life
Article
Multi-Gene Phylogeny and Morphology Reveal
Haplohelminthosporium gen. nov. and Helminthosporiella gen.
nov. Associated with Palms in Thailand and A Checklist for
Helminthosporium Reported Worldwide
Sirinapa Konta 1,2,3 , Kevin D. Hyde 1,2 , Samantha C. Karunarathna 1 , Ausana Mapook 2 ,
Chanokned Senwanna 4 , Lucas A. P. Dauner 1 , Chandrika M. Nanayakkara 5 , Jianchu Xu 1 ,
Saowaluck Tibpromma 1, * and Saisamorn Lumyong 6,7, *
1
Citation: Konta, S.; Hyde, K.D.;
Karunarathna, S.C.; Mapook, A.;
Senwanna, C.; Dauner, L.A.P.;
Nanayakkara, C.M.; Xu, J.;
2
3
4
5
6
Tibpromma, S.; Lumyong, S.
Multi-Gene Phylogeny and
7
Morphology Reveal
*
CAS Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany,
Chinese Academy of Sciences, Kunming 650201, China; sirinapakonta@gmail.com (S.K.);
kdhyde3@gmail.com (K.D.H.); samantha@mail.kib.ac.cn (S.C.K.); luke.dauner1@gmail.com (L.A.P.D.);
jxu@mail.kib.ac.cn (J.X.)
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand;
phung.ausana@gmail.com
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University,
Chiang Mai 50200, Thailand; chanokned.swn@gmail.com
Department of Plant Sciences, University of Colombo, Colombo 00300, Sri Lanka; chandi@pts.cmb.ac.lk
Research Center of Microbial Diversity and Sustainable Utilization, Faculty of Science, Chiang Mai University,
Chiang Mai 50200, Thailand
Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand
Correspondence: saowaluckfai@gmail.com (S.T.); saisamorn.l@cmu.ac.th (S.L.)
Haplohelminthosporium gen. nov. and
Helminthosporiella gen. nov.
Associated with Palms in Thailand
and A Checklist for Helminthosporium
Reported Worldwide. Life 2021, 11,
454. https://doi.org/10.3390/
life11050454
Academic Editor: Arnold J.
M. Driessen
Received: 16 March 2021
Accepted: 11 May 2021
Published: 19 May 2021
Publisher’s Note: MDPI stays neutral
Abstract: Palms (Arecaceae) are substrates for a highly diverse range of fungi. Many species are
known as saprobes and many are important plant pathogens. Over the course of our studies of microfungi from palms in Thailand, two new taxa were discovered. Morphological characteristics and
phylogenetic analyses of combined ITS, LSU, SSU, and tef1-α sequence data revealed their taxonomic
positions within Massarinaceae. There are currently ten genera identified and accepted in Massarinaceae, with the addition of the two new genera of Haplohelminthosporium and Helminthosporiella, that
are introduced in this paper. Each new genus is provided with a full description and notes, and each
new taxon is provided with an illustration for the holotype. A list of identified and accepted species
of Helminthosporium with morphology, host information, locality, sequence data, and related references of Helminthosporium reported worldwide is provided based on records in Species Fungorum
2021. This work provides a micro-fungi database of Haplohelminthosporium, Helminthosporiella, and
Helminthosporium which can be modified and validated as new data come to light.
Keywords: 4 new taxa; Massarinaceae; morphology; multi-genes; palm fungi; Thailand
with regard to jurisdictional claims in
published maps and institutional affiliations.
1. Introduction
Copyright: © 2021 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
In Thailand, a large number of novel fungi from a variety of hosts have been recently
described, adding to the region’s highly known fungal diversity [1,2]. This diversity
is supported by various factors, including host–plant species relationships, geography,
seasons, air humidity, and temperature. Many interesting fungi from Thai monocotyledons
such as bamboo (Poaceae) and Pandanaceae have been described in previous studies, and
some new taxa and records of microfungi on palms have been published, especially from
the southern region of Thailand [3–11]. However, more research on fungal diversity on
palms in Thailand is needed.
4.0/).
Life 2021, 11, 454. https://doi.org/10.3390/life11050454
https://www.mdpi.com/journal/life
Life 2021, 11, 454
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Pleosporales is the largest order in Dothideomycetes [12] with 566 genera in 91 families accepted, while 48 genera have been placed in Pleosporales genera incertae sedis with
an estimated stem age of 205 MYA [12,13]. Massarinaceae is a family within Pleosporales introduced by Munk [14] to accommodate the genus Massarina, with M. eburnea
being designated as the type species and described based on the sexual morph [15].
Hongsanan et al. [12] and Wijayawardene et al. [13] accepted nine genera in Massarinaceae
(Byssothecium, Helminthosporiella, Helminthosporium, Massarina, Pseudodidymosphaeria, Pseudosplanchnonema, Semifissispora, Stagonospora, and Suttonomyces).
Helminthosporium has the asexual morph of H. velutinum as the type species. It is
characterized by terminal and intercalary conidiogenous cells as well as solitary conidia with distosepta [16]. The members of this genus are commonly found as saprobes
and endophytes, but they are often isolated from dead corticated twigs or wood, living leaves, and soils [17–23]. Most Helminthosporium species have been described based
on their asexual morph, and only a few species have been described based on both
morphs viz., H. massarinum, H. microsorum, H. oligosporum, H. quercicola, H. quercinum,
and H. tiliae [19,21,24]. Several species in the Helminthosporium complex are polyphyletic
and have been placed in other genera viz. Bipolaris, Curvularia, and Exserohilum within
Pleosporales, other families viz. Corynesporaceae, Massarinaceae, and Mycosphaerellaceae
within Dothideomycetes, or other unrelated Ascomycetes groups that were initially based
on morphological characteristics and later on molecular data, although some species still
remain unresolved [20,25–37]. Wijayawardene et al. [13] approximated the number of taxa
in Helminthosporium at 416 species. However, this genus was not updated with the DNA
sequencesin the most recent monograph.
