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Submitted 2 July 2019, Accepted 2 December 2019, Published 28 January 2020
Corresponding Author: Prapassorn D. Eungwanichayapant – e-mail – prapassorn@mfu.ac.th 239
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A new genus Allodiatrype, five new species and a new host record of
diatrypaceous fungi from palms (Arecaceae)
Konta S1,2,3, Maharachchikumbura SSN4, Senanayake IC5, McKenzie EHC6,
Stadler M7, Boonmee S1,2, Phookamsak R1,3,11,12, Jayawardena RS1,2, Senwanna
C1,8, Hyde KD1,3,12, Elgorban AM9,10 and Eungwanichayapant PD2*
1 Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
2 School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
3 Centre for Mountain Futures (CMF), Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201,
Yunnan, People’s Republic of China
4 School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu, 611731,
People’s Republic of China
5 Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Science and Oceanography, Shenzhen
University, 3688, Nanhai Avenue, Nanshan, Shenzhen 518055, People’s Republic of China
6 Manaaki Whenua Landcare Research, Private Bag 92170, Auckland, New Zealand
7 Department of Microbial Drugs, Helmholtz-Zentrum für Infektionsforschung GmbH, Inhoffenstrasse 7, 38124
Brunswick, Germany
8 Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
9 Center of Excellence in Biotechnology Research, King Saud University P.O. Box. 2455, Riyadh 11451, Saudi Arabia
10 Department of Botany and Microbiology, College of Sciences, King Saud University P.O. Box. 2455, Riyadh 11451,
Saudi Arabia, Riyadh, Saudi Arabia
11 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy
of Sciences, Kunming 650201, Yunnan, People’s Republic of China
12 East and Central Asia Regional Office, World Agroforestry Centre (ICRAF), Kunming 650201, Yunnan, People’s
Republic of China
Konta S, Maharachchikumbura SSN, Senanayake IC, McKenzie EHC, Stadler M, Boonmee S,
Phookamsak R, Jayawardena RS, Senwanna C, Hyde KD, Elgorban AM, Eungwanichayapant PD
2020 – A new genus Allodiatrype, five new species and a new host record of diatrypaceous fungi
from palms (Arecaceae). Mycosphere 11(1), 239–268, Doi 10.5943/mycosphere/11/1/4
Abstract
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.
Keywords – 6 novel taxa – Diatrypaceae – morphology – palm fungi – phylogeny – Thai fungi –
Xylariales
Introduction
The ascomycete family Diatrypaceae Nitschke was introduced and typified by Diatrype Fr.
(Nitschke 1869). Diatrypaceous taxa have a worldwide distribution in aquatic and terrestrial
habitats (Chlebicki 1986, Glawe & Jacobs 1987, Carmarán et al. 2006, de Almeida et al. 2016,
Mycosphere 11(1): 239–268 (2020) www.mycosphere.org ISSN 2077 7019
Article
Doi 10.5943/mycosphere/11/1/4
240
Dayarathne et al. 2016, Mayorquin et al. 2016, Senwanna et al. 2017, Shang et al. 2017, 2018,
Moyo et al. 2018a). Most genera in Diatrypaceae are wood-inhabiting saprobes (Trouillas et al.
2011, Grassi et al. 2014, Mehrabi et al. 2016, Hyde et al. 2019, Phookamsak et al. 2019); some are
plant pathogens causing cankers, dieback and grapevine trunk diseases (caused by Anthostoma
decipiens (DC.) Nitschke, Cryptovalsa ampelina (Nitschke) Fuckel, Eutypa lata (Pers.) Tul. & C.
Tul., Eutypella citricola Speg, E. microtheca Trouillas, W.M. Pitt & Gubler, E. parasitica R.W.
Davidson & R.C. Lorenz (Mostert et al. 2004, Jurc et al. 2006, Luque et al. 2006, 2012, Pitt et al.
2013, Rolshausen et al. 2014, Paolinelli-Alfonso et al. 2015, Mayorquin et al. 2016, Kowalski &
Bednarz 2017, Moyo et al. 2018b); and some are endophytes such as Diatrypella frostii Peck,
Libertella Desm. and Peroneutypa scoparia (Schwein.) Carmarán & A.I. Romero (de Errasti et al.
2010, Vieira et al. 2011, Grassi et al. 2014).
The number of genera accepted in Diatrypaceae has changed over the years, and four genera
have been introduced in the past ten years e.g. Allocryptovalsa Senwanna, Phookamsak & K.D.
Hyde, Diatrypasimilis JJ. Zhou & Kohlm., Halodiatrype Dayarathne & K.D. Hyde and
Neoeutypella M. Raza, Q.J. Shang, Phookamsak & L. Cai (Chalkley et al. 2010, Liu et al. 2015,
Dayarathne et al. 2016, Senwanna et al. 2017, Phookamsak et al. 2019). Wijayawardene et al.
(2017a) included 17 genera in the family; Phookamsak et al. (2019) introduced the new genus
Neoeutypella, with two new species, while Hyde et al. (2019) introduced the new species
Diatrypella delonicis R.H. Perera & K.D. Hyde. Diatrypaceae is, however, relatively poorly studied
with regard to potential biotechnological applications, such as the production of enzymes and
beneficial secondary metabolites (Ciavatta et al. 2008, Grassi et al. 2014). The only species that
have been studied thoroughly for bioactive compounds are the plant pathogenic Eutypa lata and
Peroneutypa (syn: Eutypella) scoparia (Helaly et al. 2018).
Divergent time estimates and the evolution of major lineages in the Sordariomycetes have
indicated that Diatrypaceae has an affinity to many families, such as Graphostromataceae M.E.
Barr, J.D. Rogers & Y.M. Ju, Hypoxylaceae DC., Lopadostomataceae Daranag. & K.D. Hyde,
Microdochiaceae Hern.-Restr., Crous & J.Z. Groenew., Requienellaceae Boise., and Xylariaceae
Tul. & C. Tul. in the order Xylariales at 66–252 Mya, which is the same as the common divergence
times of most of fungal families (Samarakoon et al. 2016, Hongsanan et al. 2017). Members of
Diatrypaceae mostly have immersed to erumpent or rarely superficial, black or dark brown,
eustromatic or pseudostromatic stromata, 8-spored or polysporous asci, hyaline to light brown,
allantoid ascospores and a libertella-like asexual morph (Senanayake et al. 2015, Wijayawardene et
al. 2017b). The concepts of segregating genera in Diatrypaceae are still rather confused as
mentioned in Maharachchikumbura et al. (2016) and Shang et al. (2017, 2018). The placement of
species in each genus of Diatrypaceae is very confused with many genera being polyphyletic
(Shang et al. 2017, 2018). Hence, it is necessary to use both molecular (mostly based on ITS and
TUB2 sequence data) and morphological data for the primary identification and classification of
diatrypaceouse taxa.
The number of new microfungi that can be potentially discovered in Thailand is large (Hyde et
al. 2018). In this study, we introduce a new genus, five new species, a new combination, new host
and new geographical record for Diatrypaceae occurring on palms with morphological and
phylogenetic evidence. Detailed descriptions, illustrations, and notes for each taxon are also provided.
Materials & Methods
Collection, isolation, and identification
Fresh materials were collected from Thailand (Chiang Rai, Krabi, and Phang-Nga Provinces)
during 2014–2015 on Arenga pinnata (Wurmb) Merr., Brahea armata S. Watson, Calamus L., and
Elaeis guineensis Jacq. The taxa were identified based on morphological characteristics and DNA
sequence data. Isolations and specimen examinations were conducted following the method
provided by Konta et al. (2016). The specimens were deposited in the herbarium of Mae Fah Luang
University (MFLU) and duplicated in the herbarium of Cryptogams, Kunming Institute of Botany
241
Academia Sinica (KUN-HKAS). Cultures were deposited in Mae Fah Luang Culture Collection
(MFLUCC) at Mae Fah Luang University, Chiang Rai, Thailand. Facesoffungi and Index
Fungorum numbers were registered as outlined in Jayasiri et al. (2015) and Index Fungorum
(2019).
DNA extraction and amplification (PCR)
Genomic DNA was extracted from fungal mycelium using the Biospin Fungus Genomic
DNA extraction Kit (BioFlux, P.R. China) following the manufacturer’s protocol. The partial
nucleotide genes were subjected to PCR amplification and sequencing of internal transcribed spacer
regions and intervening 5.8S rRNA gene (ITS) of the rDNA operon (White et al. 1990), 28S rRNA
gene (LSU) (Vilgalys & Hester 1990), 18S ribosomal RNA (SSU) (White et al. 1990), translation
elongation factor 1-alpha (tef1) (Rehner & Buckley 2005), RNA polymerase II second largest
subunit (RPB2) (Liu et al. 1999, Sung et al. 2007) and β-tubulin (TUB2) (Glass & Donaldson 1995,
O’Donnell & Cigelnik 1997). For primers and conditions see Table 1.
The total volume of PCR mixtures for amplification were 25 μl containing 8.5 μl ddH2O, 12.5
μl 2× Easy Taq PCR Super Mix (mixture of Easy Taq TM DNA Polymerase, dNTPs and optimized
buffer (Beijing Trans Gen Biotech Co., Beijing, P.R. China), 2 μl of DNA template, 1 μl of each
forward and reverse primers (10 pM). The quality of PCR products was checked on 1% agarose gel
electrophoresis stained with 4S green nucleic acid (Life Science Products & Services, Shanghai,
P.R. China). Purification and sequencing of PCR products were carried out by Sangon Biotech Co.,
Shanghai, P.R. China. The resulting fragments were sequenced in both directions with primers
above. The DNA sequences generated were analyzed and consensus sequences were computed using
SeqMan software.
