it is still diﬃcult to determine the diﬀerences

between Diatrypella and Cryptovalsa based on morphological

characters (Acero et al., 2004;Vasilyeva and Stephenson, 2005).

Therefore, multilocus phylogeny including more representative

taxa are needed to clarify the relationship among species in

Diatrypella (Mehrabi et al., 2015).

Newly generated nine isolates show aﬃnities to Diatrypella

favacea clade based on phylogenetic analyses. Therefore, we

prefer the assignment of the strains to the genus Diatrypella

(Figure 1; Clade 06) preliminarily and tentatively, as it may

require future reconsideration after the typiﬁcation work on type

species of Diatrypella.

Diatrypella betulae H.Y. Zhu & X.L. Fan sp. nov. Figure 7.

MycoBank MB 837778.

Typiﬁcation: CHINA. Hubei Province, Shennongjia Forest

District, Shennong Stream, 110◦17051.5400 E 31◦28015.7900 N,

2273 m msl., from branches of Betula albosinensis, 17 Aug.

2017, Z. Du & Q. Yang, holotype BJFU CF2020501, ex-

type culture CFCC 52406. Hubei Province, Shennongjia Forest

District, Shennong Stream, 110◦17051.5400 E 31◦28015.7900 N,

2273 m msl., from branches of Betula albosinensis, 17 Aug.

2017, Z. Du & Q. Yang, isotype BJM 240507, ex-isotype culture

CFCC 52404.

Etymology: Named after the host genus from which it was

collected, Betula.

Diagnosis: Phylogenetically sister to Diatrypella shennongensis,

diﬀer by the number of perithecia.

Descriptions:Necrotrophic on branches of Betula albosinensis.

Sexual morph: Stromata solitary, immersed in the bark, erumpent

through the surface of bark, with 7–10 perithecia arranged

irregularly (1.0–)1.6–2.1 mm (av. = 1.8 ±0.2 mm, n= 10)

in diam. Ectostromatic disc orange, circular to oblong, with

7–10 ostioles arranged regularly per disc, 0.8–1.3(–1.5) mm

(av. = 1.0 ±0.3 mm, n= 10) in diam. Ostioles numerous,

scattered, umbilicate, sulcate, dark brown to black, at the same

level as the disc (125–)175–240 µm (av. = 190.4 ±38.0 µm,

n= 10) in diam. Perithecia outer surface lacking powdery

entostroma, black, ﬂask-shaped, with long discrete perithecial

necks (430–)530–830(–870) µm (av. = 680.2 ±152.7 µm, n= 10)

in diam. Asci clavate to elongate obovoid, polysporous, thin-

walled, long pedicellate, apically rounded to ﬂat, 132–140 ×7.5–

10.5(–11.5) µm (av. = 136.2 ±3×9.5 ±1.4 µm, n= 10).

Ascospores elongate-allantoid, thin-walled, pale yellowish to pale

brown at maturity, slightly curved, smooth, aseptate (4.5–)5–

7×1–2 µm (av. = 5.7 ±0.5 ×1.5 ±0.2 µm, n= 30). Asexual

morph: not observed.

Culture characteristics: Cultures are ﬂat, reaching 9 cm diam.

after 7–10 days. Colonies white, rough on surface, not produced

pigmentation on PDA media.

Known host and distribution: Known only on Betula

albosinensis in Hubei Province, China.

Additional collection examined: CHINA. Hubei Province,

Shennongjia Forest District, Shennong Stream, 110◦17051.5400

E 31◦28015.7900 N, 2,273 m msl., from branches of Betula

albosinensis, 17 Aug. 2017, Z. Du & Q. Yang, BJFU CF2020502,

living culture CFCC 52405.

Notes: Three strains representing Diatrypella betulae

appear most closely related to Da. shennongensis reported

from the same host plant Betula sp., which also clustered

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FIGURE 7 | Sexual morph of Diatrypella betulae (BJFU CF2020501, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of

ascoma. (D) Longitudinal section through ascoma. (E) Ascus and ascospores. (F) ascus. (G) Ascospores. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–G) =10µm.

in a well-supported clade (MP/ML/BI = 92/100/1). In

morphology of asci and ascospores, Diatrypella betulae

resembles Da. shennongensis by the size of asci and ascospores

(132–140 ×7.5–10.5 vs. 129–140 ×8–12; 5–7 ×1–2 vs.

5–6.5 ×1–1.5 µm), but they can be distinguished by

the number of perithecia (less than 10 vs. more than 10).

Moreover, Diatrypella betulae diﬀers from Da. shennongensis

based on ITS and tub2 loci (67/665 in ITS and 13/416

in tub2).

Diatrypella betulicola H.Y. Zhu & X.L. Fan sp. nov. Figure 8.

MycoBank MB 837779.

Typiﬁcation: CHINA. Beijing City, Mentougou District,

Mount Dongling, Xiaolongmen Forestry Centre, 115◦26051.2700

E 39◦58019.6200 N, 1,209 m msl., from branches of Betula

davurica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan, holotype

BJFU CF2020505, ex-type culture CFCC 52411. Beijing City,

Mentougou District, Mount Dongling, Xiaolongmen Forestry

Centre, 115◦26051.2700 E 39◦58019.6200 N, 1,209 m msl., from

branches of Betula davurica, 21 Aug. 2017, H.Y. Zhu & X.L. Fan,

isotype BJM 240508.

Etymology: Named after the host genus from which it was

collected, Betula.

Diagnosis:Diatrypella betulicola is diﬀerent from other

species of Diatrypella on host association and the size of

asci and ascospores.

Descriptions:Necrotrophic on branches of Betula davurica

and B. platyphylla. Sexual morph: Stromata solitary, immersed

in the bark, erumpent through the surface of bark, with

7–10 perithecia arranged regularly (1.3–)1.5–1.9(–2.1) mm

(av. = 1.7 ±0.2 mm, n= 10) in diam. Ectostromatic disc brown

to black, circular to oblong, with more than 10 ostioles arranged

regularly per disc (0.5–)0.6–1.2(–1.4) mm (av. = 0.9 ±0.3 mm,

n= 10) in diam. Ostioles numerous, scattered, umbilicate,

sulcate, dark brown to black, at the same level as the disc

(145–)185–240(–270) µm (av. = 211.0 ±31.1 µm, n= 10)

in diam. Perithecia outer surface lacking powdery entostroma,

black, ﬂask-shaped, with short discrete perithecial necks (520–

)600–730(–790) µm (av. = 667.8 ±61.1 µm, n= 10) in

diam. Asci clavate to elongate obovoid, polysporous, thin-

walled, long pedicellate, apically ﬂat,117–133 ×10–12 µm

(av. = 124.9 ±7.7 ×10.6 ±0.5 µm, n= 10). Ascospores

elongate-allantoid, thin-walled, pale yellowish to pale brown at

maturity, slightly curved, aseptate, smooth or multiguttulate,

occasionally containing one guttule per cell, 5–8 ×1–2 µm

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FIGURE 8 | Sexual morph of Diatrypella betulicola (BJFU CF2020505, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of

ascoma. (D) Longitudinal section through ascoma. (E,F) Ascus and ascospores. (G) Ascospores. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–G) =10µm.

(av. = 6.6 ±0.6 ×1.6 ±0.2 µm, n= 30). Asexual

morph: not observed.

Culture characteristics: Cultures are ﬂuﬀy, reaching 9 cm

after 7 days, becoming pale yellow at the margin after

2 weeks. Colonies dense with aerial mycelium at the center,

sparse at the margin.

Known host and distribution: Known on Betula davurica and

B. platyphylla in Beijing City, China.

Additional collection examined: CHINA. Beijing City,

Mentougou District, Mount Dongling, Xiaolongmen Forestry,

115◦26051.2700 E 39◦58019.6200 N, 1,209 m msl., from branches of

Betula platyphylla, 21 Aug. 2017, H.Y. Zhu & X.L. Fan, paratype

BJFU CF2020506, ex-paratype culture CFCC 52412.

Notes:Diatrypella betulicola was isolated from branches

of Betula davurica and Betula platyphylla in China.

Phylogenetically, two strains representing Diatrypella betulicola

cluster in a well-supported clade (MP/ML/BI = 95/96/1) in

Diatrypella 2 clade (Figure 1). Diatrypella betulae,Da. favacea,

and Da. shennongensis are the most closely related species.

Diatrypella betulicola can be diﬀerentiated from Da. betulicola

and Da. favacea by the number of ostioles (more than 10

ostioles in Diatrypella betulae and less than 10 ostioles in Da.

betulicola and Da. favacea). Moreover, Da. betulicola can be

easily distinguished from Da. shennongensis by unique size of

asci (117–133 ×10–12 vs. 129–140 ×8–12 µm) and ascospore

(5–8 ×1–2 vs. 5–6.5 ×1–1.5 µm). Moreover, it is diﬀerent from

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Zhu et al. Diatrypaceous Fungi in China

other species of Diatrypella on host association and the size of

asci and ascospores (Supplementary Table 4).

Diatrypella favacea (Fr.) Ces. & De Not., Sfer. Ital.: 29 (1863).

Figure 9.

Descriptions:Necrotrophic on branches of Betula platyphylla.

Sexual morph: Stromata solitary, consisting of an inconspicuous

pale yellow ectostromatic disc, immersed in the bark, erumpent

through the surface of bark, with 7–10 perithecia arranged

irregularly (1.9–)2.9–3.3 mm (av. = 3.1 ±0.2 mm, n= 10)

in diam. Ectostromatic disc brown to black, circular to

oblong, with 5–7 ostioles arranged irregularly per disc

(0.5–)1.8–2.3(–2.4) mm (av. = 2.1 ±0.2 mm, n= 10) in

diam. Ostioles numerous, gregarious, umbilicate, sulcate,

dark brown to black, at the same level as the disc (230–

)240–260(–280) µm (av. = 252.4 ±16.6 µm, n= 10) in

diam. Paraphyses elongate cylindrical, 118–125 ×2–3.5 µm

(av. = 121.6 ±2.7 ×2.7 ±0.7 µm, n= 10). Perithecia outer

surface lacking powdery entostroma, black, ﬂask-shaped to

spherical, with long discrete perithecial necks (690–)780–

850(940) µm (av. = 821.9 ±34.1 µm, n= 10) in diam. Asci

clavate to elongate obovoid, polysporous, thin-walled, long

pedicellate, apically rounded, 64–124 ×(9–)9.5–12(–12.5)

µm (av. = 92.7 ±14.5 ×10.3 ±1.7 µm, n= 10). Ascospores

short-allantoid, thin-walled, pale yellowish to pale brown

at maturity, slightly curved, aseptate, multiguttulate, often

containing one guttulae per cell (3.5–)4–5.5(–6) ×1.5–2 µm

(av. = 4.8 ±0.5 ×1.8 ±0.2 µm, n= 30). Asexual

morph: not observed.

Culture characteristics: Cultures are white, uniform, attaining

9 cm in 7 days. Colonies sparse at the center, medium

dense at the margin, rough on the surface, not produced

pigmentation on PDA media.

Known host and distribution: Known on various hosts with

worldwide distribution3.