Few previous studies have investigated the Helminthosporium-like taxa from plants,
particularly palms, in Thailand. In this study, we were able to isolate Helminthosporiumlike taxa from palms collected in Thailand. Morphology and multi-gene phylogenetic
analyses showed two Helminthosporium-like taxa are novel in Massarinaceae. In addition,
we provide a checklist of Helminthosporium and the name for Helminthosporiella stilbacea is
also validated.
2. Materials and Methods
2.1. Collection, Isolation, and Identification
The plant materials containing the fungal structures were collected from Krabi and
Prachuap Khiri Khan Provinces, Thailand, from living and dead parts of palm trees (Calamus sp. and Cocos nucifera). Samples were taken to the laboratory for morphological study
following the methods provided by Konta et al. [9]. Single spore isolates were obtained
following the method of Senanayake et al. [38]. Measurements were taken using an Image
Framework program. Illustrations were made in Adobe Photoshop CS6. Specimens and
cultures were deposited in the herbarium of Mae Fah Luang University (MFLU) and Mae
Fah Luang Culture Collection (MFLUCC). Faces of Fungi and Index Fungorum numbers
were registered as outlined in Jayasiri et al. [39] and Index Fungorum [40], respectively.
2.2. DNA Extraction and Amplification (PCR)
DNA extraction was performed using the Biospin Fungus genomic DNA extraction
kit-BSC14S1 (Bioflux, P.R. China) according to Dissanayake et al. [41]. Partial nucleotide
genes were subjected to PCR amplification and sequencing of the large subunit (28S,
LSU) [42], the internal transcribed spacer (ITS) [43], the small subunit (18S, SSU) [43], and
the translation elongation factor 1-alpha (tef1-α) was performed [44,45]. For primers and
conditions, see Table 1. PCR amplification and sequencing were carried out following
Konta et al. [9]. The resulting fragments were sequenced in both forward and reverse
directions, the generated DNA sequences were analysed, and the consensus sequences
were computed using SeqMan software. New sequences generated in this study were
deposited in GenBank (Table 2).
Life 2021, 11, 454
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Table 1. Details of genes/loci with PCR primers and PCR conditions.
Genes/loci
LSU
ITS
SSU
tef1-α
a
PCR Primer
(Forward/Reverse)
LR0R/LR5
ITS5/ITS4
NS1/NS4
983F/2218R
PCR Conditions
a;
95 ◦ C: 30 s, 55 ◦ C: 50 s, 72 ◦ C: 30 s (35 cycles); b
Initiation step of 95 ◦ C: 3 min; b Final elongation step of 72 ◦ C: 10 min and final hold at 4 ◦ C.
2.3. Phylogenetic Analyses
The sequences generated in this study were subjected to a BLAST search in GenBank
to identify closely related sequences. Sequence data retrieved from GenBank and recent
publications were used as references [24]. Sequence data for the ITS, LSU, SSU, and
tef1-α regions were analysed both individually and in combination. A total of 93 taxa
were used for the combined phylogenetic analyses (ITS, LSU, SSU, and tef1-α) in order to
find a natural classification placement. In addition, 103 taxa of ITS and 113 taxa of LSU
were used for phylogenetic analyses. For both the individual and combined phylogenetic
analyses, Cyclothyriella rubronotata (Cyclothyriellaceae) was selected as the outgroup taxon.
Absent sequence data (i.e., ITS, LSU, SSU, tef1-α sequence data) in the alignments were
treated with gaps as missing data. Sequence alignments were carried out with MAFFT
v.6.864b [46] and were manually improved where necessary. The single gene datasets were
combined using Mega7 [47]. Data were converted from fasta to nexus and PHYLIP format
with Alignment Transformation Environment online, https://sing.ei.uvigo.es/ALTER/
(accessed on 15 July 2020) [48]. The tree topologies obtained from single gene sequence
data were compared prior to the combined gene analysis in order to check for incongruence
in the overall topology of the phylogenetic tree. Maximum likelihood (ML) analysis was
accomplished using RAxML-HPC2 (v.8.2.12) on XSEDE in the CIPRES Science Gateway
platform (http://www.phylo.org) (accessed on 12 May 2020) [49] with GTRGAMMA
model and set as 1000 bootstrap replicates. Bayesian analysis was performed at CIPRES
using Bayesian analysis on XSEDE (v.3.2.7) as part of the “MrBayes on XSEDE” tool [49–51].
GTR+I+G model was selected by using MrModelTest 2.2 [52] under the Akaike information
criterion (AIC) as the best-fit models of the combined dataset for maximum likelihood and
Bayesian analysis [52]. Bayesian posterior probabilities (BYPP) were determined by Markov
Chain Monte Carlo sampling (MCMC) in MrBayes on XSEDE v.3.2.7. Six simultaneous
Markov chains were run for 5,000,000 generations and trees were sampled every 1000th
generation. An 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 (7502) were used for calculating posterior probabilities in
the majority rule consensus tree. Bootstrap support values for ML and BYPP are given
near to each node (Figures 1 and 2). The phylogenetic trees were configured in FigTree
v1.4.0 [53] and edited using Microsoft Office PowerPoint 2016 and Adobe Photoshop CS6
(Adobe Systems, San Jose, CA, USA).