Table 1 Details of genes/loci with PCR primers and PCR conditions.
Genes/loci
PCR primers
(forward/reverse)
PCR conditions
ITS, LSU, SSU, tef1
ITS5/ITS4, LR5/LR0R,
NS4/NS1, 983F/2218R
a; 95 °C: 30 s, 55 °C: 50 s, 72°C: 30 s (35 cycles); c
RPB2
fRPB2-5f/fRPB2–7cR
b; 95 °C: 1 min, 54 °C: 2 min, 72 °C: 1.5 min (35 cycles); c
TUB2
T1/ Bt2b
b; 94 °C: 1 min, 52 °C: 1 min, 72 °C: 1.5 min; c
a Initiation step of 95 °C: 3 min. b Initiation step of 95 °C: 5 min. c Final elongation step of 72 °C: 10 min and final hold
at 4 °C.
Phylogenetic analysis
The new sequences generated in this study were deposited in GenBank (Table 2) even if they
were not used in the phylogenetic tree. The sequences generated in this study were analysed with
additional sequences obtained from GenBank, based on BLAST searches and the literature (Hyde
et al. 2019, Phookamsak et al. 2019). Sequences of the ITS and TUB2 were analysed individually
and in combination. Only ITS and TUB2 sequence data were used in the analyses based on
previous literature and other gene sequences were deposited in GenBank for future studies.
Sequence alignments were carried out with MAFFT v.6.864b (Katoh & Standley 2013) and
alignments were manually improved where necessary. The single gene datasets were combined
using Mega7 (Kumar et al. 2016). Data were converted from fasta to nexus and PHYLIP format
with Alignment Transformation Environment online, https://sing.ei.uvigo.es/ALTER/ (Glez-Peña et
al. 2010).
The phylogenetic methods used in this study included maximum likelihood analysis (ML)
performed with RAxMl GUI v.1.0. (Stamatakis 2006, Silvestro & Michalak 2011) and Bayesian
posterior probabilities (BYPP). The latter method was performed at CIPRES using Bayesian
analysis on XSEDE (v.3.2.6) as part of the “MrBayes on XSEDE” tool (Huelsenbeck & Ronquist
2001, Miller et al. 2010). MrModelTest v. 2.2 was used to determine the best nucleotide substitution
242
model settings for the alignment for each data partition of the Bayesian analysis (Nylander 2004).
The model of evolution was performed using MrModelTest 2.2 (Nylander 2004) under the Akaike
information criterion (AIC). GTR+I+G model was selected as the best-fit models of the combined
dataset for maximum likelihood and Bayesian analysis (Nylander 2004). Bayesian posterior
probabilities (BYPP) were determined by Markov Chain Monte Carlo sampling (MCMC) in
MrBayes on XSEDE v.3.2.6. Six simultaneous Markov Chains were run for 3,000,000 generations
and trees were sampled every 1,000th generation. MCMC heated chain was set with a “temperature”
value of 0.20. All sampled topologies beneath the asymptote (25%) were discarded as part of a
burn-in procedure; the remaining trees (4,502) were used for calculating posterior probabilities in
the majority rule consensus tree. BYPP equal to/or greater than 0.90 is given near to each node
(Fig. 1). The phylogenetic trees were visualized in Fig Tree v1.4.0 (Rambaut 2006) and edited
using Microsoft Office PowerPoint 2010 and Adobe Illustrator CS6 (Adobe Systems, USA). The
alignments and respective phylogenetic trees were deposited in TreeBASE (submission ID: 25674).
Results
Phylogenetic analyses
Phylogenetic analyses of combined ITS and TUB2 sequence data based on ML and BYPP
analyses indicate that the two tree topologies are similar. The dataset consists of 117 taxa for
representative strains of species in Diatrypaceae. The total alignment length comprises 2,570
characters including gaps. The RAxML analysis resulted in a best scoring likelihood tree selected
with a final ML optimization likelihood value of -22955.866702 which is represented in Fig. 1. The
final likelihood tree was evaluated and optimized under GAMMA model parameters, with 1,547
distinct alignment patterns and 65.11% of undetermined characters or gaps. Bayesian posterior
probabilities from MCMC were evaluated with a final average standard deviation of the split
frequency of 0.011661.
The phylogram generated from the combined ITS and TUB2 sequence data supports
establishment of a new genus, five new species and six new host records of diatrypaceous fungi
within Diatrypaceae (Fig. 1). Allodiatrype species formed a basal clade to Neoeutypella, with
Diatrype enteroxantha as the sister clade. Of the five new species, Allocryptovalsa elaeidis
(MFLUCC 15-0707) formed a sister clade with A. polyspora (MFLUCC 17-0364, type species) and
A. rabenhorstii with high bootstrap support within the genus Allocryptovalsa (100% ML, 1.00
BYPP). Allodiatrype arengae (MFLUCC 15-0713) clustered with A. elaeidis with high bootstrap
support (100% ML, 1.00 BYPP). Allodiatrype elaeidis (MFLUCC 15-0708) clustered with A.
elaeidicola with 61% ML bootstrap support. Allodiatrype elaeidicola (MFLUCC 15-0737)
clustered with A. thailandica with low bootstrap support. Diatrypella elaeidis (MFLUCC 15-0279)
is sister to Diatrypella delonicis (MFLUCC 15-1014) with low bootstrap support (61% ML). Of the
new hosts and geographical records, Allodiatrype thailandica (MFLUCC 15-0711) appeared related
to the generic type of the genus (MFLUCC 14-1210). Our new isolate of Diatrypella (MFLUCC
17-0368) grouped with ex-type strain of D. heveae (MFLUCC 15-0274) with high bootstrap
support (100% ML, 1.00 BYPP).
Table 2 GenBank accession numbers of sequences used in phylogenetic analyses.
Species Strains
GenBank accession
numbers
References
ITS
TUB2
Allocryptovalsa cryptovalsoidea
HVFIG02
HQ692573
HQ692524
Trouillas et al. (2011)
Allocryptovalsa cryptovalsoidea
HVFIG05
HQ692574
HQ692525
Trouillas et al. (2011)
Allocryptovalsa elaeidis
MFLUCC 15-0707
MN308410
MN340296
This study
Allocryptovalsa polyspora T
MFLUCC 17-0364
MF959500
MG334556
Senwanna et al. (2017)
Allocryptovalsa rabenhorstii
WA07CO
HQ692620
HQ692522
Trouillas et al. (2011)
243
Table 2 Continued.
Species Strains
GenBank accession
numbers
References
ITS
TUB2
Allocryptovalsa rabenhorstii
WA08CB
HQ692619
HQ692523
Trouillas et al. (2011)
Allodiatrype arengae T
MFLUCC 15-0713
MN308411
MN340297
This study
Allodiatrype elaeidicola
MFLUCC 15-0737a
MN308415
MN340299
This study
Allodiatrype elaeidicola
MFLUCC 15-0737b
MN308416
-
This study
Allodiatrype elaeidis
MFLUCC 15-0708a
MN308412
MN340298
This study
Allodiatrype elaeidis
MFLUCC 15-0708b
MN308413
-
This study
Allodiatrype thailandica
‘Diatrype thailandica’
MFLUCC 14-1210
KU315392
-
Li et al. (2016)
Allodiatrype thailandica
MFLUCC 15-0711
MN308414
-
This study
Anthostoma decipiens T
IPV-FW349
AM399021
AM920693
Unpublished
Anthostoma decipiens T
JL567
JN975370
JN975407
Luque et al. (2012)
Cryptosphaeria eunomia T
C1C, CBS 216.87
AJ302417
-
Acero et al. (2004)
Cryptosphaeria eunomia T
C5C, CBS 223.87
AJ302421
-
Acero et al. (2004)
Cryptosphaeria ligniota
CBS 273.87
KT425233
KT425168
Acero et al. (2004)
Cryptosphaeria moravica
CBS 244.87
HM164735
HM164769
Trouillas & Gubler (2010)
Cryptosphaeria pullmanensis
ATCC 52655
KT425235
KT425170
Trouillas et al. (2015)
Cryptosphaeria pullmanensis
HBPF24
KT425202
GQ294014
Trouillas et al. (2010)
Cryptosphaeria subcutanea
CBS 240.87
KT425232
KT425167
Trouillas et al. (2015)
Cryptosphaeria subcutanea
DSUB100A
KT425189
KT425124
Trouillas et al. (2015)
Cryptovalsa ampelina
A001
GQ293901
GQ293972
Trouillas et al. (2010)
Cryptovalsa ampelina
DRO101
GQ293902
GQ293982
Trouillas et al. (2010)
Diatrype brunneospora
CNP01
HM581946
HQ692478
Trouillas et al. (2011)
Diatrype bullata
UCDDCh400
DQ006946
DQ007002
Rolshausen et al. (2006)
Diatrype decorticata
1056
KU320621
-
de Almeida et al. (2016)
Diatrype bullata
D6C, CBS 215.87
AJ302422
-
Acero et al. (2004)
Diatrype enteroxantha
HUEFS155114
KM396617
KT003700
de Almeida et al. (2016)
Diatrype enteroxantha
HUEFS155116
KM396618
KT022236
de Almeida et al. (2016)
Diatrype disciformis T
D21C, CBS 205.87
AJ302437
-
Acero et al. (2004)
Diatrype disciformis T
D7M, GB5815
AJ302423
-
Acero et al. (2004)
Diatrype macowaniana
D15C, CBS 214.87
AJ302431
-
Acero et al. (2004)
Diatrype oregonensis
DPL200
GQ293940
GQ293999
Trouillas et al. (2010)
Diatrype palmicola
MFLUCC 11-0018
KP744439
-
Liu et al. (2015)
Diatrype palmicola
MFLUCC 11-0020
KP744438
-
Liu et al. (2015)
Diatrype polycocca
D16C, CBS 213.87
AJ302432
-
Acero et al. (2004)
Diatrype spilomea
D17C
AJ302433
Acero et al. (2004)
Diatrype stigma
DCASH200
GQ293947
GQ294003
Diatrype stigma
UCD23-Oe
JX515704
JX515670
Úrbez-Torres et al. (2013)
Diatrype undulata
D20C, CBS 271.87
AJ302436
-
Acero et al. (2004)
Diatrype undulata
Olrim324
AY354239
-
Lygis et al. (2004)
Diatrype whitmanensis
CDB011
GQ293954
GQ294010
Trouillas et al. (2010)
Diatrype whitmanensis
DCHES100
GQ293951
GQ294008
Trouillas et al. (2010)
Diatrypella atlantica
HUEFS 136873
KM396614
KR259647
de Almeida et al. (2016)
Diatrypella atlantica
HUEFS 194228
KM396615
KR363998
de Almeida et al. (2016)
Diatrypella banksiae
CPC 29118
KY173402
-
Crous et al. (2013)
Diatrypella delonicis
MFLUCC 15-1014
MH812994
MH847790
Hyde et al. (2019)
Diatrypella delonicis
MFLU 16-1032
MH812995
MH847791
Hyde et al. (2019)
Diatrypella elaeidis
MFLUCC 15-0279
MN308417
MN340300
This study
Diatrypella favacea
Isolate 380
KU320616
-
de Almeida et al. (2016)
Diatrypella frostii
UFMGCB 1917
HQ377280
-
Vieira et al. (2011)
Diatrypella heveae
MFLUCC 17-0368
MF959501
MG334557
Senwanna et al. (2017)
Diatrypella heveae
MFLUCC 15-0274
MN308418
MN340301
This study
Diatrypella iranensis
KDQ18
KM245033
-
Mehrabi et al. (2015)
Diatrypella major
Isolate 1058
KU320613
-
de Almeida et al. (2016)
Diatrype oregonensis
CA117
GQ293934
GQ293996
Trouillas et al. (2010)
Diatrypella prominens
DL28A, ATCC 64182
AJ302442
-
Acero et al. (2004)
Diatrype oregonensis
DPL200
GQ293940
GQ293999
Trouillas et al. (2010)
Diatrype prominens
FJ430594
-
Unpublished
244
Table 2 Continued.