Collection examined: CHINA. Xinjiang Uygur Autonomous

Region, Bortala Mongol Autonomous Prefecture, Wenquan

County, 81◦46022.9600 E 45◦13008.4700 N, 1,439 m msl., from

branches of Betula platyphylla, July 15, 2017, C.M. Tian & R. Ma,

BJFU CF2020504, living culture CFCC 52409.

Notes:Diatrypella favacea was reported to be restricted

to Betula spp. in previous study (Saccardo, 1882;Glawe and

Rogers, 1984), but then it got involved in the problematic

species concept and delimitation with Da. verruciformis

(Farr et al., 1989). Diatrypella favacea and Da. pulvinata

clustered in a single clade until Hyde et al. (2020b) reported

Da. yunnanensis. The strain CFCC 52409 clusters with

Diatrypella favacea (CBS 198.49, DL26C, and R191) in

a separate lineage. Morphologically, our strain is similar

to those previously reported in terms of the size of asci

(64–124 ×9.5–12 vs. 70–90 ×8–12 µm) and ascospores

(4–5.5 vs. 6–8 µm) (Vasilyeva and Stephenson, 2005).

The current deﬁnition of Diatrypella favacea seems to be

diﬃcult due to the lack of type material with available

living culture or DNA sequence data. Thus, the current

3https://nt.ars-grin.gov/fungaldatabases

identiﬁcation is preliminary and awaits further studies

of typiﬁcation.

Diatrypella hubeiensis H.Y. Zhu & X.L. Fan sp. nov. Figure 10.

MycoBank MB 837781.

Typiﬁcation: CHINA. Hubei Province, Shennongjia

Forest District, Tianyan Scenic Area, 110◦27036.7100 E

31◦42059.1000 N, 2,140 m msl., from branches of Betula

davurica, 16 Aug. 2017, Z. Du & Q. Yang, holotype

BJFU CF2020507, ex-type culture CFCC 52413. Hubei

Province, Shennongjia Forest District, Tianyan Scenic Area,

110◦27036.7100 E 31◦42059.1000 N, 2,140 m msl., from branches

of Betula davurica, 16 Aug. 2017, Z. Du & Q. Yang, isotype

BJM 240510.

Etymology: Named after the location where it was

collected, Hubei Province.

Diagnosis: Phylogenetically sister to Diatrypella

yunnanensis, diﬀer by smaller size of ascospores

(6–8.5 ×1–2 vs. 18–22 ×3–4 µm).

Descriptions:Necrotrophic on branches of Betula davurica.

Sexual morph: Stromata solitary, immersed in the bark,

erumpent through the surface of bark, with 5–7 perithecia

arranged regularly (1.4–)1.7–2.5(–3.0) mm (av. = 2.1 ±0.4 mm,

n= 10) in diam. Ectostromatic disc brown to black, circular to

oblong, with 5–7 ostioles arranged irregularly per disc (0.9–)1.0–

2.0(–2.6) (av. = 1.5 ±0.4 mm, n= 10) in diam. Ostioles numerous,

gregarious, umbilicate, sulcate, dark brown to black, at the same

level as the disc (120–)130–170(–200) µm (av. = 143.1 ±30.4 µm,

n= 10) in diam. Perithecia outer surface lacking powdery

entostroma, black, ﬂask-shaped to spherical, with long discrete

perithecial necks, 500–680(–720) µm (av. = 619.3 ±71.3 µm,

n= 10) in diam. Asci clavate to elongate obovoid, occasionally

similar to an inverted volumetric ﬂask, polysporous, thin-walled,

long pedicellate, apically rounded to ﬂat, 189–240 ×18–21 µm

(av. = 213.4 ±13.3 ×19.8 ±0.5 µm, n= 10). Ascospores

elongate-allantoid, thin-walled, slightly curved, aseptate,

multiguttulate, often containing two symmetrical guttules per

cell, 6–8.5(–9) ×1–2 µm (av. = 7.4 ±0.7 ×1.6 ±0.2 µm,

n= 30). Asexual morph: not observed.

Culture characteristics: Cultures are white, ﬂuﬀy, fast growing,

attaining 9 cm in 7 days. Colonies dense, slightly raised with aerial

mycelium, not produced pigmentation on PDA media.

Known host and distribution: Known only on Betula davurica

in Hubei Province, China.

Notes: The only strain CFCC 52413 representing Diatrypella

hubeiensis clusters with Da. pulvinata and Da. yunnanensis.

However, Diatrypella pulvinata was introduced as an asexual

fungus in Quercus garryana. Diatrypella hubeiensis diﬀers from

its closest relative Da. yunnanensis by larger asci (189–240 ×18–

21 vs. 105–210 ×15–30 µm) and smaller size of ascospores

(6–8.5 ×1–2 vs. 18–22 ×3–4 µm) (Hyde et al., 2020b).

Diatrypella shennongensis H.Y. Zhu & X.L. Fan sp. nov.

Figure 11.

MycoBank MB 837780.

Typiﬁcation: CHINA. Hubei Province, Shennongjia Forest

District, Shennong Ding, Shennong Camp, 110◦17028.3900

E 31◦26034.5900 N, 2,647 m msl., from branches of Betula

albosinensis, 17 Aug. 2017, Z. Du & Q. Yang, holotype

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FIGURE 9 | Sexual morph of Diatrypella favacea (BJFU CF2020504). (A) Ascomata on the host. (B,C) Ascoma on the host. (D) Transverse section of ascoma.

(E) Longitudinal section through ascoma. (F) Asci. (G) Ascus. (H) Ascus and ascospores. (I) Ascospores. Scale bars: (A) =1mm;(B–E) = 500 µm; (F–I) =10µm.

BJFU CF2020508, ex-type culture CFCC 52415. Hubei

Province, Shennongjia Forest District, Shennong Ding,

Shennong Camp, 110◦17028.3900 E 31◦26034.5900 N, 2,647 m

msl., from branches of Betula albosinensis, 17 Aug. 2017,

Z. Du & Q. Yang, isotype BJM 240509, ex-isotype culture

CFCC 52414.

Etymology: Named after the location where it was

collected, Shennong Ding.

Diagnosis: Phylogenetically, sister to Diatrypella betulae, diﬀer

by ITS and tub2 loci (67/665 in ITS and 13/416 in tub2).

Descriptions:Necrotrophic on branches of Betula albosinensis.

Sexual morph: Stromata solitary, immersed in the bark, erumpent

through the surface of bark, causing a pustulate bark surface, with

more than 10 perithecia arranged irregularly 1.9–2.8(–3.7) mm

(av. = 2.5 ±0.6 mm, n= 10) in diam. Ectostromatic disc orange,

circular to oblong, with more than 10 ostioles arranged regularly

to irregularly per disc (1–)1.1–2.2(–2.6) mm (av. = 1.7 ±0.6 mm,

n= 10) in diam. Ostioles numerous, scattered, umbilicate, sulcate,

dark brown to black, at the same level as the disc (160–)170–

200(–215) µm (av. = 184.6 ±5.4 µm, n= 10) in diam.

Perithecia outer surface lacking powdery entostroma, black, ﬂask-

shaped to spherical, with long discrete perithecial necks (525–

)630–850(–970) µm (av. = 748.4 ±119.4 µm, n= 10) in

diam. Asci clavate to elongate obovoid, polysporous, thin-walled,

long pedicellate, apically rounded to ﬂat, 129–140 ×(5–)8–

12 µm (av. = 135 ±4.5 ×9.8 ±1.7 µm, n= 10). Ascospores

elongate-allantoid, thin-walled, pale yellowish to pale brown at

maturity, slightly curved, smooth, aseptate (4.5–)5–6.5(–7) ×1–

1.5 µm (av. = 5.8 ±0.6 ×1.3 ±0.2 µm, n= 30). Asexual

morph: not observed.

Culture characteristics: Cultures are white, dense, uniform,

ﬂuﬀy, growing up to 4 cm in diam. After 3 days, and

reaching 9 cm within 10 days. Colonies do not produced

pigmentation on PDA media.

Known host and distribution: Known only on Betula

albosinensis in Hubei Province, China.

Notes:Diatrypella shennongensis can be distinguished from its

closest relative, Da. betulae, by its number of perithecia (less than

10 vs. more than 10) in one stroma and base number diﬀerence

(67/665 in ITS and 13/416 in tub2). In addition, the multigene

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FIGURE 10 | Sexual morph of Diatrypella hubeiensis (BJFU CF2020507, holotype). (A) Ascomata on the host. (B) Ascoma on the host. (C) Transverse section of

ascoma. (D) Longitudinal section through ascoma. (E,F) ascus. (G, H) Ascospores. Scale bars: (A) =1mm;(B–D) = 500 µm; (E–H) =10µm.

phylogenetic analyses support this species as a new species with

high statistical support (MP/ML/BI = 100/100/1).

Eutypella (Nitschke) Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,

Padua, Sér. 44: 80, 1875.

Type:Eutypella cerviculata (Fr.) Sacc., Syll. Fung.

(Abellini) 1: 146, 1882.

Known distribution: Asia, Europe, North America, Oceania,

South Africa, and South America (Doidge, 1950;Rappaz, 1987;

Trouillas et al., 2011;Jayawardena et al., 2018).

Notes: The genus Eutypella was established by Saccardo

(1875) with E. cerviculata as the type species (Saccardo, 1882).

Eutypella species was mainly associated with canker diseases in

Vitis vinifera (Vasilyeva and Stephenson, 2006;Trouillas et al.,

2011;Luque et al., 2012). Although 251 species epithets of

Eutypella have been listed in Index Fungorum (2021), the most

of species are lacking DNA sequence. In phylogenetic analyses

of Diatrypaceae, it showed that Eutypella was polyphyletic

(Acero et al., 2004;Chacón et al., 2013;de Almeida et al.,

2016;Shang et al., 2017). Thus, further studies of the taxa in

Eutypella is needed.

Eutypella citricola Speg., Anales del Museo Nacional de

Buenos Aires 6: 245, 1898.

Descriptions: see Trouillas et al. (2011).

Known host and distribution: Known from Citrus limon,

C. sinensis,C. paradisi,Schinus molle var. areira,Ulmus procera

in Australia; Vitis vinifera in United States, Morus alba in China.

Collections examined: CHINA. Jiangsu Province, Yangzhou

City, 119◦28011.8100 E, 32◦47025.1000 N, 1 m msl., from branches

of Morus alba, 12 Nov. 2017, C.M. Tian & N. Jiang, BJFU

CF2020520, living culture CFCC 52433; ibid., BJFU CF2020521,

living culture CFCC 52434.

Notes: The current strains CFCC 52433 and CFCC 52434

cluster in a well-supported clade (MP/ML/BI = 100/100/1)

with Eutypella citricola (HVVIT07 and HVGRF01), residing in

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FIGURE 11 | Sexual morph of Diatrypella shennongensis (BJFU CF2020508, holotype). (A) Ascomata on the host. (B,C) Ascoma on the host. (D) Transverse section

of ascoma. (E) Longitudinal section through ascoma. (F,G) Ascus and ascospores. (H) Ascospores. Scale bars: (A) =1mm;(B–E) = 500 µm; (F–H) =10µm.