Life 2021, 11, x FOR PEER REVIEW
Life 2021, 11, 454
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Figure 1. Comparison of the topology of Maximum likelihood majority rule consensus tree for the analyses of some selected
Figure 1. Comparison of the topology of Maximum likelihood majority rule consensus tree for the analyses of some
Corynesporaceae,
Massarinaceae, Massarinaceae,
and Perioconiaceae
(A) Phylogenetic
tree
of the dataset
selected Corynesporaceae,
and isolates.
Perioconiaceae
isolates. (A)
Phylogenetic
treefor
of ITS
the sequence
dataset fordata.
ITS
(B) Phylogenetic
tree
of
the
dataset
for
LSU
sequence
data.
Bootstrap
support
values
for
maximum
likelihood
equal
sequence data. (B) Phylogenetic tree of the dataset for LSU sequence data. Bootstrap support values for(ML)
maximum
to or higher
than (ML)
50%, and
Posterior
Probabilities
(BYPP)
equal to
or greater than
0.90equal
are given
above each
likelihood
equalBayesian
to or higher
than 50%,
and Bayesian
Posterior
Probabilities
(BYPP)
to or greater
than
are given
above
each branch.
blue.The
Ex-type
are
bold. The tree
is rooted to
Cyclothybranch.0.90
Novel
taxa are
in blue.
Ex-typeNovel
strainstaxa
areare
in in
bold.
tree isstrains
rooted
toinCyclothyriella
rubronotata
strains
TR,
riella rubronotata strains TR, TR9 (Cyclothyriellaceae).
TR9 (Cyclothyriellaceae).
Life 2021, 11, x FOR PEER REVIEW
Life 2021, 11, 454
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5 of 57
Figure 2. Maximum
likelihood majority
rule
consensus
for the tree
analyses
of Massarinaceae
and sister family
PeriocoFigure 2. Maximum
likelihood
majority
rule tree
consensus
for the
analyses of Massarinaceae
and sister
family
sequence
data.
Bootstrap
support
values
for
niaceae isolates
based
on
a
dataset
of
combined
ITS,
LSU,
SSU,
and
tef1-α
Perioconiaceae isolates based on a dataset of combined ITS, LSU, SSU, and tef1-α sequence data. Bootstrap support
maximum likelihood
equal to
or higher(ML)
than 50%,
Bayesian
probabilities
(BYPP)
equal to
or greater than
values for(ML)
maximum
likelihood
equaland
to or
higherposterior
than 50%,
and Bayesian
posterior
probabilities
(BYPP)
equal
to oreach
greater
than Novel
0.90 are
given
each
branch.
Novel
taxa
are inThe
blue.
Ex-type
strains
are in bold. The
0.90 are given
above
branch.
taxa
are above
in blue.
Ex-type
strains
are
in bold.
tree
is rooted
to Cyclothyriella
is rooted
to Cyclothyriella
rubronotata strains TR, TR9 (Cyclothyriellaceae).
rubronotata tree
strains
TR, TR9
(Cyclothyriellaceae).
Life 2021, 11, 454
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Table 2. Taxa names, strain numbers and GenBank accession numbers of the sequences used in phylogenetic analyses.
Family
GenBank Accession No.
Species
Strain No.
References
ITS
LSU
SSU
tef1-α
-
GU301808
GU296144
GU349052
[54]
KF810854
-
GU296145
-
[54,55]
Corynesporaceae
Corynespora cassiicola
CBS 100,822
Corynesporaceae
Corynespora cassiicola
CCP
Corynesporaceae
Corynespora smithii
CBS 139,925
KY984299
KY984299
-
-
[21]
Corynesporaceae
Corynespora smithii
L120
KY984297
KY984297
-
KY984435
[21]
Corynesporaceae
Corynespora smithii
L130
KY984298
KY984298
KY984419
KY984436
[21]
Corynesporaceae
Corynespora smithii
L139
KY984300
KY984300
-
-
[21]
Cyclothyriellaceae
Cyclothyriella rubronotata
TR
KX650541
KX650541
-
KX650516
[56]
Cyclothyriellaceae
Cyclothyriella rubronotata
TR9 *
KX650544
KX650544
KX650507
KX650519
[56]
Massariaceae
Byssothecium circinans
CBS 675.92
-
GU205217
GU205235
GU349061
[54]
Massarinaceae
Byssothecium circinans
CBS 675.92
-
AY016357
AY016339
-
[57,58]
Massarinaceae
Haplohelminthosporium calami
MFLUCC 18-0074 *
MT928158
MT928156
MT928160
-
This study
Massarinaceae
Helminthosporium aquaticum
MFLUCC 15-0357
KU697302
KU697306
KU697310
-
[20]
Massarinaceae
Helminthosporium aquaticum
DLUCC 0758
MG098779
MG098786
MG098795
MG98585
[24]
Massarinaceae
Helminthosporium austriacum
L132 *
KY984301
KY984301
KY984420
KY984437
[21]
Massarinaceae
Helminthosporium austriacum
L169
KY984303
KY984303
-
KY984439
[21]
Massarinaceae
Helminthosporium austriacum
L137
KY984302
KY984302
-
KY984438
[21]
Massarinaceae
Helminthosporium caespitosum
L99 *
JQ044429
JQ044448
KY984421
KY984440
[21]
Massarinaceae
Helminthosporium caespitosum
L141
KY984305
KY984305
-
-
[21]
Massarinaceae
Helminthosporium caespitosum
L151
KY984306
KY984306
-
-
[21]
Massarinaceae
Helminthosporium dalbergiae
H 4628
LC014555
AB807521
AB797231
AB808497
[19]
Massarinaceae
Helminthosporium endiandrae
CBS 138902 *
KP004450
KP004478
-
-
[59]
Massarinaceae
Helminthosporium endiandrae
CBS 138,902
-
MH878637
-
-
[60]
Life 2021, 11, 454
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Table 2. Cont.