Species Strains
GenBank accession
numbers
References
ITS
TUB2
Diatrype prominens
SBen212
KU721868
-
Lawrence et al. (2017)
Diatrypella pulvinata
H048
FR715523
FR715495
de Almeida et al. (2016)
Diatrypella tectonae
MFLUCC 12-0172a
KY283084
-
Shang et al. (2017)
Diatrypella tectonae
MFLUCC 12-0172b
KY283085
KY421043
Shang et al. (2017)
Diatrypella verruciformis T
UCROK1467
JX144793
JX174093
Lynch et al. (2013)
Diatrypella verruciformis T
UCROK754
JX144783
JX174083
Lynch et al. (2013)
Diatrypella vulgaris
HVFRA02
HQ692591
HQ692503
Trouillas et al. (2011)
Diatrypella vulgaris
HVGRF03
HQ692590
HQ692502
Trouillas et al. (2011)
Eutypa armeniacae
ATCC 28120
DQ006948
DQ006975
Rolshausen et al. (2006)
Eutypa astroidea
E49C, CBS 292.87
AJ302458
DQ006966
Rolshausen et al. (2006)
Eutypa flavovirens
E48C, CBS 272.87
AJ302457
DQ006959
Rolshausen et al. (2006)
Eutypa laevata
E40C CBS 291.87
AJ302449
-
Acero et al. (2004)
Eutypa lata T
CBS 290.87
HM164736
HM164770
Trouillas & Gubler (2010)
Eutypa lata T
EP18
HQ692611
HQ692501
Trouillas et al. (2011)
Eutypa lata T
RGA01
HQ692614
HQ692497
Trouillas et al. (2011)
Eutypa lejoplaca
CBS 248.87
DQ006922
DQ226974
Rolshausen et al. (2006)
Eutypa leptoplaca
CBS 287.87
DQ006924
DQ006961
Rolshausen et al. (2006)
Eutypa maura
CBS 219.87
DQ006926
DQ006967
Rolshausen et al. (2006)
Eutypa microasca
BAFC 51550
KF964566
KF964572
Grassi et al. (2014)
Eutypa sparsa
3802 3b
AY684220
AY684201
Trouillas & Gubler (2004)
Eutypella cerviculata T
EL59C
AJ302468
-
Acero et al. (2004)
Eutypella cerviculata T
M68
JF340269
-
Arhipova et al. (2012)
Eutypella citricola
HVGRF01
HQ692579
HQ692512
Trouillas et al. (2011)
Eutypella citricola
HVVIT07
HQ692589
HQ692521
Trouillas et al. (2011)
Eutypella leprosa
EL54C, CBS 276.87
AJ302463
-
Acero et al. (2004)
Eutypella leprosa
Isolate 60
KU320622
-
de Almeida et al. (2016)
Eutypella microtheca
ADEL200
HQ692559
HQ692527
Trouillas et al. (2011)
Eutypella microtheca
BCMX01
KC405563
KC405560
Paolinelli-Alfonso et al. (2015)
Eutypella parasitica
CBS 210.39
DQ118966
-
Jurc et al. (2006)
Eutypella semicircularis
MP4669
JQ517314
-
Mehrabi et al. (2016)
Eutypella vitis
UCD2291AR
HQ288224
HQ288303
Úrbez-Torres et al. (2012)
Eutypella vitis
UCD2428TX
FJ790851
GU294726
Úrbez-Torres et al. (2009)
Halodiatrype avicenniae
MFLUCC 15-0953
KX573916
KX573931
Dayarathne et al. (2016)
Halodiatrype salinicola T
MFLUCC 15-1277
KX573915
KX573932
Dayarathne et al. (2016)
Kretzschmaria deusta
CBS 826.72
KU683767
KU684190
U’Ren et al. (2016)
Monosporascus cannonballus T
CMM3646
JX971617
-
Unpublished
Monosporascus cannonballus T
ATCC 26931
FJ430598
-
Unpublished
Neoeutypella baoshanensis T
EL51C, CBS 274.87
AJ302460
-
Acero et al. (2004)
Neoeutypella baoshanensis T
LC 12111
MH822887
MH822888
Hyde et al. (2019)
Pedumispora rhizophorae T
BCC44877
KJ888853
-
Klaysuban et al. (2014)
Pedumispora rhizophorae T
BCC44878
KJ888854
-
Klaysuban et al. (2014)
Peroneutypa alsophila
EL58C, CBS 250.87
AJ302467
-
Acero et al. (2004)
Peroneutypa comosa
BAFC 393
KF964568
-
Grassi et al. (2014)
Peroneutypa curvispora
HUEFS 136877
KM396641
-
de Almeida et al. (2016)
Peroneutypa diminutiasca
MFLUCC 17-2144
MG873479
-
Shang et al. (2018)
Peroneutypa diminutispora
HUEFS 192196
KM396647
-
de Almeida et al. (2016)
Peroneutypa kochiana
EL53M
AJ302462
-
Acero et al. (2004)
Peroneutypa longiasca
MFLUCC 17-0371
MF959502
MG334558
Senwanna et al. (2017)
Peroneutypa mackenziei
MFLUCC 16-0072
KY283083
KY706363
Shang et al. (2017)
Peroneutypa mangrovei
NFCCI-4246
MG844286
MH094409
Phookamsak et al. (2019)
Peroneutypa rubiformis
MFLUCC 17-2142
MG873477
-
Shang et al. (2018)
Peroneutypa scoparia
MFLUCC 11-0478
KU940151
-
Dai et al. (2016)
Peroneutypa scoparia
MFLUCC 18-1111
MK603519
MK101307
Hyde et al. (2019)
Quaternaria quaternata
EL60C, CBS 278.87
AJ302469
-
Acero et al. (2004)
Quaternaria quaternata
GNF13
KR605645
-
Mehrabi et al. (2016)
Xylaria hypoxylon
CBS 122620
AM993141
KX271279
Peršoh et al. (2009)
Note: Newly generated sequences are in bold; T denotes the type species of the genus.
245
Figure 1 – Bayesian analyses the majority rule consensus tree of selected species in Diatrypaceae
generated from combined ITS and TUB2 sequence data. Bootstrap support values for maximum
246
likelihood (ML) greater than 50%, and Bayesian posterior probabilities (BYPP) greater than 0.90
are given at the nodes. Branches with 100% ML and 1.00 BYPP are shown with a blue dot. Ex-type
strains are in bold. Newly generated sequences are in red. Novel taxa are in red bold. The asterisks
represent unstable species.
Figure 1 – Continued.
Taxonomy
Allocryptovalsa Senwanna, Phookamsak & K.D. Hyde, in Senwanna, Phookamsak, Doilom, Hyde
& Cheewangkoon, Mycosphere 8(10): 1839 (2017)
Type species – Allocryptovalsa polyspora C. Senwanna, Phookamsak & K.D. Hyde
Notes – Allocryptovalsa was introduced and is typified by A. polyspora Senwanna,
Phookamsak & K.D. Hyde, which was collected from Hevea brasiliensis (Senwanna et al. 2017).