Allocryptovalsa/Eutypella sensu lato (Figure 1; Clade 12). In

morphology, this fungus can be identiﬁed by its 3–4 sulcate

ostioles, 8-spored, 55–80 ×7.5–9 µm asci and (9–)10.5–

12(–13) ×2–3 µm ascospores (Trouillas et al., 2011). This

study represents the ﬁrst record of this species from Morus alba.

Species of Allocryptovalsa represents similarity to Cryptovalsa

in morphology by having polysporous asci and is diﬀerent

from the eight spored asci of Eutypella (Senwanna et al., 2017).

Unfortunately, we did not observe any microscopic feature

from premature ascostromata. Thus, the current identiﬁcation

is preliminary and tentative, as it requires typiﬁcation before a

stable species concept achieved.

Other Genera Included in Diatrypaceae

In here, we follow Hyde et al. (2020a); Wijayawardene et al.

(2020) to list the genera in Diatrypaceae. A taxonomic key to

distinguish 23 genera of Diatrypaceae is provided.

Allodiatrype Konta & K.D. Hyde, Mycosphere

11(1): 247, 2020.

MycoBank MB 1814.

Type:Allodiatrype arengae Konta & K.D. Hyde, Mycosphere

11(1): 249, 2020.

Known distribution: Thailand (Li et al., 2016;Konta et al.,

2020).

Notes:Allodiatrype was established by Konta et al. (2020)

and typiﬁed by A. arengae. In the meanwhile, A. elaeidicola,

A. elaeidis, and A. thailandica were also accommodated to this

genus (Konta et al., 2020). The genus shares most similarities with

Diatrype, whereas they can be distinguished by the shape and size

of stromata (Konta et al., 2020).

Anthostoma Nitschke, Pyrenomyc. Germ. 1: 110, 1867.

MycoBank MB 224.

Type:Anthostoma decipiens (DC.) Nitschke, Pyrenomyc.

Germ. 1: 111, 1867.

Known distribution: Asia, Europe, North America, and South

America (Cash, 1952;Ahmad, 1969;Farr, 1973;Rappaz, 1995).

Notes: Both sexual and asexual morphs of the type species

Anthostoma decipiens were studied by Rappaz (1992, 1993).

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Zhu et al. Diatrypaceous Fungi in China

Phylogenetic analyses of sequence data also supported these

results (MP/ML/BI = 100/100/1) (Rocchi et al., 2010;Jaklitsch

et al., 2014). Species of Anthostoma own dark brown to dark,

globose to subglobose ascomata, cylindrical, prominent ostioles,

cylindrical to clavate asci with apically rounded to truncate

apices and a short pedicel and brown to black-brown ascospores

(Nitschke, 1867).

Cryptosphaeria Ces. & De Not., Comm. Soc. Crittog. Ital.

1(4): 231, 1863.

MycoBank MB 26092.

Type:Cryptosphaeria millepunctata Grev., Fl. Edin.: 360, 1824.

Known distribution: Asia, Europe, North America, Oceania,

and South Africa (Teodoro, 1937;Trouillas et al., 2015;

Jayawardena et al., 2018;Moyo et al., 2019).

Notes:Cryptosphaeria is widely accepted in Diatrypaceae

(Nitschke, 1867, 1870;Rappaz, 1989;Trouillas et al., 2010b,

2015). The genus comprises corticolous species and is

characterized by widely eﬀuse and poorly developed stromata,

often covered by a periderm, with separately emerging ostioles,

spindle-shaped, long pedicellate asci, and sub-olivaceous to

brown ascospores (Glawe, 1984;Rappaz, 1987).

Cryptovalsa Ces. & De Not. ex Fuckel, Jb. Nassau. Ver.

Naturk. 23–24: 212, 1870.

MycoBank MB 1340.

Type:Cryptovalsa protracta (Pers.) De Not.,

Hedwigia 2: 178, 1863.

Known distribution: Asia, Europe, North America, Oceania,

South Africa, and South America (Teodoro, 1937;Trouillas et al.,

2010b, 2011;Jayawardena et al., 2018;Moyo et al., 2019).

Notes:Cryptovalsa was established by Cesati and De

Notaris (1863) to accommodate C.protracta,C.ampelina,C.

nitschkei, and C.eﬀusa. The genus is characterized by eutypoid

stromata that are rather variable, when erumpent separately

diatrypelloid, often immersed in wood, but sometimes invading

bark tissues. Asci are cylindrical or clavate, polysporous, with

short or long pedicels. Ascospores are crowded, allantoid,

and yellowish (Spooner, 1981;Vasilyeva and Stephenson,

2005). Fifty-ﬁve species epithets of Cryptovalsa are listed

in Index Fungorum (2021), but only Cryptovalsa ampelina

has sequence data.

Diatrypasimilis J.J. Zhou & Kohlm., Mycologia

102(2): 432, 2010.

MycoBank MB 515026.

Type:Diatrypasimilis australiensis J.J. Zhou & Kohlm.,

Mycologia 102(2): 432, 2010.

Known distribution: Australia (Chalkley et al., 2010).

Notes:Diatrypasimilis was established to accommodate

D. australiensis as type species from mangroves based on

conventional taxonomic criteria and molecular phylogeny

by Chalkley et al. (2010). The genus is characterized by

carbonaceous, black stromata, 8-spored, cylindrical asci, and

ellipsoidal, dark brown ascospores with a germ slit.

Dothideovalsa Speg., Anal. Mus. Nac. B. Aires, Ser.

3(12): 414, 1909.

MycoBank MB 1697.

Type:Dothideovalsa tucumanensis Speg., Anal. Mus. Nac. B.

Aires, Ser. 3(12): 414, 1909.

Known distribution: Argentina, Brazil, and United States

(Rappaz, 1987;Hanlin, 1992).

Notes:Dothideovalsa was introduced to accommodate D.

diantherae,D.eutypoides,D.tucumanensis, and D.turnerae. At

present, only four species epithets of Dothideovalsa were listed

in Index Fungorum (2021). However, there was no available

sequence data in GenBank. Therefore, this genus is still doubtful

and needs further studies.

Endoxylina Romell, Bot. Notiser 1892: 173, 1892.

MycoBank MB 1814.

Type:Endoxylina stellulata Romell, Bot. Notiser: 173 (1892).

Known distribution: Asia, Europe, and North America (Tai,

1979;Ju et al., 1996).

Notes:Endoxylina was introduced and assigned to Diatrypales

(Current name: Xylariales ﬁde Kirk et al., 2008) without

assigning the familial position by Romell (1892). Based on

previous morphological literature and herbarium studies,

Hyde et al. (2017) transferred Endoxylina to Diatrypaceae.

The concept of genus Endoxylina is rather broad, and it

is characterized as having stromata of valsoid or eutypoid

conﬁgurations and 8-spored, long pedicellate, asci with J-,

apical ring, as well as uni- to triseptate, ascospores (Romell,

1892;Ju et al., 1996;Vasilyeva, 2010;Hyde et al., 2017).

At present, 21 species epithets of Endoxylina have been

described in Index Fungorum (2021), however, some of

these Endoxylina species have now been transferred and

synonymized with other genera (Ellis and Everhart, 1892;

Wehmeyer, 1975;Rappaz, 1987;Barr, 1993). Therefore,

the number of species recognized for this genus is

not well-established.

Eutypa Tul. & C. Tul., Select. Fung. Carpol. (Paris) 2: 52, 1863.

MycoBank MB 1950.

Type:Eutypa lata (Pers.) Tul. & C. Tul., Select. Fung. Carpol.

(Paris) 2: 56, 1863.

Known distribution: Asia, Europe, North America, Oceania,

South Africa, and South America (Ahmad, 1978;Rappaz, 1987;

Schmid-Heckel, 1988;Eriksson and Yue, 1998;Mulenko et al.,

2008;Moyo et al., 2019).

Notes: Species of Eutypa are the causal agents of dieback

of grapevine, apricots, and cherries (Trouillas and Gubler,

2004, 2010;Baumgartner et al., 2010;Camps et al., 2010;

Blanco-Ulate et al., 2013;Camps et al., 2014). The genus

is characterized by stromata which are irregular in shape,

as conﬂuent bumps, with conspicuous, scattered, roundish to

prominent ostioles on the host surface. Asci are 8-spored, clavate,

apically rounded to truncate, with indistinct apical rings and long

pedicels. Ascospores are allantoid to ellipsoidal, aseptate, and

pale yellowish.

Halocryptosphaeria Dayar., Devadatha, V.V. Sarma & K.D.

Hyde, Mycosphere 11(1): 136, 2020.

MycoBank MB 556800.

Type:Halocryptosphaeria bathurstensis (K.D. Hyde & Rappaz)

Dayar. & K.D. Hyde, Mycosphere 11(1): 136, 2020.

Known distribution: India (Dayarathne et al., 2020a).

Notes:Halocryptosphaeria was established by Dayarathne et al.

(2020a) to accommodate only one species, H. bathurstensisspecies.

Although Halocryptosphaeria has some morphological

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Zhu et al. Diatrypaceous Fungi in China

similarities with Cryptosphaeria, they can be easily distinguished

by phylogenetical analyses.

Halocryptovalsa Dayar. & K.D. Hyde, Cryptog. Mycol.

41(3): 49, 2020.

MycoBank MB 824308.

Type:Halocryptovalsa avicenniae (Abdel-Wahab, Bahkali

& E.B.G. Jones) Dayar. & K.D. Hyde, Cryptog. Mycol.

41(3): 50, 2020.

Known distribution: India, Saudi Arabia, and Thailand (Abdel-

Wahab et al., 2017;Dayarathne et al., 2020a,b).

Notes:Halocryptovalsa was established by Dayarathne et al.

(2020b) to accommodate species resembling Cryptovalsa from

marine environments namely Cryptovalsa avicenniae and a new

species Halocryptovalsa salicorniae. The genus is characterized

by poorly developed stromata and poly-spored asci, with a J-,

cylindrical, conspicuous apical or subapical ring. Ascospores are

hyaline or yellow-brown to brown, allantoid, with small, fat

globules at the end (Dayarathne et al., 2020b).

Halodiatrype Dayar. & K.D. Hyde, Phytotaxa 7(5): 617, 2016.

MycoBank MB 552254.

Type:Halodiatrype salinicola Dayar. & K.D. Hyde,

Mycosphere 7(5): 617, 2016.

Known distribution: Thailand (Dayarathne et al., 2016, 2020b).

Notes:Halodiatrype was established by Dayarathne et al.

(2016) to accommodate H.avicenniae and H.salinicola isolated

from mangroves. The characteristic of this genus is having

ascomata lacking stromatal tissues, 8-spored, cylindrical to

clavate, pedicellate asci, oblong to allantoid or sub-inequilateral,

larger ascospores with septa, and libertella-like asexual morphs,

which can easily identify Halodiatrype from other genera in

Diatrypaceae (Dayarathne et al., 2016; 2020b).

Leptoperidia Rappaz, Mycol. Helv. 2(3): 544, 1987.