Family
GenBank Accession No.
Species
Strain No.
ITS
LSU
SSU
tef1-α
References
Massarinaceae
Helminthosporium endiandrae
SM64
MT279335
-
-
-
Unpublished
Massarinaceae
Helminthosporium endiandrae
SM61
MT279339
-
-
-
Unpublished
Massarinaceae
Helminthosporium endiandrae
SM64
MT279340
-
-
-
Unpublished
Massarinaceae
Helminthosporium endiandrae
SM61
MT279336
-
-
-
Unpublished
Massarinaceae
Helminthosporium endiandrae
AKRM1
MN880136
-
-
-
Unpublished
Massarinaceae
Helminthosporium erythrinicola
CBS 145,569
MK876391
MK876432
-
-
[22]
Massarinaceae
Helminthosporium genistae
L128
KY984308
KY984308
KY984422
-
[21]
Massarinaceae
Helminthosporium genistae
L129
KY984309
KY984309
KY984423
-
[21]
Massarinaceae
Helminthosporium genistae
L142 *
KY984310
KY984310
-
-
[21]
Massarinaceae
Helminthosporium hispanicum
L109 *
KY984318
KY984318
KY984424
KY984441
[21]
Massarinaceae
Helminthosporium italicum
MFLUCC 17-0241
KY797638
KY815015
-
KY815021
[61]
Massarinaceae
Helminthosporium juglandinum
L97
KY984322
KY984322
KY984425
KY984445
[21]
Massarinaceae
Helminthosporium juglandinum
L118 *
KY984321
KY984321
-
KY984444
[21]
Massarinaceae
Helminthosporium leucadendri
CBS 135133 *
KF251150
KF251654
-
KF253110
[62]
Massarinaceae
Helminthosporium magnisporum
H 4627 *
AB811452
AB807522
AB797232
AB808498
[19]
Massarinaceae
Helminthosporium massarinum
KT 1564 *
AB809629
AB807524
AB797234
AB808500
[19]
Massarinaceae
Helminthosporium massarinum
KT 838
AB809628
AB807523
AB797233
AB808499
[19]
Massarinaceae
Helminthosporium microsorum
L94
KY984327
KY984327
KY984426
KY984446
[21]
Massarinaceae
Helminthosporium microsorum
L95
KY984328
KY984328
-
KY984447
[21]
Massarinaceae
Helminthosporium microsorum
L96 *
KY984329
KY984329
KY984427
KY984448
[21]
Massarinaceae
Helminthosporium oligosporum
L92
KY984332
KY984332
KY984428
KY984450
[21]
Massarinaceae
Helminthosporium oligosporum
L93 *
KY984333
KY984333
-
KY984451
[21]
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Table 2. Cont.
Family
GenBank Accession No.
Species
Strain No.
ITS
LSU
SSU
tef1-α
References
Massarinaceae
Helminthosporium oligosporum
L106
KY984330
KY984330
-
KY984449
[21]
Massarinaceae
Helminthosporium quercinum
L90 *
KY984339
KY984339
KY984429
KY984453
[21]
Massarinaceae
Helminthosporium quercinum
L91
KY984340
KY984340
-
KY984454
[21]
Massarinaceae
Helminthosporium solani
CBS 365.75
KY984341
KY984341
KY984430
KY984455
[21]
Massarinaceae
Helminthosporium solani
CBS 640.85
KY984342
KY984342
-
-
[21]
Massarinaceae
Helminthosporiella stilbacea
CPHmZC-01
KX228298
KX228355
-
-
[63]
Massarinaceae
Helminthosporiella stilbacea
COAD 2126
MG668862
-
-
-
[64]
Massarinaceae
Helminthosporiella stilbacea
MFLUCC 15-0813 *
MT928159
MT928157
MT928161
MT928151
This study
Massarinaceae
Helminthosporium submersum
MFLUCC 16-1360 *
-
MG098787
MG098796
MG098586
[24]
Massarinaceae
Helminthosporium submersum
MFLUCC 16-1290
MG098780
MG098788
MG098797
MG098587
[24]
Massarinaceae
Helminthosporium submersum
DLUCC 0805
MG098781
MG098789
MG098798
-
[24]
Massarinaceae
Helminthosporium syzygii
CBS 145,570 *
MK876392
MK876433
-
-
[22]
Massarinaceae
Helminthosporium tiliae
L88 *
KY984345
KY984345
KY984431
KY984457
[21]
Massarinaceae
Helminthosporium tiliae
L89
KY984346
KY984346
-
-
[21]
Massarinaceae
Helminthosporium tiliae
L171
KY984343
KY984343
-
KY984456
[21]
Massarinaceae
Helminthosporium velutinum
yone 38
-
AB807527
AB797237
AB808502
[19]
Massarinaceae
Helminthosporium velutinum
yone 63
-
AB807528
AB797238
AB808503
[19]
Massarinaceae
Helminthosporium velutinum
MFLUCC 15-0423
KU697300
KU697304
KU697308
-
[20]
Massarinaceae
Helminthosporium velutinum
MFLUCC 15-0428
KU697299
KU697303
KU697307
-
[20]
Massarinaceae
Helminthosporium velutinum
H 4626
LC014556
AB807530
AB797240
AB808505
[19]
Massarinaceae
Helminthosporium velutinum
L117
KY984349
KY984349
-
KY984460
[21]
Massarinaceae
Helminthosporium velutinum
L126
KY984350
KY984350
-
KY984461
[21]
Massarinaceae
Helminthosporium velutinum
L131 *
KY984352
KY984352
KY984432
KY984463
[21]
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Table 2. Cont.