This genus comprises three species, A. cryptovalsoidea Trouillas, W.M. Pitt & Gubler ex
Senwanna, Phookamsak & K.D. Hyde, A. polyspora and A. rabenhorstii (Nitschke) C. Senwanna,
Phookamsak & K.D. Hyde (Index Fungorum 2019).
Allocryptovalsa elaeidis Konta & K.D. Hyde, sp. nov. Fig. 2
Index Fungorum number: IF556570, Facesoffungi number: FoF05116
Etymology – Epithet refers to host genus, Elaeis
Holotype – MFLU 15-1438
247
Saprobic on dead petiole of Elaeis guineensis (Arecaceae). Sexual morph – Stromata (285–
)330–730(–950) μm long, (250–)350–890(–1030) μm wide (x
̄ = 568 × 600 μm, n = 20), mostly
solitary, sometimes gregarious, surrounded by black circle on host surface, immersed to erumpent
in the bark, black, raised, pustulate, dome-shaped, 1–2-ascomata, with umbilicate ostiole appearing
on the surface of stroma. Ascomata (including neck) 325–460 μm high, 315–515 μm diam. (x
̄ = 365
× 400 μm, n = 20), perithecial, immersed in the stroma, covered with the epidermis of plant tissue,
delimited by a black zone in host tissues, globose to subglobose, glabrous, ostiole individual, with a
short neck. Ostiolar canal 180–250 μm high, 160–230 μm diam. (x
̄ = 250 × 230 μm, n = 5), sulcate,
with periphyses. Peridium 25–86 μm wide (x
̄ = 46 μm, n = 30), composed of two sections, outer
layer dark brown, thick-walled cells, arranged in textura angularis, inner layer hyaline, thin-walled
cells of textura angularis. Hamathecium composed of 3–13 μm wide (x
̄ = 6 μm, n = 40), filiform,
septate, hyaline, unbranched, paraphyses. Asci (including stalks) (55–)68–147(−157) × 14–26 μm
(x
̄ = 97 × 18 μm, n = 30), apex-bearing part (1.7–)2–4(−5.5) μm long (x
̄ = 3 μm, n = 30),
polysporous, unitunicate, clavate, with moderately short stalks, stalk-bearing part 35–56 μm long (x
̄
= 50 μm, n = 10). Ascospores (6–)7.5–9(−10.5) × 2–4 μm (x
̄ = 9 × 3 μm, n = 100), overlapping,
yellowish to brown, ellipsoidal to cylindrical or elongate-allantoid, aseptate, smooth-walled.
Asexual morph – Undetermined.
Geographical distribution – Thailand.
Culture characters – Ascospores germinated on Malt Extract Agar (MEA) within 24 hours.
Colonies on MEA, dense but thinner towards the edge, margin diffuse, white of upper surface (Fig.
2q). Additional sequence data – LSU (MN308401), SSU (MN308419) (MFLUCC 15-0707).
Material examined – THAILAND, Krabi Province, on dead petiole of Elaeis guineensis Jacq.
(Arecaceae), 3 December 2014, S. Konta, KBM01f (MFLU 15-1438, holotype); ex-type living
culture = MFLUCC 15-0707.
Notes – Allocryptovalsa elaeidis is morphologically most similar to A. polyspora,
overlapping in the number of ascomata per stroma, size of ascomata and asci (Senwanna et al.
2017). Although A. elaeidis and A. polyspora are morphologically similar, the phylogenetic
analyses strongly support these collections as two distinct species. Allocryptovalsa elaeidis differs
from A. cryptovalsoidea, A. polyspora, and A. rabenhorstii in host association, that is Elaeis
guineensis (A. elaeidis), Ficus carica (A. cryptovalsoidea), Hevea brasiliensis (A. polyspora),
Robinia L. and Vitis vinifera (A. rabenhorstii), respectively (Saccardo 1882, Trouillas et al. 2011,
Senwanna et al. 2017). The species of this genus have been reported from Australia, Germany,
Thailand, and USA (Saccardo 1882, Trouillas et al. 2011, Senwanna et al. 2017). Comparisons of
the nucleotide between Allocryptovalsa species are shown in Table 5. Thus, A. elaeidis is
introduced as the fourth species in Allocryptovalsa based on its different morphology coupled with
high support values from the phylogenetic analysis (100% ML, 1.00 BYPP, Fig. 1).
Allodiatrype Konta & K.D. Hyde, gen. nov.
Index Fungorum number: IF556641; Facesoffungi number: FoF06299
Etymology – In reference to the morphological resemblance to Diatrype
Saprobic on dead petiole of palm (Arecaceae), and on a dead stem of unidentified plants.
Sexual morph: Stromata scattered or aggregated on the host, erumpent, arising through cracks in
the bark, irregularly shaped or circular, orbicular, convex surface, 1–10-ascomata immersed in one
stroma, with or without a black stromatic zone. Ostiole opening through host bark and appearing as
black spots, surrounded with a ring-like, ostiolar opening, composed of an outer layer of dark
brown, small, tightly packed, thin parenchymatous cells and an inner layer of yellowish, large,
loosely packed, parenchymatous cells. Ascomata perithecial, immersed in stromatic tissue,
aggregated, brown, globose to sub-globose, narrowing towards the apex and very narrow at the
base of ostiolar canal, thin-walled, ostiolate; ostiolar canal, with periphyses, ostiolar opening
covered with carbonaceous, black cells; periphyses hyaline, filamentous. Peridium comprising an
outer layer of yellow-brown, thick-walled cells of textura angularis and a thin, inner stratum of
248
yellow, thick-walled cells of textura angularis. Hamathecium composed of septate, hyaline
paraphyses. Asci unitunicate, 8-spored, with long, narrow, thin-walled stalk, with cylindrical, thick-
walled, swollen upper portion, apex flat, with J-, cylindrical, conspicuous apical ring. Ascospores
seriate, hyaline becoming yellowish at maturity, allantoid, unicellular, thin-walled, with small fat
globules at each end, smooth-walled. Asexual morph: Undetermined.
Figure 2 – Allocryptovalsa elaeidis (MFLU 15-1438, holotype) a Stromata on host substrate. b
Close up of stroma. c Section of ascoma. d Peridium. e Ostiolar canal. f Paraphyses. g–j Asci. k–p
Ascospores. q Colony on MEA. Bars: a = 500 μm, b, c = 200 μm, d, e = 20 μm, f = 10 μm, g–j = 50
μm, k–p = 5 μm.
249
Geographical distribution – Thailand.
Type species – Allodiatrype arengae Konta & K.D. Hyde
Notes – Allodiatrype is introduced to accommodate Allodiatrype arengae, A. elaeidicola, A.
elaeidis, and A. (syn. Diatrype) thailandica. Allodiatrype is typified by A. arengae, which was
collected from Arenga pinnata (Arecaceae). The morphology of Allodiatrype species is closely
similar to that of Diatrype species. However, Allodiatrype differs in having 1–10-ascomata
immersed in one stroma, and with or lacking a black stromatic zone, while stromata of Diatrype
mostly spread over a large area, sometimes covering the host surface. As becomes evident from
Fig. 1, strains of both genera appear in distinct clades in a phylogeny based on multiple strains of
both genera, thereby justifying the erection of the new genus Allodiatrype.
Allodiatrype arengae Konta & K.D. Hyde, sp. nov. Fig. 3
Index Fungorum number: IF556929, Facesoffungi number: FoF05117
Etymology – Epithet refers to host genus, Arenga
Holotype – MFLU 15-1444
Saprobic on petiole of Arenga pinnata (Arecaceae). Sexual morph: Stromata 690–940 μm
long, 370–935 μm wide (x
̄ = 830 × 700 μm, n = 10), with well-developed interior, solitary,
superficial, black, without black stromatic, glabrous, subglobose to irregular, pustulate, 1–5-
ascomata, with umbilicate ostioles appearing on the surface of the stroma. Ostiole opening through
host bark and appearing as black spots, surrounded with a ring-like structure, composed of an outer
layer of dark brown, small, tightly packed, thin parenchymatous cells and an inner layer of yellow,
large, loosely packed, parenchymatous cells. Ascomata (excluding necks) 250–400 μm high, 240–
400 μm diam. (x
̄ = 340 × 300 μm, n = 25), perithecial, immersed in the stroma, globose to
subglobose, glabrous, ostioles individual, with a short neck. Ostiolar canal 100–170 μm high, 70–
130 μm diam. (x
̄ = 130 × 100 μm, n = 20), cylindrical, sulcate, with periphyses. Peridium 12–25
μm wide, (x
̄ = 20 μm, n = 40), composed of two sections, outer layer of brown to dark brown, thin-
walled cells, arranged in textura angularis, inner layer of hyaline thin-walled cells of textura
angularis. Hamathecium composed of 3–7 μm wide (x
̄ = 5 μm, n = 40), septate, hyaline
paraphyses. Asci (excluding stalks), spore-bearing part (14–)20(−45) × (4–)6–10(−12) μm (x
̄ = 30 ×
8 μm, n = 80), apically rounded, with J-apical ring, apex-bearing part (1.5–)3–5(−7.5) μm long (x
̄ =
4 μm, n = 40), 8-spored, unitunicate, clavate, with long stalks, (28–)34–89(–103) μm long (x
̄ = 64
μm, n = 60). Ascospores (6–)7–10(−12) × 2–3 μm (x
̄ = 10 × 2 μm, n = 120), overlapping, yellowish
to light-brown, ellipsoidal to cylindrical or elongate-allantoid, aseptate, smooth-walled. Asexual
morph: Undetermined.