MycoBank MB 25186.

Type:Leptoperidia macropunctata (Rehm) Rappaz, Mycol.

Helv. 2(3): 545, 1987.

Known distribution: Congo, Mexico, Philippines (Ellis and

Everhart, 1896;Rehm, 1913).

Notes:Leptoperidia was introduced to accommodate L.

applanata,L.asperrima,L.macropunctata, and L.triﬁda (Rappaz,

1987). The genus is characterized by relatively small stroma, asci

and ascospores, perithecia with very thin and slightly melanized

walls (Rappaz, 1987). At present, only four species epithets of

Leptoperidia are listed in Index Fungorum (2021). Sequence data

are unavailable in GenBank.

Libertella Desm., Annls Sci. Nat., Bot. 19: 275, 1830.

MycoBank MB 8769.

Type:Libertella betulina Desm., Annls Sci. Nat.,

Bot. 19: 276, 1830.

Known distribution: Asia, Europe, North America, Oceania,

South Africa, and South America (Doidge, 1950;Conners, 1967;

Ahmad, 1969;Farr, 1973;Sosnowski et al., 2007;Mulenko et al.,

2008).

Notes:Libertella was introduced by Desmazières (1830) to

accommodate L. betulina,L.faginea, and L.rosae. This genus was

mostly reported as the asexual morph of Diatrypella, however,

some species were reported as the asexual morph of Eutypa,

Eutypella,Diaporthe, and Polystigma (Kirk et al., 2001). The

genus is characterized by subcortical, erumpent and yellow to red

acervula conidiomata, and branched conidiophores that produce

hyaline, 1-celled, ﬁliform conidia (Barnett and Hunter, 1972;

Sutton, 1980;von Arx, 1981).

Monosporascus Pollack & Uecker, Mycologia 66(2): 348, 1974.

MycoBank MB 3260.

Type:Monosporascus cannonballus Pollack & Uecker,

Mycologia 66(2): 348, 1974.

Known distribution: Africa, Asia, Europe, North

America, and South America (Pande, 2008;Sales et al.,

2010;Chew-Madinaveitia et al., 2012;Salem et al., 2013;

Aleandri et al., 2017).

Notes:Monosporascus was introduced by Pollack and

Uecker (1974) with M.cannonballus as the type species. The

genus is characterized by pyriform asci and the formation

of one (rarely two) single large, sphaerical ascospores

(Pollack and Uecker, 1974).

Neoeutypella M. Raza, Q.J. Shang, Phook. & L. Cai, Fungal

Diversity 95: 167, 2019.

MycoBank MB 555373.

Type:Neoeutypella baoshanensis M. Raza, Q.J. Shang, Phook.

& L. Cai, Fungal Diversity 95: 168, 2019.

Known distribution: China and France

(Phookamsak et al., 2019).

Notes:Neoeutypella was introduced by Phookamsak et al.

(2019) to accommodate two fungal strains under the name

Eutypella caricae and a new strain isolated from Pinus armandii

(Pinaceae). Neoeutypella is characterized by carbonaceous

stromata, erumpent through host epidermis, producing yellow

pigments surrounding the stroma, 8-spored, spindle-shaped asci

with long pedicellate, and overlapping 1–3-seriate, allantoid,

aseptate, slightly or moderately curved ascospores, with a

libertella-like asexual morph (Phookamsak et al., 2019).

Pedumispora K.D. Hyde & E.B.G. Jones, Mycol. Res.

96(1): 78, 1992.

MycoBank MB 25433.

Type:Pedumispora rhizophorae K.D. Hyde & E.B.G. Jones,

Mycol. Res. 96: 78, 1992.

Known distribution: Australia, India, and Thailand (Hyde and

Jones, 1992;Pande, 2008;Dayarathne et al., 2020b).

Notes:Pedumispora was established by Hyde and

Jones (1992) to accommodate a taxon from mangrove

habitats. A phylogenetic study based on nuclear ITS

and LSU regions showed that the taxonomic position

of Pedumispora was in Diatrypaceae (Klaysuban et al.,

2014). The genus is characterized by 8-spored, fusiform,

pedicellate, unitunicate, apically truncate asci. Ascospores

are ﬁliform, mutli-septate, curved, longitudinally striate,

with tapering poles, with one or both ends crook-like

(Hyde and Jones, 1992).

Peroneutypa Berl., Icon. Fung. 3: 80, 1902.

MycoBank MB 3834.

Type:Peroneutypa bellula (Desm.) Berl., Icon.

Fung. 3: 81, 1902.

Known distribution: Asia, Europe, South Africa, and South

America (Teodoro, 1937;Moyo et al., 2018a,Castilla-Cayuman

et al., 2019;Moyo et al., 2019).

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Notes:Peroneutypa was introduced by Berlese (1902) to

accommodate P. bellula,P.corniculata, and P.heteracantha

without designating the type species. Rappaz (1987)

proposed P. bellula as the type species and synonymized

Peroneutypa under Eutypella. However, Carmarán et al. (2006)

resurrected Peroneutypa based on morphological characters and

phylogenetic. The genus is described as valsoid stromata, with

packed, long prominent necks, sessile to long pedicels, small asci

with truncate apices, and allantoid ascospores (Carmarán et al.,

2006, 2014;Senwanna et al., 2017;Shang et al., 2017).

Quaternaria Tul. & C. Tul., Select. Fung. Carpol. 2: 104, 1863.

MycoBank MB 4632.

Type:Quaternaria persoonii Tul. & C. Tul., Select. Fung.

Carpol. 2: 105, 1863.

Known distribution: Asia, Europe, and South America (Mujica

and Vergara, 1945;Munk, 1957;Srinivasulu and Sathe, 1970).

Notes:Quaternaria was introduced by Tulasne and Tulasne

(1863) and was typiﬁed by Q. persoonii.Clements and Shear

(1931) lectotypiﬁed the illegitimate name Q.quaternata to Q.

persoonii and considered Quaternaria as a synonym of Eutypella

(Tulasne and Tulasne, 1863). Based on molecular phylogeny and

the discussion of Gams (1994),Quaternaria was considered to

be an independent genus by Acero et al. (2004). The genus is

characterized by stromata was cryptosphaeroid in appearance

and developed within the bark parenchyma.

Rostronitschkia Fitzp., Mycologia 11(4): 165, 1919.

MycoBank MB 4793.

Type:Rostronitschkia nervincola Fitzp., Mycologia

11(4): 166, 1919.

Known distribution: Puerto Rico and Virgin Islands

(Stevenson, 1975).

Notes:Rostronitschkia was introduced to accommodate the

type species R.nervincola. At present, only the type species

was listed in Index Fungorum (2021). However, there were no

available sequence data and strains. Therefore, this genus is still

doubtful and needs to further study.

Key to genera of Diatrypaceae

1 Sexual morph absent................................................................ 2

1 Sexual morph present..............................................................3

2 Conidiomata acervuli, yellow to red....................... Libertella

2 Conidiomata pycnidial, brownish yellow, watery, bubble-

like................................................................................ Diatrype

3 Ascospores globose, fusiform, or oblong to ellipsoidal...... 4

3 Ascospores allantoid............................................................... 7

4 Ascospores fusiform, septate............................ Pedumispora

4 Ascospores aseptate................................................................. 5

5 Ascospores globose, with 1–2 spores in each ascus............

........................................................................... Monosporascus

5 Ascospores oblong to ellipsoidal, with 8 spores in each

ascus.......................................................................................... 6

6 Ascospores with a germ slit........................... Diatrypasimilis

6 Ascospores lacking a germ slit............................ Anthostoma

7 Asci with more than 8 spores................................................ 8

7 Asci with 8 spores.................................................................... 9

8 Stromata erumpent through host surface, discoid.......

................................................................................. Diatrypella

8 Stromata immersed in wood but sometimes invading bark

tissues, eutypoid..................................................................... 10

9 Perithecia outer surface lacking powdery entostroma..........

........................................................................... Allocryptovalsa

9 Perithecia outer surface coated with white, powdery

entostroma............................................................. Cryptovalsa

10 Habitats marine............................................ Halocryptovalsa

10 Habitats terrestrial................................................................. 11

11 Ascospores 0-1 septate.......................................................... 12

11 Ascospores uni- to triseptate................................ Endoxylina

12 Stromata semi-immersed to erumpent through the host

periderm (ectostromatic)..................................................... 13

12 Stromata deeply immersed in the host periderm

(entostromatic)....................................................................... 15

13 Asci lacking an apical ring................................. Halodiatrype

13 Asci with J-, cylindrical, conspicuous apical ring............. 14

14 Ascospores hyaline becoming yellowish at maturity........

................................................................................. Allodiatrype

14 Ascospores hyaline..................................................... Diatrype

15 Ascomata clustered, forming valsoid conﬁguration,

breaking through entostroma by short to long necks..... 16

15 Ascomata scattered, arranged in linear entostroma, with

short to long necks................................................................ 18

16 Entostromata immersed in the host, with individually

protruding necks at the center.......................... Quaternaria

16 Entostromata slightly raised on the host, with long

cylindrical, packed necks.................................................... 17

17 Hosts range wide.................................................... Eutypella

17 Host Pinus armandii.......................................... Neoeutypella

18 Ascomata forming very long necks, through the host

surface.................................................................. Peroneutypa

18 Ascomata forming short papilla protruding the host

surface................................................................................... 19

19 Peridium thin-walled, composed of a single layer

of melanized cells, diﬃcult to separate from the

entostroma......................................................... Leptoperidia

19 Peridium thick-walled, composed of not one distinct layers,

separating from entostroma................................................ 20

20 Asci cylindric-clavate, with pale yellow ascospores... Eutypa

20 Asci generally spindle-shaped, with sub-olivaceous to

brown ascospores.................................................................. 21

21 Peridium compose of two distinct layers... Cryptosphaeria

21 Peridium compose of three distinct layers........................

.................................................................... Halocryptosphaeria

Key to Diatrypaceae species on Betula spp.

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1 Sexual morph absent.................................... Diatrype betulae

1 Sexual morph present........................................................... 2

2 Asci containing 8 biseriate ascospores................................. 3

2 Asci polysporous..................................................................... 5

3 Apical ring amyloid................................... Diatrype undulata

3 Apical ring indistinguishable.................................................. 4

4 Size of asci more than 25 µm.......... Cryptosphaeria venusta

4 Size of asci less than 25 µm................... Diatrype platystoma

5 Stromata immersed in wood but sometimes invading bark

tissues, eutypoid...................................... Eutypella halseyana

5 Stromata erumpent through host surface, discoid.............. 6

6 Asci cylindrical....................................... Diatrypella favacea

6 Asci clavate to spindle shaped................................................ 7

7 More than 10 perithecia arranged irregularly..........

........................................................ Diatrypella shennongensis

7 Less than 10 perithecia arranged regularly......................... 8

8 More than one host.............................. Diatrypella betulicola

8 Only one host............................................................................ 9

9 Length of asci less than 140 µm........................... Diatrypella

betulae

9 Length of asci more than 140 µm.....................................

............................................................... Diatrypella hubeiensis

DISCUSSION

In this study, 21 isolates were identiﬁed as diatrypaceous

fungi from Betula albosinensis,B. davurica, and B. platyphylla

(Betulaceae), Castanea mollissima and Quercus mongolica

(Fagaceae), Juglans regia (Juglandaceae), and Morus alba

(Moraceae). Morpho-molecular analyses conﬁrmed that these

strains belong in four genera (viz.Allocryptovalsa,Diatrype,

Diatrypella, and Eutypella) including nine novel species (viz.