Family
GenBank Accession No.
Species
Strain No.
ITS
LSU
SSU
tef1-α
References
Massarinaceae
Helminthosporium velutinum
CPC 26297= CBS 141,504
KX306757
KX306785
-
-
[65]
Massarinaceae
Helminthosporium velutinum
yone 96
LC014558
AB807529
AB797239
AB808504
[19]
Massarinaceae
Helminthosporium velutinum
H 4739
LC014557
AB807525
AB797235
AB808501
[19]
Massarinaceae
Helminthosporium velutinum
L115
KY984347
KY984347
-
KY984458
[21]
Massarinaceae
Helminthosporium velutinum
L116
KY984348
KY984348
-
KY984459
[21]
Massarinaceae
Helminthosporium velutinum
L127
KY984351
KY984351
-
KY984462
[21]
Massarinaceae
Helminthosporium velutinum
L98
KY984359
KY984359
KY984433
KY984466
[21]
Massarinaceae
Helminthosporium velutinum
H 4743
-
AB807526
AB797236
-
[19]
Massarinaceae
Helminthosporium velutinum
MFLUCC 16-1096
MG098783
MG098791
MG098799
MG098588
[24]
Massarinaceae
Helminthosporium velutinum
MFLUCC 16-1282
MG098784
MG098792
MG098800
MG098589
[24]
Massarinaceae
Helminthosporium velutinum
MFLUCC 17-1707
MG098785
MG098793
MG098801
MG098590
[24]
Massarinaceae
Helminthosporium velutinum
MFLUCC 17-1321
-
MG098794
MG098802
MG098591
[24]
Massarinaceae
Helminthosporium velutinum
S-076
KU697301
KU697305
KU697309
-
[20]
Massarinaceae
Helminthosporium velutinum
MFLUCC 15-0243
KU697301
KU697305
KU697309
-
[20]
Massarinaceae
Helminthosporium velutinum
MFLUCC 16-1300
MG098782
MG098790
-
-
[24]
Massarinaceae
Massarina albocarnis
CBS119345
LC194503
LC194379
LC194337
LC194416
[66]
Massarinaceae
Massarina cisti
CBS 266.62 *
LC014568
AB807539
AB797249
AB808514
[19]
Massarinaceae
Massarina cisti
CBS 266.62
-
FJ795447
FJ795490
-
[67]
Massarinaceae
Massarina eburnea
CBS 473.64
AF383959
GU301840
AF164367
-
[60,68]
Massarinaceae
Massarina eburnea
JCM 14422
LC014569
AB521735
AB521718
AB808517
[19]
Massarinaceae
Massarina igniaria
CBS 845.96
-
FJ795452
FJ795494
-
[67]
Massarinaceae
Massarina pandanicola
MFLUCC 17-0596
MG646958
MG646947
MG646979
MG646986
[4]
Massarinaceae
Massarina phragmiticola
CBS 110,446
-
DQ813510
DQ813512
-
[69]
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Table 2. Cont.
Family
GenBank Accession No.
Species
Strain No.
ITS
LSU
SSU
tef1-α
References
Massarinaceae
Neottiosporina paspali
CBS 331.37
-
EU754172
EU754073
-
[70]
Massarinaceae
Pseudodidymosphaeria spartii
CBS 183.58
-
GU205225
GU205250
-
[71]
Massarinaceae
Pseudodidymosphaeria spartii
MFLUCC 13-0273
KP325434
KP325436
KP325438
-
[72]
Massarinaceae
Pseudodidymosphaeria spartii
MFLUCC 14-1212
KP325435
KP325437
KP325439
-
[72]
Massarinaceae
Pseudosplanchnonema phorcioides
MFLUCC 14-0618
KP683372
KP683373
KP683374
-
[72]
Massarinaceae
Pseudosplanchnonema phorcioides
MFLUCC 13-0533
-
KM875454
KM875455
-
[73]
Massarinaceae
Pseudosplanchnonema phorcioides
L16
KY984360
-
KY984434
KY984467
[21]
Massarinaceae
Pseudosplanchnonema phorcioides
MFLUCC 13-0611
KP683375
KP683376
KP683377
-
[21]
Massarinaceae
Semifissispora natalis
CPC 25383
KT950846
KT950858
-
KT950878
[21]
Massarinaceae
Semifissispora natalis
CBS 140659
-
MH878157
-
-
[21]
Massarinaceae
Semifissispora rotundata
CPC 549
KT950847
KT950859
-
-
[21]
Massarinaceae
Semifissispora tooloomensis
CBS143431
MG38607
MG386124
-
-
[21]
Massarinaceae
Stagonospora perfecta
KT 1726A
AB809642
AB807579
AB797289
AB808555
[19]
Massarinaceae
Stagonospora cf. paludosa
CBS 130,005
KF251254
KF251757
-
-
[62]
Massarinaceae
Stagonospora duoseptata
CBS 135,093
KF251255
KF251758
-
-
[62]
Massarinaceae
Stagonospora imperaticola
MFLUCC 15-0026
KY706143
KY706133
KY706138
KY706146
[74]
Massarinaceae
Stagonospora multiseptata
MFLUCC 15-0449
KX965735
KX954404
-
-
[74]
Massarinaceae
Stagonospora paludosa
CBS 135088 *
KF251257
KF251760
-
KF253207
[62]
Massarinaceae
Stagonospora perfecta
CBS 135,099
KF251258
KF251761
-
-
[62]
Massarinaceae
Stagonospora perfecta
KT 1726A
AB809642
AB807579
AB797289
AB808555
[19]
Massarinaceae
Stagonospora pseudocaricis
CBS 135,132
KF251259
KF251763
-
-
[62]
Massarinaceae
Stagonospora pseudopaludosa
CPC 22,654
KF777188
KF777239
-
-
[62]
Massarinaceae
Stagonospora pseudoperfecta
KT 889 *
AB809641
AB807577
AB797287
AB808553
[19]
Massarinaceae
Stagonospora sp.