Geographical distribution – Thailand.
Culture characters – Ascospores germinated on MEA within 24 hours. Colonies on MEA,
white in beginning, dense but thinning towards the edge, margin diffuse, reverse pale yellow in the
middle (Fig. 3q).
Additional sequence data – LSU (MN308402), SSU (MN308420), tef1 (MN525596), RPB2
(MN542886) (MFLUCC 15-0713).
Material examined – THAILAND, Phang-Nga Province, on dead petiole of Arenga pinnata
(Wurmb) Merr. (Arecaceae), 4 December 2014, S. Konta, PHR01b (MFLU 15-1444, holotype); ex-
type living culture = MFLUCC 15-0713.
Notes – Allodiatrype arengae was collected from a dead petiole of Arenga pinnata from
Phang-Nga Province, Thailand. Allodiatrype arengae is phylogenetically distinct from its sister
species A. elaeidicola, A. elaeidis, and A. thailandica with high statistical support (100% ML, 1.00
BYPP) (Fig. 1). A comparison of the nucleotide between Allodiatrype species and Diatrype
disciformis is given in Table 6. Morphologically, A. arengae has superficial stromata lacking a
black stromatic zone (Fig. 3a), while A. elaeidicola and A. elaeidis formed erumpent stromata,
arising through the cracks in bark or epidermis with a black stromatic zone (Figs 4a, b, 5a),
sometimes it covers the host surface (Fig. 4a, b). Other characters such as ascomata, asci and
250
ascospores are mostly similar to A. thailandica, and their sizes also overlap. Our new strain is also
recorded on a different host substrate from other species.
Figure 3 – Allodiatrype arengae (MFLU 15-1444, holotype) a Stromata on host substrate. b Close
up of stroma (ostiole opening surrounded with a ring-like structure). c, d Section of stroma. e
Ostiolar canal. f Peridium. g–j Asci. k–p Ascospores. q Colony on MEA. Bars: a = 500 μm, b, c =
200 μm, d, g–j = 50 μm, e, f = 20 μm, k–p = 5 μm.
251
Allodiatrype elaeidicola Konta & K.D. Hyde, sp. nov. Fig. 4
Index Fungorum number: IF556930, Facesoffungi number: FoF05118
Etymology – Epithet refers to host genus, Elaeis
Holotype – MFLU 15-1468
Saprobic on petiole of Elaeis guineensis (Arecaceae). Sexual morph: Stromata 1.2–2.8 mm
long, 0.96–1.66 mm diam. (x
̄ = 1.86 × 1.19 mm, n = 15), with well-developed interior, solitary to
gregarious, erumpent, black, with black stromatic zone extending down to the host surface,
glabrous, irregular in shape, pustulate, multi-ascomata, with umbilicate ostioles appearing on the
surface of the stroma. Ostiole opening through host bark and appearing as black spots, surrounded
with a ring-like structure, composed of an outer layer of dark brown, small, tightly packed, thin
parenchymatous cells and an inner layer of yellowish to orange, large, loosely packed,
parenchymatous cells. Ascomata (excluding necks) 280–430 μm high, 180–435 μm diam. (x
̄ = 370
× 270 μm, n = 30), perithecial, immersed in the stroma, globose to subglobose, glabrous, ostioles
individual, with a short neck. Ostiolar canal 120–185 μm high, 60–120 μm diam. (x
̄ = 140 × 95
μm, n = 30), cylindrical, sulcate, with periphyses. Peridium 14–40 μm wide (x
̄ = 29 μm, n = 30),
composed of two section layers, outer part; brown to dark brown, thick-walled cells, arranged in
textura angularis, inner layer; hyaline, thick-walled cells of textura angularis. Hamathecium
undermined. Asci (excluding stalks), spore-bearing part (17–)20–31(−43) × 4–7 μm (x
̄ = 26 × 6 μm,
n = 50), apically rounded, with J-apical ring, apex-bearing part (1.6–)2–3(−5) μm long (x
̄ = 3, n =
20), 8-spored, unitunicate, clavate, with long stalks, stalk-bearing part (32–)40–60(–76) μm long (x
̄
= 50 μm, n = 20). Ascospores (6.5–)8–10(−11) × 1.5–3 μm (x
̄ = 9 × 2 μm, n = 100), overlapping,
yellowish to brown, ellipsoidal to cylindrical or elongate-allantoid, aseptate, smooth-walled.
Asexual morph: Undetermined.
Geographical distribution – Thailand.
Culture characters – Ascospores germinated on MEA within 24 hours and germ tube was
produced from end cell. Colonies on MEA, white when beginning, dense but thinner towards edge,
margin diffuse, reverse coloration yellow (Fig. 4r).
Additional sequence data – LSU (MN308406), SSU (MN308424), tef1 (MN525598), RPB2
(MN542889) (MFLUCC 15-0737a); LSU (MN308407), SSU (MN308425), RPB2 (MN542890)
(MFLUCC 15-0737b).
Material examined – THAILAND, Phang-Nga Province, on dead petiole of Elaeis guineensis
Jacq. (Arecaceae), 5 December 2014, S. Konta, PHR10f (MFLU 15-1468, holotype, HKAS95035,
Fig. 4); ex-type living culture = MFLUCC 15-0737.
Notes – In the phylogenetic analyses, Allodiatrype elaeidicola is related to A. elaeidis and A.
thailandica with low bootstrap support (Fig. 1). However, the taxon is different in having yellow to
dark orange inner cells in the stromata (Fig. 4c, d) and thinner walled asci (Fig. 4i), while other
species have white to yellow inner cells in the stromata. A comparison of SSU, ITS, TUB2, RBP2
nucleotides to the type species, A. arengae shows that A. elaeidicola is significantly different from
A. arengae (SSU, 1/1030 bp (0.09%); ITS, 13/605 bp (2.14%); TUB2, 28/1586 bp (1.76%); RBP2,
3/1139 bp (0.26%) (Table 6).
Allodiatrype elaeidis Konta & K.D. Hyde, sp. nov. Fig. 5
Index Fungorum number: IF556931, Facesoffungi number: FoF05119
Etymology – Epithet refers to host genus, Elaeis
Holotype – MFLU 15-1439
Saprobic on petiole of Elaeis guineensis (Arecaceae). Sexual morph – Stromata 470–860 μm
long, 440–710 μm diam. (x
̄ = 630 × 550 μm, n = 10), with well-developed interior, solitary to
gregarious, erumpent, black, with black stromatic zone on host surface, glabrous, irregular in shape,
pustulate, bi- to multi-ascomata, with umbilicate ostioles appearing on the surface of the stroma.
Ostiole opening through host bark and appearing as black spots, surrounded with a ring-like
structure, composed of an outer layer of dark brown, small, tightly packed, thin parenchymatous
cells and an inner layer of white to light-yellow, large, loosely packed, parenchymatous cells.
252
Figure 4 – Allodiatrype elaeidicola (MFLU 15-1468, holotype) a Stromata on host substrate. b
Close up of stromata (ostiole opening surrounded with a ring-like structure). c, d Section of stroma.
e, i Mature asci. f–h Immature asci stained in cotton blue. j Ostiolar canal. k Peridium. l–q
Ascospores. r Colony on MEA. Bars: a, b = 1000 μm, c = 500 μm, d = 200 μm, e–k = 50 μm, l–q =
5 μm.
253
Ascomata (excluding necks) 250–350 μm high, 230–300 μm diam. (x
̄ = 330 × 280 μm, n = 10),
perithecial, immersed in the stroma, globose to subglobose, glabrous, ostioles individual, with a
short neck. Ostiolar canal 100–130 μm high, 95–115 μm diam. (x
̄ = 120 × 110 μm, n = 10),
cylindrical, sulcate, with periphyses. Peridium 20–40 μm wide (x
̄ = 30 μm, n = 50), composed of
two section layers, outer part; brown to dark brown, thick-walled cells, arranged in textura
angularis, inner layer; hyaline, thick-walled cells of textura angularis. Hamathecium composed of
2–7 μm wide (x
̄ = 4 μm, n = 60), filiform, longer than asci, septate, branch, hyaline, paraphyses.
Asci (excluding stalks), spore-bearing part (17–)20–30(−39) × 9–11(−14) μm (x
̄ = 25 × 11 μm, n =
60), apically rounded, with J- apical ring, apex-bearing part (1.5–)2–3(−5) μm long (x
̄ = 3 μm, n =
60), 8-spores, unitunicate, clavate, with moderately long stalks, stalk-bearing part (28–)36–65(–94)
μm long (x
̄ = 50 μm, n = 60). Ascospores (6–)8–10(−11) × 1.5–3) μm (x
̄ = 9 × 2 μm, n = 120),
overlapping, yellowish to pale-brown, ellipsoidal to cylindrical or elongate-allantoid, aseptate,
smooth-walled. Asexual morph – Undetermined.
Geographical distribution – Thailand.
Culture characters – Ascospores germinated on MEA within 24 hours and germ tube was
produced from end cell. Colonies on MEA, white at beginning, thinner towards edge, margin
diffuse, reverse coloration pale yellow (Fig. 5n).
Additional sequence data – LSU (MN308403), SSU (MN308421), tef1 (MN525597), RPB2
(MN542887) (MFLUCC 15-0708a); LSU (MN308404), SSU (MN308422), RPB2 (MN542888)
(MFLUCC 15-0708b).