Allocryptovalsa castaneae,A. castaneicola,Diatrype betulae,

D. castaneicola,D. quercicola,Diatrypella betulae,Da. betulicola,

Da. hubeiensis, and Da. shennongensis) and two known species

(viz.Diatrypella favacea and Eutypella citricola). Eutypella

citricola was reported from Morus host for the ﬁrst time.

The generic concepts of Diatrypaceae have been unstable,

thus many species were transferred from one genus to another

(Phookamsak et al., 2019;Konta et al., 2020). The current

study revised the Diatrypaceae and accepted 23 genera in this

family (Supplementary Table 1). However, there only exist 18

genera in current phylogenetic analyses due to availability of

molecular data (Figure 1). In China, 11 genera and 62 species

belong in Diatrypaceae have been recorded (Supplementary

Table 5). However, 40 species (66.67%) do not have molecular

data until now.

Birch is of high economic, medicinal, and ornamental value.

Six species of Diatrypaceae were recorded from Betula spp.

with DNA sequences in the current study, including Diatrype

betulae,Diatrypella betulae,Da. betulicola,Da. favacea,Da.

hubeiensis, and Da. shennongensis. All species of Diatrypella 2

clade were isolated from Betula spp., except for Da. pulvinata

and Da. yunnanensis isolated from an unidentiﬁed plant. It

showed that many Diatrypella species may have obvious host

speciﬁcity. The other four species (viz.Cryptosphaeria venusta,

Diatrype platystoma,D. undulata, and Eutypella halseyana) of

Diatrypaceae were recorded from Betula spp. in China but

no materials with DNA sequences, including Cryptosphaeria

venusta,Diatrype platystoma,D. undulata, and Eutypella

halseyana (Teng, 1996;Vasilyeva and Ma, 2014). A morphological

key was provided to separate them in the current study.

The Allocryptovalsa species clustered within the same clade

as Eutypella species in our phylogenetic analyses (Figure 1;

Clade 12: Allocryptovalsa/Eutypella sensu lato). Allocryptovalsa

was introduced by Senwanna et al. (2017) which resembles

Cryptovalsa in morphology by having polysporous asci diﬀerent

from the 8-spored asci of Eutypella. The number of ascospores

per ascus (eight spores vs. multiple spores) has been used

traditionally to diﬀerentiate the genera of Diatrypaceae (Diatrype

vs. Diatrypella and Cryptovalsa vs. Eutypella). However, the

recent studies indicated that the polysporous ascus feature maybe

not signiﬁcant in Diatrypaceae (Acero et al., 2004;Trouillas et al.,

2011;Chacón et al., 2013;Liu et al., 2015). A thorough revision

is needed to resolve the problematic situation of Diatrypaceae,

which includes a mass of misidentiﬁed genus/species, as a result

of the unstable phylogenetic frame with type materials. Therefore,

it seems to better if the future work could treate the strains from

clade 12 into one genus Allocryptovalsa.

The strains of Diatrype formed a clade with high support

values (Figure 1; Clade 09: Diatrype sensu stricto). However,

in this clade, some strains of Diatrypella (Da. quercina,Da.

iranensis, and Da. macrospora) were placed between Diatrype

species. Acero et al. (2004) reported that Cryptosphaeria,

Diatrype,Diatrypella,Eutypa, and Eutypella were polyphyletic

and confused probably due to lack of tub2 gene sequences

or misidentiﬁed species. However, these ﬁve genera are still

polyphyletic within the family from previous studies (de Almeida

et al., 2016;Shang et al., 2017;Mehrabi et al., 2019;Dayarathne

et al., 2020a,b;Konta et al., 2020) based on the ITS and

tub2 sequences data. Allodiatrype and Halodiatrype reported

their morphological resemblance to Diatrype (Dayarathne et al.,

2016;Konta et al., 2020). However, asci of Halodiatrype lack

an apical ring, the asci of Allodiatrype and Diatrype have

J-, cylindrical, conspicuous apical ring (Dayarathne et al.,

2016;Maharachchikumbura et al., 2016;Konta et al., 2020).

Additionally, Diatrype can be diﬀerentiated from Allodiatrype

by the color of ascospores. The ascospores of Diatrype are

hyaline becoming yellowish ascospores at maturity, whereas the

ascospores of Allodiatrype are hyaline (Maharachchikumbura

et al., 2016;Konta et al., 2020). Though Diatrype and Diatrypella

are not polyphyletic any more, some strains of Diatrype and

Diatrypella species still need further study.

In some cases, some strains of Diatrype species

(D. brunneospora and D. palmicola), Diatrypella species

(Da. banksiae), Eutypa species (E. ﬂavovirens and E. guttulata)

and Eutypalla species (E. cearensis) formed distinct lineages

within Diatrypaceae (Figure 1;Incertae sedis). Diatrypella

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Zhu et al. Diatrypaceous Fungi in China

banksiae is closely related to the genus Neoeutypella

with high support values from the phylogenetic analyses

(MP/ML/BI = 98/100/1) (Figure 1), which produced an

asexual morph (Crous et al., 2016;Phookamsak et al., 2019).

Nevertheless, Diatrypella banksiae having spindle-shaped conidia

can be easily diﬀerentiated from Neoeutypella baoshanensis

having ﬁliform conidia (Crous et al., 2016;Phookamsak et al.,

2019). In the future study, sexual morph of Diatrypella banksiae

also remains to be studied, because the asexual morphs of

Diatrypaceae are not generally useful in separating species

(Glawe and Rogers, 1982, 1984;Rappaz, 1987;Konta et al.,

2020). Therefore, Diatrypella banksiae probably belongs to the

genus Neoeutypella. Further study of the relationship between

Neoeutypella and Diatrypella banksiae is needed.

Eutypa guttulata is closely related to the genus Halodiatrype

and Pedumispora from the phylogenetic analyses (Figure 1) as a

basal branch. Eutypa guttulata can be distinguished by fusiform

ascospores, whereas Pedumispora has allantoid ascospores (Hyde

and Jones, 1992). It is also diﬀerent from Halodiatrype because

of lacking an apical ring (Dayarathne et al., 2016). Diatrype

brunneospora is closed to Eutypa guttulata (Figure 1), which is

morphologically similar to members of Eutypa spp. (Trouillas

et al., 2010b). Therefore, the assignment of this isolate to the

genus Diatrype may require reconsideration in the future. Eutypa

ﬂavovirens is closely related to the genus Cryptosphaeria with no

support (Figure 1). Diatrypella can be distinguished by hyaline to

subhyaline, rarely pale olivaceous ascospores, whereas Diatrype

palmicola has pale yellowish to pale brown ascospores at maturity

(Liu et al., 2015). For those species, the assignment of these

isolates still remain unclear, which may require reconsideration

in the future.

Eutypa microasca appeared in a strongly supported clade

along with two Peroneutypa species with fusiform asci in

Figure 1 (Clade 19: Peroneutypa). Carmarán et al. (2006)

suggested that the morphology of the ascus could explain

the phylogenetic relationships within Diatrypaceae better than

stromata, although our study indicates that it is not entirely

supported in Peroneutypa. The phylogenetic signal of the ascus

shape and the phylogenetic placement of Eutypa microasca

should be further tested in future study.

DATA AVAILABILITY STATEMENT

The datasets presented in this study can be found in online

repositories. The names of the repository/repositories and

accession number(s) can be found in the article/Supplementary

Material.

AUTHOR CONTRIBUTIONS

XF and CT conceived and designed the experiments. HZ,

MP, and RM performed the experiment. HZ and MP

analyzed the data. NW and DD provided some materials

and polished the language. HZ wrote the manuscript. XF

revised and approved the ﬁnal version of the manuscript.

All authors contributed extensively to the work presented

in the manuscript.

FUNDING

This study was ﬁnanced by the National Natural Science

Foundation of China (No. 31670647) and the Fundamental

Research Funds for the Central Universities (No. 2019ZY23).

ACKNOWLEDGMENTS

The authors want to thank the Experimental Teaching

Centre (College of Forestry, Beijing Forestry University)

for providing installed scientiﬁc equipment during the

whole process. NW and DD thank Province Universities

of the Diversity and Ecological Adaptive Evolution for

Animals and Plants on the Yun-Gui Plateau, the National

Natural Science Foundation of China, and the Thousand

Talents Plan, Youth Project of Yunnan Provinces. NW

gratefully acknowledges the State Key Laboratory of

Functions and Applications of Medicinal Plants, Guizhou

Medical University; Science and Technology Department of

Guizhou Province.

SUPPLEMENTARY MATERIAL

The Supplementary Material for this article can be found

online at: https://www.frontiersin.org/articles/10.3389/fmicb.

2021.646262/full#supplementary-material

Supplementary Table 1 | Placement of genera in Diatrypaceae by

different authors.

Supplementary Table 2 | Isolates and GenBank accession numbers used in the

phylogenetic analyses of Diatrypaceae.

Supplementary Table 3 | Synopsis of species of Allocryptovalsa.

Supplementary Table 4 | Synopsis of species of Diatrypella from Betula spp.

Supplementary Table 5 | Distribution of diatrypaceous fungi on plants in China.

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Frontiers in Microbiology | www.frontiersin.org 26 May 2021 | Volume 12 | Article 646262

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47854

Conﬂict of Interest: The authors declare that the research was conducted in the

absence of any commercial or ﬁnancial relationships that could be construed as a

potential conﬂict of interest.

is an open-access article distributed under the terms of the Creative Commons

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is permitted, provided the original author(s) and the copyright owner(s) are credited

and that the original publication in this journal is cited, in accordance with accepted

academic practice. No use, distribution or reproduction is permitted which does not

comply with these terms.

Frontiers in Microbiology | www.frontiersin.org 27 May 2021 | Volume 12 | Article 646262

Taxonomy and pathogenicity of fungi associated with oak decline in northern and central Zagros forests of Iran with emphasis on coelomycetous species

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Apr 2024

Oak decline is a complex disorder that seriously threatens the survival of Zagros forests. In an extensive study on taxonomy and pathology of fungi associated with oak decline in the central and northern part of Zagros forests, 462 fungal isolates were obtained from oak trees showing canker, gummosis, dieback, defoliation, and partial or total death symptoms. Based on inter-simple sequence repeat (ISSR) fingerprinting patterns, morphological characteristics, and sequences of ribosomal DNA (28S rDNA and ITS) and protein coding loci (acl1, act1, caM, tef-1α, rpb1, rpb2, and tub2), 24 fungal species corresponding to 19 genera were characterized. Forty percent of the isolates were placed in eight coelomycetous species from seven genera, namely, Alloeutypa, Botryosphaeria, Cytospora, Didymella, Gnomoniopsis, Kalmusia, and Neoscytalidium. Of these, four species are new to science, which are introduced here as taxonomic novelties: Alloeutypa iranensis sp. nov., Cytospora hedjaroudei sp. nov., Cytospora zagrosensis sp. nov., and Gnomoniopsis quercicola sp. nov. According to pathogenicity trials on leaves and stems of 2-year-old Persian oak (Quercus brantii) seedlings, Alternaria spp. (A. alternata, A. atra, and A. contlous), Chaetomium globosum, and Parachaetomium perlucidum were recognized as nonpathogenic. All coelomycetous species were determined as pathogenic in both pathogenicity trials on leaves and seedling stems, of which Gnomoniopsis quercicola sp. nov., Botryosphaeria dothidea, and Neoscytalidium dimidiatum were recognized as the most virulent species followed by Biscogniauxia rosacearum.