CBS 135,096
KF251263
KF251766
-
-
[62]
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Table 2. Cont.
Family
GenBank Accession No.
Species
Strain No.
ITS
LSU
SSU
tef1-α
References
Massarinaceae
Stagonospora tainanensis
KT 1866
AB809643
AB807580
AB797290
AB808556
[19]
Massarinaceae
Stagonospora trichophoricola
CBS 136,764
KJ869110
KJ869168
-
-
[75]
Massarinaceae
Stagonospora uniseptata
CPC 22,150
KF251266
KF251769
-
-
[62]
Massarinaceae
Stagonospora uniseptata
CBS 135,090
KF251264
KF251767
-
-
[62]
Massarinaceae
Suttonomyces clematidis
MFLUCC 14-0240
-
KP842917
KP842920
-
[76]
Massarinaceae
Suttonomyces rosae
MFLUCC 15-0051
MG828973
MG829085
MG829185
-
[77]
Periconiaceae
Periconia byssoides
H 4600
LC014581
AB807570
AB797280
AB808546
[19]
Periconiaceae
Periconia digitata
CBS 510.77
LC014584
AB807561
AB797271
AB808537
[19]
Periconiaceae
Periconia macrospinosa
CBS 135,663
KP183999
KP184038
KP184080
-
[78]
Periconiaceae
Periconia pseudodigitata
KT 1395 *
LC014591
AB807564
AB797274
AB808540
[19]
* = The asterisks after the strain number represent the ex-type strains from the holotype specimens.
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3. Results and Discussion
3.1. Phylogenetic Analyses
The individual datasets for ITS and LSU regions comprised selected isolates from
closely related families (Figure 1). The RAxML analyses of the ITS dataset yielded
the best-scoring trees with a final ML optimization likelihood value of -9830.778478
(Figure 1A). The matrix had 531 distinct alignment patterns with 51.80% undetermined
characters or gaps. Estimated base frequencies were as follows: A = 0.227770, C = 0.273565,
G = 0.243931, T = 0.254733; substitution rates AC = 2.172295, AG = 3.427213, AT = 2.029849,
CG = 0.957843, CT = 5.859679, GT = 1.000000; and gamma distribution shape parameter
α = 0.350193. In Figure 1A, the novel taxon Haplohelminthosporium calami grouped within
Massarinaceae and was well separated from other genera but without good bootstrap
support. Helminthosporiella stilbacea (MFLUCC 15-0813) is closely related to Hel.stilbacea
(strains CPHmZC-01 and COAD 2126) with 100% ML/1.00 BYPP.
The RAxML analyses of the LSU dataset yielded the best-scoring trees with a final
ML optimization likelihood value of −4283.882978 (Figure 1B). The matrix had 307 distinct alignment patterns with 12.16% undetermined characters or gaps. Estimated base
frequencies were as follows: A = 0.246483, C = 0.214075, G = 0.309890, T = 0.229553; substitution rates AC = 1.828869, AG = 4.019496, AT = 3.119987, CG = 0.662100, CT = 12.098644,
GT = 1.000000; and gamma distribution shape parameter α = 0.159335. In Figure 1B, the
novel taxon Haplohelminthosporium calami was also well separated within Massarinaceae
and clustered with Helminthosporium and Helminthosporiella. Helminthosporiella stilbacea
(MFLUCC 15-0813) is closely related to Hel. stilbacea (strain CPHmZC-01) with 100%
ML/1.00 BYPP.
The RAxML analysis of the combined (ITS, LSU, SSU, and tef1-α) dataset yielded a
best scoring tree with a final ML optimization likelihood value of -22122.846454 (Figure 2).
The matrix had 1363 distinct alignment patterns, with 41.38% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.241467, C = 0.241603,
G = 0.271551, T = 0.245380; substitution rates AC = 1.860804, AG = 3.064520, AT = 1.916442,
CG = 1.009390, CT = 7.530432, GT = 1.000000; and gamma distribution shape parameter α
= 0.183588. In the phylogenetic analyses (Figure 2), twelve genera are included in the tree.
The novel taxon of Haplohelminthosporium calami grouped within Massarinaceae without
strong bootstrap support. Haplohelminthosporium calami is closely related to H. endiandrae
(CBS 138902, MH878637), but this is statistically unsupported. Helminthosporiella stilbacea
(MFLUCC 15-0813) constitutes a sister phylogenetic affiliation to Hel. stilbacea (strains
CPHmZC-01 and COAD 2126) with 100% ML/1.00 BYPP statistical support.
The phylogenetic analyses (Figures 1 and 2) showed several topologies of the tree
had generally rather low support (ML ≤50% and BYPP ≤0.90). This reflects the relatively
high amount of homoplasy in the data. Most Helminthosporium-like taxa did not have
SSU and tef1-α sequence data for the phylogenetic analyses. In the future, divergent time
estimations will be needed for Helminthosporium-like taxa to resolve taxonomic confusion
and placement.
3.2. Taxonomy
3.2.1. Haplohelminthosporium Konta & K.D. Hyde, gen. nov
Index Fungorum number: IF557873; Facesoffungi number: FoF09169
Etymology—Haplo in Greek means single, which refers to the single conidium in each
conidiophore. It is a close relative of Helminthosporium.