Material examined – THAILAND, Krabi Province, on dead petiole of Elaeis guineensis Jacq.
(Arecaceae), 3 December 2014, S. Konta, KBM01g (MFLU 15-1439, holotype, Fig. 5); ex-type
living culture = MFLUCC 15-0708a.
Notes – Multigene phylogenetic analyses (Fig. 1( show that A. elaeidis (MFLUCC 15-0708(
forms a distinct lineage within the Allodiatype clade and is related to A. elaeidicola with 61% ML
bootstrap support. Allodiatrype elaeidis is morphologically similar to A. arengae, A. elaeidicola
and A. thailandica in ascospore size. However, the species differs in its wider asci (9–14 μm). A
comparison of ITS, TUB2, RBP2 nucleotides with A. arengae shows that A. elaeidis is significantly
different from A. arengae (ITS, 13/621 bp (2.09%(; TUB2, 16/1579 bp (1.01%); RBP2, 2/1138 bp
(0.17%) (Table 6).
Allodiatrype thailandica (R.H. Perera, Jian K. Liu & K.D. Hyde) Konta & K.D. Hyde, comb. nov.
Index Fungorum number: IF556932, Facesoffungi number: FoF01797
≡ Diatrype thailandica R.H. Perera, Jian K. Liu & K.D. Hyde, Fungal Diversity 78: 1–237.
10.1007/s13225-016-0366-9, [105] (2016)
Description: For original description see Li et al. )2016(.
Additional sequence data – LSU (MN308405), SSU (MN308423) (MFLUCC 15-0711).
Material examined – THAILAND, Phang-Nga Province, on dead petiole of Calamus sp.
(Arecaceae), 6 December 2014, S. Konta, DNH05f (MFLU 15-1442, Fig. 6); living culture =
MFLUCC 15-0711.
Notes – According to our analysis in Fig. 1, Diatrype thailandica grouped together with
Allodiatrype species without bootstrap support. This species only has LSU, ITS and SSU sequence
data and multigene analysis of ITS and TUB2 sequence data could not be resolved it from the ex-
type of A. elaeidicola. ITS is unlikely to provide good resolution in Diatrypaceae (Hongsanan et al.
2018). Thus, Diatrype thailandica is synonymized under Allodiatrype. Diatrype thailandica was
collected on stems of an unidentified plant from Chiang Rai, Thailand (Li et al. 2016) and in this
study, we collected it from Calamus (Arecaceae) in Phang-Nga Province. Morphological characters
obtained from the fresh specimen is similar to the description provided by Li et al. (2016).
254
Figure 5 – Allodiatrype elaeidis (MFLU 15-1439, holotype) a Stromata on host substrate. b Close
up of stromata (ostiole opening surrounded with a ring-like structure). c, d Section of stromata. e
Ostiolar canal. f Peridium. g, h Asci. i–m Ascospores. n Colony on MEA. Bars: a, b = 500 μm, c =
200 μm, d, g, h = 50 μm, e, f = 20 μm, i–m = 5 μm.
255
Figure 6 – Allodiatrype thailandica (MFLU 15-1442) a Stromata on host substrate. b Close up of
stromata. c, d Section of stroma. e Ostiolar canal. f Peridium. g–k Asci. l–o Ascospores. q Colony
on MEA. Bars: a = 1000 μm, b = 500 μm, c= 200 μm, e = 50 μm, f, h–k = 20 μm, l–o = 5 μm.
256
Diatrypella (Ces. & De Not.) De Not
Type species – Diatrypella verruciformis (Ehrh.) Nitschke
= Diatrypella favacea (Fr.) Ces. & De Not., Sfer. Ital.: 29 (1863))
Notes – The polyphyletic nature of Diatrypella has been reported based on the phylogeny of
ITS and TUB2 genes (de Almeida et al. 2016, Mehrabi et al. 2016, Senwanna et al. 2017, Shang et
al. 2017, Hyde et al. 2019). Currently, 114 epithets are listed under Diatrypella (Index Fungorum
2019). Recently an additional species, Diatrypella delonicis R.H. Perera & K.D. Hyde. was
introduced by Hyde et al. (2019).
Diatrypella heveae Senwanna, Phookamsak & K.D. Hyde, in Senwanna, Phookamsak, Doilom,
Hyde & Cheewangkoon, Mycosphere 8(10): 1846 (2017) Fig. 7
Index Fungorum number: IF553859, Facesoffungi number: FoF05121
Saprobic on petiole of Brahea armata (Arecaceae). Sexual morph: Stromata 660–2155 μm
long, 285–860 μm wide (x
̄ = 1430 × 500 μm, n = 11), with well-developed interior, solitary to
gregarious, erumpent in the bark, black, fusiform in shape, pustulate, 1–2-ascomata. Ascomata
(excluding necks) 70–300 μm high, 90–260 μm diam. (x
̄ = 181 × 210 μm, n = 20), immersed in the
stroma, sub globose to irregular, ostioles individual, with a short neck. Ostiolar canal 110–180 μm
high, 80–140 μm diam. (x
̄ = 140 × 110 μm, n = 20), cylindrical, sulcate, with periphyses. Peridium
10–35 μm wide (x
̄ = 22 μm, n = 30), composed of two layers, outer layer of brown, thin-walled
cells, arranged in textura angularis, inner layer of hyaline, thick-walled cells of textura angularis.
Hamathecium composed of 2–5 μm wide (x
̄ = 3 μm, n = 30), filiform, longer than asci, septate,
hyaline paraphyses. Asci (including stalks), spore-bearing part (55–)64–90(−105) ×(12–)13–
16(−18) μm (x
̄ = 80 × 15 μm, n = 50), apically rounded, with J- apical ring, apex-bearing part 2–
4(−6) μm long (x
̄ = 3, n = 35), polysporous, unitunicate, clavate, with moderately short stalks, stalk-
bearing part 24–36 μm long (x
̄ = 35 μm, n = 20). Ascospores (5–)6–10(−13) × 1−2.5 μm (x
̄ = 7 × 2
μm, n = 130), overlapping, yellowish to brown, ellipsoidal to cylindrical or elongate-allantoid,
aseptate, smooth-walled. Asexual morph: Undetermined.
Geographical distribution – Thailand, Chiang Rai, Wiang Chiang Rung District (Senwanna et
al. 2017).
Culture characters – Ascospores germinated on MEA within 24 hours, germ tube produced
from end cell. Colonies on MEA smooth, white, dense towards the edge, margin diffuse (Fig. 7v).
Additional sequence data – LSU (MN308409), SSU (MN308427), tef1 (MN525600), RPB2
(MN542892) (MFLUCC 15-0274).
Material examined – THAILAND, Chiang Rai Province, Mae Chan District, on dead petiole
of Brahea armata S. Watson. (Arecaceae), 25 November 2014, S. Konta, HR01a (MFLU 15-0020);
living culture = MFLUCC 15-0274.
Notes – We collected and illustrated Diatrypella heveae (MFLU 15-0020) from Chiang Rai
Province, Thailand associated with Brahea armata (Arecaceae). This is the second record of this
species and the first record of D. heveae on a palm; the holotype was collected on rubber from the
same province (Senwanna et al. 2017). The phylogenetic results suggested that our strain is the
same species, D. heveae. The morphological characteristics largely resemble those of D. heveae
(e.g., the measurements of ascomata, ostiolar canals, peridia, asci, and ascospores, revealed
overlapping sizes). However, MFLU 15-0020 differs in having fusiform stromata while the type
specimen of D. heveae has rounded to irregular stromata and fewer ascomata within a stroma (1–2
versus 4–5) (Senwanna et al. 2017). The sequence data of LSU, SSU, ITS, tef1, RPB2, and TUB2
are almost identical to those of the ex-type.
Diatrypella elaeidis Konta & K.D. Hyde, sp. nov. Fig. 8
Index Fungorum number: IF556572, Facesoffungi number: FoF05122
Etymology – Refers to host genus, Elaeis
Holotype – MFLU 15-0025
257
Figure 7 – Diatrypella heveae (MFLU 15-0020) a Stromata on host substrate. b, c Close up of
stromata. d Section of stroma. e Ostiolar canal. f Peridium. g–n Asci. o–u Ascospores. v Colony on
MEA. Bars: a = 1000 μm, b, c = 500 μm, d = 50 μm, e, f, g–n = 20 μm, o–u = 5 μm.
Saprobic on petiole of Elaeis guineensis (Arecaceae). Sexual morph: Stromata 1025–3965
μm long, 285–860 μm wide (x
̄ = 1950 × 620 μm, n = 35), with well-developed interior, solitary to
gregarious, erumpent, black, glabrous, fusiform or lenticular in shape, pustulate, 1–2-ascomata.
Ascomata 242–600 μm high, 240–424 μm diam. (x
̄ = 350 × 335 μm, n = 25), perithecial, immersed
in the stroma, globose to subglobose, ostioles individual, with a short neck. Ostiolar canal 100–200
μm high, 70–190 μm diam. (x
̄ = 140 × 120 μm, n = 20), cylindrical, sulcate, with periphyses.