Fungal diversity notes 1512–1610: taxonomic and phylogenetic contributions on genera and species of fungal taxa

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Full-text available

Feb 2023

This article is the 14th in the Fungal Diversity Notes series, wherein we report 98 taxa distributed in two phyla, seven classes, 26 orders and 50 families which are described and illustrated. Taxa in this study were collected from Australia, Brazil, Burkina Faso, Chile, China, Cyprus, Egypt, France, French Guiana, India, Indonesia, Italy, Laos, Mexico, Russia, Sri Lanka, Thailand, and Vietnam. There are 59 new taxa, 39 new hosts and new geographical distributions with one new combination. The 59 new species comprise Angustimassarina kunmingense, Asterina lopi, Asterina brigadeirensis, Bartalinia bidenticola, Bartalinia caryotae, Buellia pruinocalcarea, Coltricia insularis, Colletotrichum flexuosum, Colletotrichum thasutense, Coniochaeta caraganae, Coniothyrium yuccicola, Dematipyriforma aquatic, Dematipyriforma globispora, Dematipyriforma nilotica, Distoseptispora bambusicola, Fulvifomes jawadhuvensis, Fulvifomes malaiyanurensis, Fulvifomes thiruvannamalaiensis, Fusarium purpurea, Gerronema atrovirens, Gerronema flavum, Gerronema keralense, Gerronema kuruvense, Grammothele taiwanensis, Hongkongmyces changchunensis, Hypoxylon inaequale, Kirschsteiniothelia acutisporum, Kirschsteiniothelia crustaceum, Kirschsteiniothelia extensum, Kirschsteiniothelia septemseptatum, Kirschsteiniothelia spatiosum, Lecanora immersocalcarea, Lepiota subthailandica, Lindgomyces guizhouensis, Marthe asmius pallidoaurantiacus, Marasmius tangerinus, Neovaginatispora mangiferae, Pararamichloridium aquisubtropicum, Pestalotiopsis piraubensis, Phacidium chinaum, Phaeoisaria goiasensis, Phaeoseptum thailandicum, Pleurothecium aquisubtropicum, Pseudocercospora vernoniae, Pyrenophora verruculosa, Rhachomyces cruralis, Rhachomyces hyperommae, Rhachomyces magrinii, Rhachomyces platyprosophi, Rhizomarasmius cunninghamietorum, Skeletocutis cangshanensis, Skeletocutis subchrysella, Sporisorium anadelphiae-leptocomae, Tetraploa dashaoensis, Tomentella exiguelata, Tomentella fuscoaraneosa, Tricholomopsis lechatii, Vaginatispora flavispora and Wetmoreana blastidiocalcarea. The new combination is Torula sundara. The 39 new records on hosts and geographical distribution comprise Apiospora guiyangensis, Aplosporella artocarpi, Ascochyta medicaginicola, Astrocystis bambusicola, Athelia rolfsii, Bambusicola bambusae, Bipolaris luttrellii, Botryosphaeria dothidea, Chlorophyllum squamulosum, Colletotrichum aeschynomenes, Colletotrichum pandanicola, Coprinopsis cinerea, Corylicola italica, Curvularia alcornii, Curvularia senegalensis, Diaporthe foeniculina, Diaporthe longicolla, Diaporthe phaseolorum, Diatrypella quercina, Fusarium brachygibbosum, Helicoma aquaticum, Lepiota metulispora, Lepiota pongduadensis, Lepiota subvenenata, Melanconiella meridionalis, Monotosporella erecta, Nodulosphaeria digitalis, Palmiascoma gregariascomum, Periconia byssoides, Periconia cortaderiae, Pleopunctum ellipsoideum, Psilocybe keralensis, Scedosporium apiospermum, Scedosporium dehoogii, Scedosporium marina, Spegazzinia deightonii, Torula fici, Wiesneriomyces laurinus and Xylaria venosula. All these taxa are supported by morphological and multigene phylogenetic analyses. This article allows the researchers to publish fungal collections which are important for future studies. An updated, accurate and timely report of fungus-host and fungus-geography is important. We also provide an updated list of fungal taxa published in the previous fungal diversity notes. In this list, erroneous taxa and synonyms are marked and corrected accordingly.

Additions to Diatrypaceae (Xylariales): Novel Taxa and New Host Associations

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Full-text available

Nov 2023
J. Fungi

Diatrypaceae members have a broad distribution and are commonly found on decaying wood. Despite taxonomic and morphological challenges within this group, there has been a growing interest in Diatrypaceae in recent years. The dead branches of several plant hosts with fungal fruiting bodies were collected from Doi Tung National Park, Chiang Rai, and the Narathiwat Provinces in Thailand. Their morphological characteristics, coupled with a molecular phylogeny of combined ITS and tub2 sequence data, were used to introduce two novel Allodiatrype species (A. dalbergiae and A. eleiodoxae) and one new Melanostictus species (M. chiangraiensis). Moreover, four new host records, Diatrypella heveae, D. major, Melanostictus thailandicus, and Paraeutypella citricola on Microcos paniculata, Nayariophyton zizyphifolium, Dalbergia cultrata, and M. paniculata, respectively, as well as a new geographical record of D. major are reported. This research provides detailed descriptions of macro- and microcharacteristics, coupled with a phylogenetic tree for the newly introduced species and host records. The morphological features of Allodiatrype and Melanostictus are listed in the synoptic table.

Chemical structures, biological activities, and biosynthetic analysis of secondary metabolites of the Diatrypaceae family: A comprehensive review

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Apr 2024

Fungal diversity notes 1611–1716: taxonomic and phylogenetic contributions on fungal genera and species emphasis in south China

Article

Full-text available

Nov 2023

This article is the 15th contribution in the Fungal Diversity Notes series, wherein 115 taxa from three phyla, nine classes, 28 orders, 48 families, and 64 genera are treated. Fungal taxa described and illustrated in the present study include a new family, five new genera, 61 new species, five new combinations, one synonym, one new variety and 31 records on new hosts or new geographical distributions. Ageratinicolaceae fam. nov. is introduced and accommodated in Pleosporales. The new genera introduced in this study are Ageratinicola, Kevinia, Pseudomultiseptospora (Parabambusicolaceae), Marasmiellomycena, and Vizzinia (Porotheleaceae). Newly described species are Abrothallus altoandinus, Ageratinicola kunmingensis, Allocryptovalsa aceris, Allophoma yuccae, Apiospora cannae, A. elliptica, A. pallidesporae, Boeremia wisteriae, Calycina papaeana, Clypeococcum lichenostigmoides, Coniochaeta riskali-shoyakubovii, Cryphonectria kunmingensis, Diaporthe angustiapiculata, D. campylandrae, D. longipapillata, Diatrypella guangdongense, Dothiorella franceschinii, Endocalyx phoenicis, Epicoccum terminosporum, Fulvifomes karaiensis, F. pannaensis, Ganoderma ghatensis, Hysterobrevium baoshanense, Inocybe avellaneorosea, I. lucida, Jahnula oblonga, Kevinia lignicola, Kirschsteiniothelia guangdongensis, Laboulbenia caprina, L. clavulata, L. cobiae, L. cosmodisci, L. nilotica, L. omalii, L. robusta, L. similis, L. stigmatophora, Laccaria rubriporus, Lasiodiplodia morindae, Lyophyllum agnijum, Marasmiellomycena pseudoomphaliiformis, Melomastia beihaiensis, Nemania guangdongensis, Nigrograna thailandica, Nigrospora ficuum, Oxydothis chinensis, O. yunnanensis, Petriella thailandica, Phaeoacremonium chinensis, Phialocephala chinensis, Phytophthora debattistii, Polyplosphaeria nigrospora, Pronectria loweniae, Seriascoma acutispora, Setoseptoria bambusae, Stictis anomianthi, Tarzetta tibetensis, Tarzetta urceolata, Tetraploa obpyriformis, Trichoglossum beninense, and Tricoderma pyrrosiae. We provide an emendation for Urnula ailaoshanensis Agaricus duplocingulatoides var. brevisporus introduced as a new variety based on morphology and phylogeny.