Saprobic on living leaves and petioles of Calamus sp. On living leaves, small spots,
circular to irregular, yellow in the beginning, later becoming red brown surrounded by
yellow. Colonies on natural substrate forming black patches on the upper leaf, petiole
surfaces. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on
natural substrate forming black patches on the upper leaf, petiole surfaces. Mycelium
mostly immersed, partly on the surface forming small stroma-like aggregations of red
brown pseudoparenchymatous cells. Conidiophores arising singly or fasciculate from stroma
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cells, erect, simple, unbranched, straight, curved and swollen at apex, septate, thick-walled,
cylindrical, smooth, bulbous at base, hyaline in the middle, brown to yellow-brown at
1–2-cells above the base, pale brown to yellow-brown at apical cell. Conidiogenous cells
monotretic, terminal, determinate, cylindrical, wide and yellow-brown with a well-defined,
small, noncicatrized pore at the apex. Conidia one for each conidiophore, obpyriform to
lageniform, straight or curved, smooth, olive-brown, distoseptate, with a dark scar at
the base.
Type species—Haplohelminthosporium calami Konta & K.D. Hyde
Notes: Haplohelminthosporium is established as a monotypic genus with Hap. calami as
the type species. ITS phylogenetic analyses separated this genus from other genera, while in
the LSU and multigene analyses it clustered with Helminthosporium and Helminthosporiella,
but both without good statistical support (Figures 1 and 2). Haplohelminthosporium is
presented herein as an asexual morph (hyphomycete) similar to Helminthosporium and
Helminthosporiella in that it is hyphomycete with an erect conidiophore, monotretic conidiogenous cell and distoseptate conidia [19,22,63]. The type species of Helminthosporium has
pale to dark brown, septate conidiophores, with terminal and intercalary polytretic conidiogenous cells, noncicatrized pores at the apex and upper 3–4 cells, solitary or short catenate
conidia that are subhyaline to brown, distoseptate, and is dark brown to black scar at the
base [19]. Helminthosporiella has brown to red-brown conidiophores with terminal, polytretic conidiogenous cells, with catenate and easily disarticulating chains of conidia that are
medium brown, striated at surface and distoseptate [63]. However, Haplohelminthosporium
is distinguished by its unbranched conidiophores arising solitarily or fasciculate from
the stroma-like bulbous basal cells that are hyaline in the middle, brown to red-brown
at 1–2-cells above the base, pale brown to red-brown and curved at the apical cell with
well-defined non-cicatrized small pores and with a single olive-brown conidium arising
from each conidiophore (Figure 3). In the BLAST search of GenBank, the closest match of
the LSU, ITS, and SSU sequence data were identical to Helminthosporium spp. Based on
distinguishing morphological characteristics together with single/multigene phylogenetic
analyses we introduce the newly described strain as a new genus Haplohelminthosporium
in Massarinaceae.
Haplohelminthosporium calami Konta & K.D. Hyde, sp. nov.
Index Fungorum number: IF557874, Facesoffungi number: FoF09170, Figure 3
Etymology: Referring to the genus of palm trees Calamus L.
Holotype: MFLU 20-0520.
Saprobic on living leaves and petioles of Calamus sp. On living leaves, small spots,
circular to irregular, yellow in the beginning, later becoming red-brown surrounded by
yellow. Colonies on natural substrate forming black patches on the upper leaf, petiole
surfaces. Sexual morph: Undetermined. Asexual morph: Mycelium mostly immersed, on
the surface forming small stroma-like aggregations of red brown pseudoparenchymatous
stromal cells (7–)10–14(–20) µm (x = 12 µm). Conidiophores (110–)140–175(–215) × (4–)5–7(–8)
µm (x = 160 × 6 µm, n = 50), wide at the base and apex, macronematous, mononematous,
arising singly or fasciculate from the stroma cells, erect, simple, unbranched, straight,
curved and swollen at the apex, thick-walled, cylindrical, smooth, bulbous at base, hyaline
in the middle, brown to red-brown at 1–2-cells above the base, pale brown to red brown at the
last cell of the apex, (3–)4–5(–6) septa. Conidiogenous cells monotretic, terminal, determinate,
cylindrical, with well-defined small noncicatrized pores at the apex, wide and yellow-brown
at the apex. Conidia (55–)70–100(–120) × (13–)17–20(–23) µm (x = 80 × 20 µm, n = 60), one
on each conidiophore, obpyriform to lageniform, straight or curved, smooth, olive-brown,
(3–)4–6(–7)-distoseptate, with a dark scar at the base.
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Figure 3.
holotype)
(A)(A)
The
forest
in Krabi
Province.
(B–E)
Fresh
and herFigure
3. Haplohelminthosporium
Haplohelminthosporiumcalami
calami(MFLU
(MFLU20-0520,
20-0520,
holotype)
The
forest
in Krabi
Province.
(B–E)
Fresh
and
barium palm
samples.
(F,G)
Colonies
on living
leaf.leaf.
(H–L)
Conidiophores.
(M–U)
Conidia.
(V,W)
Germinated
conidia.
(X)
herbarium
palm
samples.
(F,G)
Colonies
on living
(H–L)
Conidiophores.
(M–U)
Conidia.
(V,W)
Germinated
conidia.
Culture
on on
PDA.
(Y)(Y)
Conidiophore
and
conidia
ononculture.
(X)
Culture
PDA.
Conidiophore
and
conidia
culture.(Z)
(Z)Conidiogenesis.
Conidiogenesis.(AA)
(AA)Conidiophores.
Conidiophores. (AB,AC)
(AB,AC) Conidia.
Conidia.
Scale bars: C, E =2 cm, H–W, Y–AC = 50 μm.
Scale bars: C, E =2 cm, H–W, Y–AC = 50 µm.