Peridium 22–55 μm wide (x
̄ = 33 μm, n = 45), composed of two layers, outer part of brown, thick-
walled cells, arranged in textura angularis, inner layers of hyaline, thick-walled cells of textura
angularis. Hamathecium composed of 2–8 μm wide (x
̄ = 4 μm, n = 70), filiform, longer than asci,
septate, hyaline paraphyses. Asci (excluding stalks), spore-bearing part (29–)40–70(−98) × (7–)11–
258
15(−17) μm (x
̄ = 60 × 12 μm, n = 70), apically rounded, with J-apical ring, apex-bearing part (0.5–
)1–3(−5) μm long (x
̄ = 2 μm, n = 35), polysporous, unitunicate, cylindrical, with moderately short
stalks, stalk-bearing part (18–)25–40(–57) μm long (x
̄ = 33 μm, n = 20). Ascospores (3.5–)5–7(−10)
× 1.5−3 μm (x
̄ = 6 × 2 μm, n = 130), overlapping, yellowish to brown, ellipsoidal to cylindrical or
elongate-allantoid, aseptate, smooth-walled. Asexual morph: Undetermined.
Geographical distribution – Thailand.
Culture characters – Ascospores germinated on MEA within 24 hours. Colonies on MEA
smooth, white, dense towards the edge, margin diffuse (Fig. 8r).
Additional sequence data – LSU (MN308408), SSU (MN308426), tef1 (MN525599), RPB2
(MN542891) (MFLUCC 15-0279).
Material examined – THAILAND, Chiang Rai Province, Mae Chan District, on dead petiole
of Elaeis guineensis Jacq. (Arecaceae), 25 November 2014, S. Konta, HR03a (MFLU 15-0025,
holotype); ex-type living culture = MFLUCC 15-0279.
Notes – Diatrypella elaeidis clearly differs from other species in the genus by its fusiform
stroma with 1–2-ascomata immersed in the stroma and their dimensions. Diatrypella elaeidis forms
a sister clade to D. delodicis with bootstrap support of 68% ML, 0.90 BYPP (Fig. 1). However, D.
elaeidis is distinguished from D. delodicis in having black long fusiform stroma, while, D.
delodicis has pale to dark brown globose to subglobose stroma with a flattened base. Additionally,
the number of perithecia per stroma, D. elaeidis is less than those in D. delodicis (1–2 vs 3–4), asci
and hamathecium of D. elaeidis are smaller than those of D. delodicis (asci; 61 × 12 μm vs 100 ×
18 μm, hamathecium 4 µm vs 9.8 μm wide) (Hyde et al. 2019). Diatrypella delodicis was found in
Chiang Rai on dried seed pods of Delonix regia (Fabaceae) (Hyde et al. 2019), while D. elaeidis
was collected from Elaeis guineensis. A comparison of the nucleotide of D. elaeidis to D. delodicis,
D. heveae, and D. verruciformis (type species) is given in Table 7.
Discussion
Diatrypaceae species are difficult to identify based on morphology due to overlapping
phenotypic characters (Glawe & Rogers 1984, Trouillas et al. 2011, Dayarathne et al. 2016,
Senwanna et al. 2017, Shang et al. 2017). Many studies have revealed several new hosts of species
and have contributed to our knowledge of their geographical distribution (Moyo et al. 2019). In this
study, we introduce a new genus and five new species for family Diatrypaceae based on
morphology combined with phylogeny. New hosts and new geographical distribution records are
also provided. This supports the high novelty of fungal species in Thailand in this family (Hyde et
al. 2018). The asexual morphs of Diatrypaceae are usually coelomycetes, but are not generally
useful in separating species (Glawe & Rogers 1982, 1984). Allocryptovalsa (Allocryptovalsa clade,
Fig. 1) appears to be a monophyletic group in Diatrypaceae, as was established in previous
phylogenetic studies (Senwanna et al. 2017, Hyde et al. 2019). Molecular data are available for A.
cryptovalsoidea, A. polyspora and A. rabenhorstii (Trouillas et al. 2011, Senwanna et al. 2017).
In our phylogenetic analyses, a novel genus Allodiatrype is introduced. Three novel taxa and
a novel combination grouped together but constitute separate lineages (Allodiatrype clade, Fig. 1).
The strains of Diatrypella form a well-resolved clade (Fig. 1). However, in this clade, some strains
of Diatrype (D. oregonensis, D. prominens) are placed between Diatrypella species. In same case,
some strains of Diatrypella species such as D. banksiae, D. decorticata, D. favacea and D.
iranensis often form distinct lineages within Diatrypaceae (Fig. 1). This may be due to lack of
TUB2 gene sequences or misidentified species. Hence, fresh collections and sequence data are
required to resolve their phylogenetic placement within the family.
Nine genera of Diatrypaceae out of 18 genera (Table 3) have been recorded from palms
worldwide (Arecaceae). Of these, seven genera and 20 species (Table 4) have been recorded in
Thailand. In addition, our taxa were isolated from dead parts of palms (Arecaceae). They were
collected in the same period (November to December 2014) but from different habitats. However,
our collection of diatrypaceous fungi are distributed in different hosts as well as different parts of
the palms (Arecaceae) and different locations. Moyo et al. (2019) have discussed these aspects and
259
Figure 8 – Diatrypella elaeidis (MFLU 15-0025, holotype) a Stromata on host substrate. b Close
up of stromata. c, d Sections of stromata. e, f Ostiolar canal. g Peridium. h–l Asci. m–q Ascospores.
r Colony on MEA. Scale bars: a, b = 1000 μm, c = 500 μm, d = 200 μm, e, f, h–l = 50 μm, g = 20
μm, m–q = 5 μm.
concluded that broad sampling across the globe might be required to fully comprehend the host
associations and distribution of Diatrypaceae.
Our analysis (Fig. 1) showed that Peroneutypa forms a monophyletic clade with high statistical
support (96% ML, 1.00 BYPP). A putatively named strain, Eutypa microasca (BAFC 51550) is
placed within this clade but without good statistical support, as shown in the original publication
and other studies (Grassi et al. 2014, Shang et al. 2018, Hyde et al. 2019, Phookamsak et al. 2019).
260
This is just one of many cases showing that the generic concepts of Diatrypaceae are in need of revision. In addition, many genera appear paraphyletic
or even polyphyletic in our current phylogeny based on the available material. This situation can only be changed by substantial amounts of fieldwork
aimed at collection and culturing of the “missing” taxa, along with careful morphological studies and multi-locus phylogenies.
Table 3 World distribution of Diatrypaceae on palms (Arecaceae).
Genera
Species name
Hosts
Countries
References
Allocryptovalsa
A. elaeidis Konta & K.D. Hyde.
Elaeis guineensis
Thailand
This study
Allodiatrype
A. arengae Konta & K.D. Hyde.
Arenga pinnata
Thailand
This study
A. elaeidicola Konta & K.D. Hyde.
Elaeis guineensis
Thailand
This study
A. elaeidis Konta & K.D. Hyde.
Elaeis guineensis
Thailand
This study
A. thailandica (R.H. Perera, Jian K. Liu & K.D. Hyde) Konta &
K.D. Hyde.
≡ Diatrype thailandica R.H. Perera, Jian K. Liu & K.D. Hyde
Calamus sp.
Thailand
This study
Anthostoma
A. cocois Höhn.
Cocos nucifera
Samoa
Dingley et al. (1981)
A. yatay Speg.
Cocos yatay
Argentina
Farr (1973)
Cryptovalsa
C. deusta (Ellis & G. Martin) Petr.
≡ Diatrypella deusta Ellis & G. Martin.
Sabal serrulata
China, U.S.A.
(Florida)
Cash (1952), Petrak (1953),
Teng (1996)
C. protracta (Pers.) De Not.
= Diatrypella nitschkei (Fuckel) L.C. Tiffany & J.C. Gilman.
Sabal palmetto
U.S.A. (Florida)
Petrak (1953)
Diatrype
D. chlorosarca Berk. & Broome.
Archontophoenix alexandrae,
Archontophoenix sp., Trachycarpus
fortunei, Trachycarpus sp.
China, Hong
Kong
Lu et al. (2000), Zhuang
(2001), Taylor & Hyde
(2003)
D. euterpes (Henn.) Rappaz.
≡ Eutypa euterpes Henn.
Euterpe oleracea
Brazil
Rappaz (1987), Mendes et
al. (1998)
D. palmarum Rick.
Phoenix sylvestris
India
Patil & Patil (1983)
D. palmicola Jian K. Liu & K.D. Hyde.
Caryota urens
Thailand
Liu et al. (2015)
Diatrype sp.
Rhopalostylis sp.
New Zealand
McKenzie et al. (2004)
Diatrypella
D. heveae Senwanna, Phookamsak & K.D. Hyde.
Brahea armata
Thailand
This study
D. borassi Chona & Munjal.
Trachycarpus fortunei,
Archontophoenix alexandrae,
Archontophoenix sp.
Australia, Hong
Kong
Lu et al. (2000), Zhuang
(2001), Taylor & Hyde
(2003)
D. caryotae R.K. Verma.
Caryota urens
India
Verma (1996)
D. elaeidis Konta & K.D. Hyde.
Elaeis guineensis
Thailand
This study
D. tuberculata Ellis & Catkins ex Ellis & Everh.
Sabal serrulata
U.S.A. (Florida)
Cash (1952)
Diatrypella sp.
Rhopalostylis sp.
New Zealand
McKenzie et al. (2004)
Eutypa
E. rattanicola J. Fröhl. & K.D. Hyde.
Calamus moti
Australia
Fröhlich & Hyde (2000)
Eutypella
E. arecae (Syd. & P. Syd.) Rappaz.
Areca catechu, Calamus tetradactylus,
Calamus sp., Trachycarpus fortunei
China, Hong
Kong,
Rappaz (1987), Fröhlich &
Hyde (2000), Lu et al. (2000),
261
Table 3 Continued.