Fungal Planet description sheets: 1478–1549

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Jun 2023

Fungal Planet Description Sheets: 1478–1549

Article

Full-text available

Jun 2023
Persoonia

Novel species of fungi described in this study include those from various countries as follows: Australia, Aschersonia mackerrasiae on whitefly, Cladosporium corticola on bark of Melaleuca quinquenervia, Penicillium nudgee from soil under Melaleuca quinquenervia, Pseudocercospora blackwoodiae on leaf spot of Persoonia falcata, and Pseudocercospora dalyelliae on leaf spot of Senna alata. Bolivia, Aspicilia lutzoniana on fully submersed siliceous schist in high-mountain streams, and Niesslia parviseta on the lower part and apothecial discs of Erioderma barbellatum onatwig. Brazil, Cyathus bonsai on decaying wood, Geastrum albofibrosum from moist soil with leaf litter, Laetiporus pratigiensis on a trunk of a living unknown hardwood tree species, and Scytalidium synnematicum on dead twigs of unidentified plant. Bulgaria, Amanita abscondita on sandy soil in a plantation of Quercus suber. Canada, Penicillium acericola on dead bark of Acer saccharum, and Penicillium corticola on dead bark of Acer saccharum. China, Colletotrichum qingyuanense on fruit lesion of Capsicum annuum. Denmark, Helminthosphaeria leptospora on corticioid Neohypochnicium cremicolor. Ecuador (Galapagos), Phaeosphaeria scalesiae on Scalesia sp. Finland, Inocybe jacobssonii on calcareouss oils in dry forests and park habitats. France, Cortinarius rufomyrrheus on sandy soil under Pinus pinaster, and Periconia neominutissima on leaves of Poaceae. India, Coprinopsis fragilis on decaying bark of logs, Filoboletus keralensis on unidentified woody substrate, Penicillium sankaranii from soil, Physisporinus tamilnaduensis on the trunk of Azadirachta indica, and Poronia nagaraholensis on elephant dung. Iran, Neosetophoma fic on infected leaves of Ficus elastica. Israel, Cnidariophoma eilatica (incl. Cnidariophoma gen. nov.) from Stylophora pistillata. Italy, Lyophyllum obscurum on acidic soil. Namibia, Aureobasidium faidherbiae on dead leaf of Faidherbia albida, and Aureobasidium welwitschiae on dead leaves of Welwitschia mirabilis. Netherlands, Gaeumannomycella caricigena on dead culms of Carex elongata, Houtenomyces caricicola (incl. Houtenomyces gen. nov.) on culms of Carex disticha, Neodacampia ulmea (incl. Neodacampia gen. nov.) on branch of Ulmus laevis, Niesslia phragmiticola on dead standing culms of Phragmites australis, Pseudopyricularia caricicola on culms of Carex disticha, and Rhodoveronaea nieuwwulvenica on dead bamboo sticks. Norway, Arrhenia similis half-buried and moss-covered pieces of rotting wood in grass-grownpath. Pakistan, Mallocybe ahmadii on soil. Poland, Beskidomyces laricis (incl. Beskidomyces gen. nov.) from resin of Larix decidua ssp. polonica, Lapidomyces epipinicola from sooty mould community on Pinus nigra, and Leptographium granulatum from a gallery of Dendroctonus micans on Picea abies. Portugal, Geoglossum azoricum on mossy areas of laurel forest areas planted with Cryptomeria japonica, and Lunasporangiospora lusitanica from a biofilm covering a bio deteriorated limestone wall. Qatar, Alternaria halotolerans from hypersaline sea water, and Alternaria qatarensis from water sample collected from hypersaline lagoon. South Africa, Alfaria thamnochorti on culm of Thamnochortus fraternus, Knufia aloeicola on Aloe gariepensis, Muriseptatomyces restionacearum (incl.Muriseptatomyces gen. nov.) on culms of Restionaceae, Neocladosporium arctotis on nest of cases of bagworm moths(Lepidoptera, Psychidae) on Arctotis auriculata, Neodevriesia scadoxi on leaves of Scadoxus puniceus, Paraloratospora schoenoplecti on stems of Schoenoplectus lacustris, Tulasnella epidendrea from the roots of Epidendrum × obrienianum, and Xenoidriella cinnamomi (incl. Xenoidriella gen. nov.) on leaf of Cinnamomum camphora. South Korea, Lemonniera fraxinea on decaying leaves of Fraxinus sp. frompond. Spain, Atheniella lauri on the bark of fallen trees of Laurus nobilis, Halocryptovalsa endophytica from surface-sterilised, asymptomatic roots of Salicornia patula, Inocybe amygdaliolens on soil in mixed forest, Inocybe pityusarum on calcareous soil in mixed forest, Inocybe roseobulbipes on acidic soils, Neonectria borealis from roots of Vitis berlandieri × Vitis rupestris, Sympoventuria eucalyptorum on leaves of Eucalyptus sp., and Tuber conchae fromsoil. Sweden, Inocybe bidumensis on calcareous soil. Thailand, Cordyceps sandindaengensis on Lepidoptera pupa, buried in soil, Ophiocordyceps kuchinaraiensis on Coleoptera larva, buried in soil, and Samsoniella winandae on Lepidoptera pupa, buriedinsoil. Taiwan region (China), Neophaeosphaeria livistonae on dead leaf of Livistona rotundifolia. Türkiye, Melanogaster anatolicus on clay loamy soils. UK, Basingstokeomyces allii (incl. Basingstokeomyces gen. nov.) on leaves of Allium schoenoprasum. Ukraine, Xenosphaeropsis corni on recently dead stem of Cornus alba. USA, Nothotrichosporon aquaticum (incl. Nothotrichosporon gen. nov.) from water, and Periconia philadelphiana from swab of coil surface. Morphological and culture characteristics for these new taxa are supported by DNA barcodes.

Over the footprints of Italian mycology with emphasis on plant-associated Ascomycota

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Full-text available

May 2023

plant-associated Italian Ascomycota

Taxonomic and phylogenetic contributions to Diatrypaceae from southeastern Tibet in China

Article

Full-text available

Mar 2023

In this study, we investigated the diversity of diatrypaceous fungi from southeastern Tibet in China. The phylogenetic analyses were carried out based on ITS and β-tubulin sequences of 75 taxa of Diatrypaceae from around the world. Based on a combination of morphological features and molecular evidence, a new genus-Alloeutypa, with three new species-A. milinensis, Diatrype linzhiensis, and Eutypella motuoensis, and a new combination-A. flavovirens, were revealed by the materials in China. Alloeutypa is characterized by stromatal interior olivaceous buff, stromata producing well-developed discrete, and ascospores allantoid, subhyaline. These characteristics separate the new genus from the similar genus Eutypa. Comprehensive morphological descriptions, illustrations, and a phylogenetic tree to show the placement of new taxa are provided. All novelties described herein are morphologically illustrated and phylogeny investigated to better integrate taxa into the higher taxonomic framework and infer their phylogenetic relationships as well as establish new genera and species. Our results indicate that the diatrypaceous fungi harbor higher species diversity in China.

Diversity, morphology, and molecular phylogeny of Diatrypaceae from southern China

Article

Full-text available

Apr 2023

During an investigation of Diatrypaceae from southern China, 10 xylariales-like taxa have been collected. Morphological and multi-gene analyses confirmed that these taxa reside in Diatrypaceae and represent eight novel taxa and two new records belonging to six genera (viz ., Allocryptovalsa, Diatrype, Diatrypella, Paraeutypella, Peroneutypa , and Vasilyeva gen. nov.). Vasilyeva gen. nov. was proposed to accommodate Vasilyeva cinnamomi sp. nov. Among the other collections, seven new species were introduced ( viz ., Diatrype camelliae-japonicae sp. nov., Diatrype rubi sp. nov., Diatrypella guiyangensis sp. nov., Diatrypella fatsiae-japonicae sp. nov., Paraeutypella subguizhouensis sp. nov., Peroneutypa hainanensis sp. nov., and Peroneutypa qianensis sp. nov.), while two were reported as new records from China ( Allocryptovalsa rabenhorstii and Diatrype enteroxantha ). For Diatrypaceae , the traditional taxonomic approach based on morphology may not be applicable.

Modern Taxonomic Approaches to Identifying Diatrypaceous Fungi from Marine Habitats, with a Novel Genus Halocryptovalsa Dayarathne & K.D.Hyde, Gen. Nov.

Article

Full-text available

Mar 2020

Marine diatrypaceous fungi are associated with many host species, including Avicennia spp., Halosarceia halocnemoides (Nees) Paul G.Wilson, Kandelia candel (L.) Druce, Salicornia sp., Rhizophora apiculata Blume and Suaeda fruticosa (L.) Forssk. Accurate morphological identification of individual taxa of this family is difficult, as previous classifications generally lack sequence data. We provide a detailed account of all known marine diatrypaceous taxa and determine their phylogenetic and morphological relationships. A novel genus Halocryptovalsa Dayarathne & K.D.Hyde, gen. nov. is established to accommodate Halocryptovalsa salicorniae Dayarathne & K.D.Hyde, sp. nov., and Halocryptovalsa avicenniae, comb. nov. (synonym of Cryptovalsa avicenniae). Three novel species Cryptosphaeria avicenniae Devadatha & V.V.Sarma, sp. nov., Cryptosphaeria halophila Dayarathne & K.D.Hyde, sp. nov., and Diatrype mangrovei Dayarathne & K.D.Hyde, sp. nov., are introduced from mangroves. Halodiatrype is amended with asexual morph characters.

Outline of Fungi and fungus-like taxa

Article

Full-text available

Mar 2020

This article provides an outline of the classification of the kingdom Fungi (including fossil fungi. i.e. dispersed spores, mycelia, sporophores, mycorrhizas). We treat 19 phyla of fungi. These are Aphelidiomycota, Ascomycota, Basidiobolomycota, Basidiomycota, Blastocladiomycota, Calcarisporiellomycota, Caulochytriomycota, Chytridiomycota, Entomophthoromycota, Entorrhizomycota, Glomeromycota, Kickxellomycota, Monoblepharomycota, Mortierellomycota, Mucoromycota, Neocallimastigomycota, Olpidiomycota, Rozellomycota and Zoopagomycota. The placement of all fungal genera is provided at the class-, order- and family-level. The described number of species per genus is also given. Notes are provided of taxa for which recent changes or disagreements have been presented. Fungus-like taxa that were traditionally treated as fungi are also incorporated in this outline (i.e. Eumycetozoa, Dictyosteliomycetes, Ceratiomyxomycetes and Myxomycetes). Four new taxa are introduced: Amblyosporida ord. nov. Neopereziida ord. nov. and Ovavesiculida ord. nov. in Rozellomycota, and Protosporangiaceae fam. nov. in Dictyosteliomycetes. Two different classifications (in outline section and in discussion) are provided for Glomeromycota and Leotiomycetes based on recent studies. The phylogenetic reconstruction of a four-gene dataset (18S and 28S rRNA, RPB1, RPB2) of 433 taxa is presented, including all currently described orders of fungi.

Fungal diversity notes 1151–1276: taxonomic and phylogenetic contributions on genera and species of fungal taxa

Article

Full-text available

Mar 2020

Fungal diversity notes is one of the important journal series of fungal taxonomy that provide detailed descriptions and illustrations of new fungal taxa, as well as providing new information of fungal taxa worldwide. This article is the 11th contribution to the fungal diversity notes series, in which 126 taxa distributed in two phyla, six classes, 24 orders and 55 families are described and illustrated. Taxa in this study were mainly collected from Italy by Erio Camporesi and also collected from China, India and Thailand, as well as in some other European, North American and South American countries. Taxa described in the present study include two new families, 12 new genera, 82 new species, five new combinations and 25 new records on new hosts and new geographical distributions as well as sexual-asexual reports. The two new families are Eriomycetaceae (Dothideomycetes, family incertae sedis) and Fasciatisporaceae (Xylariales, Sordariomycetes). The twelve new genera comprise Bhagirathimyces (Phaeosphaeriaceae), Camporesiomyces (Tubeufiaceae), Eriocamporesia (Cryphonectriaceae), Eriomyces (Eriomycetaceae), Neomonodictys (Pleurotheciaceae), Paraloratospora (Phaeosphaeriaceae), Paramonodictys (Parabambusicolaceae), Pseudoconlarium (Diaporthomycetidae, genus incertae sedis), Pseudomurilentithecium (Lentitheciaceae), Setoapiospora (Muyocopronaceae), Srinivasanomyces (Vibrisseaceae) and Xenoanthostomella (Xylariales, genera incertae sedis). The 82 new species comprise Acremonium chiangraiense, Adustochaete nivea, Angustimassarina camporesii, Bhagirathimyces himalayensis, Brunneoclavispora camporesii, Camarosporidiella camporesii, Camporesiomyces mali, Camposporium appendiculatum, Camposporium multiseptatum, Camposporium septatum, Canalisporium aquaticium, Clonostachys eriocamporesiana, Clonostachys eriocamporesii, Colletotrichum hederiicola, Coniochaeta vineae, Conioscypha verrucosa, Cortinarius ainsworthii, Cortinarius aurae, Cortinarius britannicus, Cortinarius heatherae, Cortinarius scoticus, Cortinarius subsaniosus, Cytospora fusispora, Cytospora rosigena, Diaporthe camporesii, Diaporthe nigra, Diatrypella yunnanensis, Dictyosporium muriformis, Didymella camporesii, Diutina bernali, Diutina sipiczkii, Eriocamporesia aurantia, Eriomyces heveae, Ernakulamia tanakae, Falciformispora uttaraditensis, Fasciatispora cocoes, Foliophoma camporesii, Fuscostagonospora camporesii, Helvella subtinta, Kalmusia erioi, Keissleriella camporesiana, Keissleriella camporesii, Lanspora cylindrospora, Loratospora arezzoensis, Mariannaea atlantica, Melanographium phoenicis, Montagnula camporesii, Neodidymelliopsis camporesii, Neokalmusia kunmingensis, Neoleptosporella camporesiana, Neomonodictys muriformis, Neomyrmecridium guizhouense, Neosetophoma camporesii, Paraloratospora camporesii, Paramonodictys solitarius, Periconia palmicola, Plenodomus triseptatus, Pseudocamarosporium camporesii, Pseudocercospora maetaengensis, Pseudochaetosphaeronema kunmingense, Pseudoconlarium punctiforme, Pseudodactylaria camporesiana, Pseudomurilentithecium camporesii, Pseudotetraploa rajmachiensis, Pseudotruncatella camporesii, Rhexocercosporidium senecionis, Rhytidhysteron camporesii, Rhytidhysteron erioi, Septoriella camporesii, Setoapiospora thailandica, Srinivasanomyces kangrensis, Tetraploa dwibahubeeja, Tetraploa pseudoaristata, Tetraploa thrayabahubeeja, Torula camporesii, Tremateia camporesii, Tremateia lamiacearum, Uzbekistanica pruni, Verruconis mangrovei, Wilcoxina verruculosa, Xenoanthostomella chromolaenae and Xenodidymella camporesii. The five new combinations are Camporesiomyces patagoniensis, Camporesiomyces vaccinia, Camposporium lycopodiellae, Paraloratospora gahniae and Rhexocercosporidium microsporum. The 22 new records on host and geographical distribution comprise Arthrinium marii, Ascochyta medicaginicola, Ascochyta pisi, Astrocystis bambusicola, Camposporium pellucidum, Dendryphiella phitsanulokensis, Diaporthe foeniculina, Didymella macrostoma, Diplodia mutila, Diplodia seriata, Heterosphaeria patella, Hysterobrevium constrictum, Neodidymelliopsis ranunculi, Neovaginatispora fuckelii, Nothophoma quercina, Occultibambusa bambusae, Phaeosphaeria chinensis, Pseudopestalotiopsis theae, Pyxine berteriana, Tetraploa sasicola, Torula gaodangensis and Wojnowiciella dactylidis. In addition, the sexual morphs of Dissoconium eucalypti and Phaeosphaeriopsis pseudoagavacearum are reported from Laurus nobilis and Yucca gloriosa in Italy, respectively. The holomorph of Diaporthe cynaroidis is also reported for the first time.

Refined families of Sordariomycetes

Article

Full-text available

Feb 2020

This is a continuation of the papers “Towards a classification of Sordariomycetes” (2015) and “Families of Sordariomycetes” (2016) in which we compile a treatment of the class Sordariomycetes. The present treatment is needed as our knowledge has rapidly increased, from 32 orders, 105 families and 1331 genera in 2016, to 45 orders, 167 families and 1499 genera (with 308 genera incertae sedis) at the time of publication. In this treatment we provide notes on each order, families and short notes on each genus. We provide up-to-date DNA based phylogenies for 45 orders and 163 families. Three new genera and 16 new species are introduced with illustrations and descriptions, while 23 new records and three new species combinations are provided. We also list 308 taxa in Sordariomycetes genera incertae sedis. For each family we provide general descriptions and illustrate the type genus or another genus, the latter where the placement has generally been confirmed with molecular data. Both the sexual and asexual morphs representative of a family are illustrated where available. Notes on ecological and economic considerations are also given.

A new genus Allodiatrype, five new species and a new host record of diatrypaceous fungi from palms (Arecaceae)

Article

Full-text available

Jan 2020

Diatrypaceous fungi on palms (Arecaceae) in Thailand were collected and identified based on morphological characteristics as well as combined DNA sequence analyses (ITS and TUB2). One new genus Allodiatrype, and five new species, Allocryptovalsa elaeidis, Allodiatrype arengae, A. elaeidicola, A. elaeidis and Diatrypella elaeidis are introduced. A checklist of Diatrypaceae occurring on palms (Arecaceae) and Thai diatrypaceous fungi is also provided.

Discovery of Cytospora species associated with canker disease of tree hosts from Mount Dongling of China

Article

Full-text available

Feb 2020

Members of Cytospora encompass important plant pathogens, saprobes and endophytes on a wide range of woody hosts with a worldwide distribution. In the current study, we obtained seven representative isolates from six tree hosts of Betulaceae, Juglandaceae, Rosaceae, Tiliaceae and Ulmaceae in Mount Dongling of China. Based on morphological comparison and phylogenetic analyses using partial ITS, LSU, act , rpb2 , tef1-α and tub2 gene sequences, we identified two known species ( Cytospora leucostoma and C. pruinopsis ) and two novel species ( C. coryli and C. spiraeicola ). These results represent the first study on Cytospora species associated with canker disease from Mount Dongling of China.

Morpho-molecular characterization of microfungi associated with marine based habitats

Article

Full-text available

Jan 2020

Our investigation of saprobic marine fungi in India, Thailand, Sweden and the UK yielded 57 species accommodated in 26 families. In the present study, we describe two new genera, 37 new species and 15 new host records. Novel genera, Halocryptosphaeria and Halotestudina are introduced within Diatrypaceae (Xylariales) and Testudinaceae (Xenoacremonium brunneosporum are introduced based on multigene analyses and morphological studies. This study also provides insights into the diversity of fungi from marine based habitats and confirm that they occupy diverse marine niches. We also demonstrate how marine based substrates, including sand dunes, are fascinating substrates for discovering novel taxa. All taxa described herein are based on morphological examination of fresh specimens supported by multigene phylogenies to better integrate taxa into higher taxonomic framework and infer their phylogenetic relationships as well as establish new species.

Fungal diversity notes 929–1035: taxonomic and phylogenetic contributions on genera and species of fungi

Article

Aug 2020
FUNGAL DIVERS

This article is the ninth in the series of Fungal Diversity Notes, where 107 taxa distributed in three phyla, nine classes, 31 orders and 57 families are described and illustrated. Taxa described in the present study include 12 new genera, 74 new species, three new combinations, two reference specimens, a re-circumscription of the epitype, and 15 records of sexualasexual morph connections, new hosts and new geographical distributions. Twelve new genera comprise Brunneofusispora, Brunneomurispora, Liua, Lonicericola, Neoeutypella, Paratrimmatostroma, Parazalerion, Proliferophorum, Pseudoastrosphaeriellopsis, Septomelanconiella, Velebitea and Vicosamyces. Seventy-four new species are Agaricus memnonius, A. langensis, Aleurodiscus patagonicus, Amanita flavoalba, A. subtropicana, Amphisphaeria mangrovei, Baorangia major, Bartalinia kunmingensis, Brunneofusispora sinensis, Brunneomurispora lonicerae, Capronia camelliaeyunnanensis, Clavulina thindii, Coniochaeta simbalensis, Conlarium thailandense, Coprinus trigonosporus, Liua muriformis, Cyphellophora filicis, Cytospora ulmicola, Dacrymyces invisibilis, Dictyocheirospora metroxylonis, Distoseptispora thysanolaenae, Emericellopsis koreana, Galiicola baoshanensis, Hygrocybe lucida, Hypoxylon teeravasati, Hyweljonesia indica, Keissleriella caraganae, Lactarius olivaceopallidus, Lactifluus midnapurensis, Lembosia brigadeirensis, Leptosphaeria urticae, Lonicericola hyaloseptispora, Lophiotrema mucilaginosis, Marasmiellus bicoloripes, Marasmius indojasminodorus, Micropeltis phetchaburiensis, Mucor orantomantidis, Murilentithecium lonicerae, Neobambusicola brunnea, Neoeutypella baoshanensis, Neoroussoella heveae, Neosetophoma lonicerae, Ophiobolus malleolus, Parabambusicola thysanolaenae, Paratrimmatostroma kunmingensis, Parazalerion indica, Penicillium dokdoense, Peroneutypa mangrovei, Phaeosphaeria cycadis, Phanerochaete australosanguinea, Plectosphaerella kunmingensis, Plenodomus artemisiae, P. lijiangensis, Proliferophorum thailandicum, Pseudoastrosphaeriellopsis kaveriana, Pseudohelicomyces menglunicus, Pseudoplagiostoma mangiferae, Robillarda mangiferae, Roussoella elaeicola, Russula choptae, R. uttarakhandia, Septomelanconiella thailandica, Spencermartinsia acericola, Sphaerellopsis isthmospora, Thozetella lithocarpi, Trechispora echinospora, Tremellochaete atlantica, Trichoderma koreanum, T. pinicola, T. rugulosum, Velebitea chrysotexta, Vicosamyces venturisporus, Wojnowiciella kunmingensis and Zopfiella indica. Three new combinations are Baorangia rufomaculata, Lanmaoa pallidorosea and Wojnowiciella rosicola. The reference specimens of Canalisporium kenyense and Tamsiniella labiosa are designated. The epitype of Sarcopeziza sicula is re-circumscribed based on cyto- and histochemical analyses. The sexual-asexual morph connection of Plenodomus sinensis is reported from ferns and Cirsium for the first time. In addition, the new host records and country records are Amanita altipes, A. melleialba, Amarenomyces dactylidis, Chaetosphaeria panamensis, Coniella vitis, Coprinopsis kubickae, Dothiorella sarmentorum, Leptobacillium leptobactrum var. calidus, Muyocopron lithocarpi, Neoroussoella solani, Periconia cortaderiae, Phragmocamarosporium hederae, Sphaerellopsis paraphysata and Sphaeropsis eucalypticola.

Some species of the genus Diatrype from the Czech Republic preserved in PRM, BRNM and KRAM.

Article

Aug 2005

Andrzej Chlebicki

Two new species of Eutypella and a new combination in the genus Peroneutypa (Diatrypaceae)

Article

Aug 2019
MYCOL PROG

We describe two new species of Eutypella, E. persica from dead branch of Alnus sp. inGuilan province, and E. quercina from dead branch of Quercus sp. in East Azerbaijanprovince of Iran, using morphological and molecular data. Eutypella persica has largestromata with 20–70 perithecia in a valsoid arrangement, sulcate to smooth ostioles,and produces an asexual structure on both natural substrate and culture media.Eutypella quercina, lacking the asexual structure, is characterized by mostly circularstromata with 10–50 perithecia in a valsoid arrangement and smooth ostioles.Phylogenetic relationships of Eutypella with other genera of the Diatrypaceae are hereinferred using maximum parsimony and neighbor-joining analyses of the ITS rDNA andpartial β-tubulin gene. The new combination, Peroneutypa iranica, is proposed basedon morphological features.