Genera
Species name
Hosts
Countries
References
Philippines,
Switzerland
Zhuang (2001), Taylor &
Hyde (2003)
E. rehmiana (Henn.) Höhn.
Areca catechu, Areca sp., Calamus sp.
Philippines
Reinking (1918, 1919),
Teodoro (1937)
E. sabalina Cooke.
Arecaceae, Chamaerops humilis, Sabal
minor, S. palmetto (≡ Sabal
blackburniana), Sabal sp.
Bermuda, China,
Georgia, U.S.A.
(Alabama, Florida,
Louisiana)
Vizioli (1923),
Anonymous (1960), Tai
(1979), Alfieri et al.
(1984), Rappaz (1987),
Teng (1996), Glawe &
Jones (1989), Zhuang
(2001)
Eutypella sp.
Rhopalostylis sp.
New Zealand
McKenzie et al. (2004)
Peroneutypa
Peroneutypa sp.
Cocos nucifera
Cuba
Urtiaga (1986)
Table 4 Distribution of diatrypaceous fungi on plants in Thailand.
Genera Species name Hosts Collection site
(Province) Collection date References
Allocryptovalsa
A. polyspora Senwanna, Phookamsak & K.D. Hyde.
Dead twig of Hevea brasiliensis
Phayao
29 January 2017
Senwanna et al. (2017)
A. elaeidis Konta & K.D. Hyde.
Dead petiole of Elaeis guineensis
Krabi
3 December 2014
This study
Allodiatrype
A. arengae Konta & K.D. Hyde.
Dead petiole of Arenga pinnata
Phang-Nga
4 December 2014
This study
A. elaeidicola Konta & K.D. Hyde.
Dead petiole of Elaeis guineensis
Phang-Nga
5 December 2014
This study
A. elaeidis Konta & K.D. Hyde.
Dead petiole of Elaeis guineensis
Krabi
3 December 2014
This study
A. thailandica (R.H. Perera, Jian K. Liu & K.D. Hyde)
Konta & K.D. Hyde.
≡ Diatrype thailandica R.H. Perera, Jian K. Liu &
K.D. Hyde
Dead petiole of Calamus sp.
Stems of unidentified plant
Phang-Nga
Chiang Rai
6 December 2014
12 March 2015
This study
Li et al. (2016)
Diatrype
D. palmicola Jian K. Liu & K.D. Hyde.
Dead branch of Caryota urens
Chiang Rai
6 September 2010
Liu et al. (2015)
Diatrypella
D. tectonae M. Doilom, Q.J. Shang & K.D. Hyde.
Dead branch of Tectona grandis
Chiang Rai
5 February 2012
Shang et al. (2017)
D. heveae Senwanna, Phookamsak & K.D. Hyde.
Dead twig of Hevea brasiliensis
Dead petiole of Brahea armata
Chiang Rai
Chiang Rai
1 November 2016
25 November 2014
Senwanna et al. (2017)
This study
D. vulgaris Trouillas, W.M. Pitt & Gubler.
Stems of unidentified plant
Chiang Rai
1 January 2015
Hyde et al. (2017)
D. delonicis R.H. Perera & K.D. Hyde.
Dried seed pods of Delonix regia
Chiang Rai
10 December 2014
Hyde et al. (2019)
D. elaeidis Konta & K.D. Hyde.
Dead petiole of Elaeis guineensis
Chiang Rai
25 November 2014
This study
Eutypa
E. flavovirens (Pers.) Tul. & C. Tul.
Decaying twigs
Chiang Rai
15 November 2012
Senanayake et al. (2015)
262
Table 4 Continued.
Genera Species name Hosts
Collection site
(Province)
Collection date References
Halodiatrype
H. avicenniae Dayarathne & K.D. Hyde.
Intertidal decayed wood of
Avicennia sp. at a mangrove
stand
Phetchaburi
28 August 2015
Dayarathne et al. (2016)
H. salinicola Dayarathne & K.D. Hyde.
Submerged marine wood
Phang-Nga
7 December 2014
Dayarathne et al. (2016)
Peroneutypa
P. diminutiasca Q.J. Shang, Phookamsak & K.D.
Hyde.
Undetermined deadwood
Chiang Mai
27 January 2017
Shang et al. (2018)
P. longiasca Senwanna, Phookamsak & K.D. Hyde.
Dead twig of Hevea brasiliensis
Chiang Rai
1 November 2016
Senwanna et al. (2017)
P. mackenziei Q.J. Shang, Phookamsak & K.D. Hyde.
Undetermined decaying wood
Chiang Rai
22 January 2015
Shang et al. (2017)
P. rubiformis Q.J. Shang, Phookamsak & K.D. Hyde.
Undetermined deadwood
Chiang Mai
27 January 2017
Shang et al. (2018)
P. scoparia Carmarán & A.I. Romero.
Dead culms of bamboo
Dead culms of bamboo
Undetermined deadwood
Dieback diseased marine wood
Chiang Rai
Chiang Rai
Chiang Mai
Phetchaburi
16 July 2011
11 August 2011
27 January 2017
11 January 2018
Dai et al. (2016)
Dai et al. (2016)
Shang et al. (2018)
Hyde et al. (2019)
Table 5 Comparison of the nucleotides of Allocryptovalsa elaeidis to A. polyspora, A. cryptovalsoidea and A. rabenhorstii.
Allocryptovalsa spp.
LSU
SSU
ITS
TUB2
RBP2
tef1
References
A. polyspora MFLUCC17-0364T
1/823 (0.12%)
-
3/486 (0.62%)
12/516 (2.32%)
-
-
Senwanna et al. (2017)
A. cryptovalsoidea HVFIG02
-
-
4/488 (0.81%)
0
-
-
Trouillas et al. (2011)
A. rabenhorstii WA07CO
-
-
23/560 (4.1%)
27/369 (7.31%)
-
-
Trouillas et al. (2011)
Notes – ‘-’ do not have sequence; ‘0’ no base pair similarity; T type species; base pair differences included gaps.
Table 6 Comparison of the nucleotides of Allodiatrype arengae to all species of Allodiatrype, and type species of the genus Diatrype.
Species
LSU
SSU
ITS
TUB2
RBP2
tef1
References
A. elaeidicola MFLUCC 15-0737
0
1/1030 (0.09%)
13/605 (2.14%)
28/1586 (1.76%)
3/1139 (0.26%)
0
This study
A. elaeidis MFLUCC 15-0708
0
0
13/621 (2.09%)
16/1579 (1.01%)
2/1138 (0.17%)
0
This study
A. thailandica MFLUCC 14-1210
-
-
9/526 (1.71%)
-
-
-
Li et al. (2016)
A. thailandica MFLUCC 15-0711
0
9/1040 (0.86%)
11/619 (1.77%)
-
-
-
This study
Diatrype disciformis T
21/905 (2.32%)
2/894 (0.22%)
56/536 (10.44%)
-
314/1137 (27.6%)
69/957 (7.2%)
Acero et al. (2004)
Notes – ‘-’ do not have sequence; ‘0’ no base pair similarity; T type species of Diatrype; base pair differences included gaps.
263
Table 7 Comparison of the nucleotides of Diatrypella elaeidis to D. delodicis, D. heveae, and D. verruciformis.
Diatrypella spp.
LSU
SSU
ITS
TUB2
RBP2
tef1
References
D. delonicis MFLUCC 15-1014
-
-
1/600 (0.16%)
7/370 (1.89%)
-
-
Hyde et al. (2019)
D. heveae MFLUCC 15-0274
1/889 (0.11%)
6/1036 (0.57%)
10/603 (1.65%)
15/810 (1.85%)
26/800 (3.25%)
9/954 (0.94%)
This study
D. verruciformis UCROK754
-
-
15/607 (2.47%)
35/367 (9.53%)
-
-
Lynch et al. (2013)
Notes – ‘-’ do not have sequence; ‘0’ no base pair similarity; base pair differences included gaps.
Acknowledgments
We are grateful to the National Research Council of Thailand (project no. 61215320023), the Mushroom Research Foundation and the Thailand
Research Fund entitled. The authors extend their appreciation to The Researchers supporting project number (RSP-2019/56) King Saud University,
Riyadh, Saudi Arabia. Kevin D. Hyde thanks The future of specialist fungi in a changing climate: baseline data for generalist and specialist fungi
associated with ants, Rhododendron species and Dracaena species (grant number: DBG6080013) and Impact of climate change on fungal diversity
and biogeography in the Greater Mekong Subregion (grant number: RDG6130001). Saranyaphat Boonmee would like to thank the Thailand Research
Fund (No. TRG6180001), Plant Genetic Conservation Project under the Royal Initiation of Her Royal Highness Princess Maha Chakri Sirindhorn-Mae
Fah Luang University and Mae Fah Luang University grant Number 631C15001. Rungtiwa Phookamsak thanks CAS President’s International
Fellowship Initiative (PIFI) for young staff (grant no. 2019FYC0003), The Yunnan Provincial Department of Human Resources and Social Security
(grant no. Y836181261), and National Science Foundation of China (NSFC) project code 31850410489 for financial support. Sirinapa Konta is
grateful to Ausana Mapook, Dr. Paul Kirk, Dr. Shaun Pennycook, Dr. Saowaluck Tibpromma, Dr. Mingkwan Doilom, Li Junfu, Dr. Samantha C.
Karunarathna, Yuanpin Xiao, Li Wenjing, Sirilak Radbouchoom, Zeng Ming, Qiuju Shang, Monika C. Dayarathne, and Milan C. Samarakoon for their
valuable suggestion and help.
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