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Culture characteristics: Culture on PDA, colony yellow-gray-brown at the center, turning dull creamy white toward to margin, smooth, dense, zonate at the margin (Figure 3X).
Material examined: THAILAND, Krabi Province, on living leaves and petioles of
Calamus sp. (Arecaceae), 14 December 2015, Sirinapa Konta, KHNPR-2 (MFLU 20-0520,
holotype); ex-type living culture, MFLUCC 18-0074.
Notes: BLAST search of the ITS sequence of the newly described strain (Haplohelminthosporium calami) shows 88.89% similarity with Helminthosporium juglandinum (L118),
the LSU sequence shows 98.75% similarity with H. aquaticum (MFLUCC 15-0357), and the
SSU sequence shows 99.52% similarity with H. quercinum (L90). Based on ITS phylogenetic analysis, Haplohelminthosporium calami formed a single branch at the basal clades of
Helminthosporiella and Helminthosporium (Figure 1A), while based on LSU analysis, Hap.
calami clustered together with H. juglandinum (L97), H. endiandrae (CBS 138902, MH878637),
and Hel. stilbacea with no strong statistical support for both analyses. The phylogenetic
results of the combined dataset indicated that Hap. calami clustered with H. endiandrae
(CBS 138902, MH878637) without strong bootstrap support (Figure 2). Comparison of base
pair differences between LSU loci for isolates of Hap. calami strains MFLUCC 18-0074 and
H. endiandrae strains CBS 138,902 (KP004478; Ex-type from the holotype, and MH878637;
sister strain) including gaps showed 1.74% (15/861 bp) differences, and the position of
each base pair difference is shown in Table 3. Other H. endiandrae strains (AKMR1, CBS
138902; ex-type from the holotype, and SM61) grouped together in Helminthosporium, as the
other strains have an ITS region, but the H. endiandrae (CBS 138902, MH878637) strain that
grouped with our new collection lacks the ITS region. Therefore, we compared the morphology of these two species and found that Hap. calami differs from H. endiandrae with respect
to its smaller conidiophores ((110–)140–175(–215) × (4–)5–7(–8) vs. 200–300 × 5–7 µm),
number of conidiophore septa ((3–)4–5(–6) vs. 8–16 septa), larger conidia ((55–)70–100(–120)
× (13–)17–20(–23) vs. (35–)37–45(–57) × (7–)8(–9) µm), solitary conidium per conidiophore,
and higher number of distoseptate ((3–)4–6(–7)-distoseptate vs. 3(–4)-distoseptate). The
results show the placement of Haplohelminthosporium calami within Massarinaceae, and that
this species is distinct from other known species. Therefore, we introduce Hap. Calami as a
new species based on both morphological and phylogenetic data.
Table 3. Polymorphic nucleotides from sequence data of the LSU loci for isolates of Haplohelminthosporium calami MFLUCC
18-0074 and Helminthosporium endiandrae CBS 138,902 (KP004478, MH878637).
Species
LSU
Strain
6
34
74
270
400
412
419
427
480
484
490
491
524
644
843
Haplohelminthosporium calami
(this study)
MFLUCC
18-0074
-
A
A
T
T
T
C
C
A
C
A
T
T
T
G
Helminthosporium endiandrae
(Ex-type from the holotype)
CBS 138,902
(KP004478)
-
C
C
C
C
C
T
T
C
T
T
G
C
G
G
H. endiandrae (sister strain in
Figures 1B and 2)
CBS 138,902
(MH878637)
C
A
C
C
C
C
T
T
C
T
T
G
C
G
-
3.2.2. Helminthosporiella Konta & K.D. Hyde, gen. nov.
Index Fungorum number: IF558311, Facesoffungi number: FoF09171
Helminthosporiella Hern.-Restr., Sarria & Crous, in Crous et al., Persoonia 36: 437 (2016),
MycoBank MB816988, Nom. inval., Art. 40.3 (Shenzhen)
Saprobic on dead petiole of Cocos nucifera.Sexual morph: Undetermined. Asexualmorph:Colony on natural substrate black, hairy. Mycelium mostly immersed, at the surface
forming small stroma-like aggregations of dark brown pseudoparenchymatous cells. Conidiophores macronematous, wide at the apex and base, arising singly from the stroma cells,
erect, simple, unbranched, straight or flexuous, thick-walled, cylindrical, smooth-walled,
dark brown, becoming pale brown at the apex, septate. Conidiogenous cells terminal and
intercalary, polytretic, with well-defined thick, pale brown pores. Conidia obpyriform to
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lageniform, straight or curved, smooth-walled, subhyaline to light brown, distoseptate,
with a thick scar at the base.
Type species—Helminthosporiella stilbacea Konta & K.D. Hyde
Notes: Helminthosporiella was introduced by Crous et al. [63] to accommodate a new
combination of Hel. stilbacea Hern.-Restr., Sarria & Crous, in Massarinaceae, the basionym
of the type species was not provided a Latin diagnosis [63]. In this paper we accept
Helminthosporiella as a distinct genus, presently with a single species Helminthosporiella
stilbacea. Since a Latin diagnosis is no longer required, we provide an English diagnosis
and priority was given to the previous genus and species names. Furthermore, this study
provides the holotype to validate the genus and species, and reports the first host record
of Hel. stilbacea associated with coconut tree (Arecaceae) in Thailand. In particular, based
on the present morphology and DNA sequence data, Helminthosporiella is identified as a
monotypic genus, with Hel. stilbacea as the type species. The members of Helminthosporiella
were found associated with leaf spots on oil palm (Arecaceae) [64].
Helminthosporiella stilbacea Konta & K.D. Hyde, sp. nov.
Index Fungorum number: IF558312, Facesoffungi number: FoF09172, Figure 4.
=Cercospora palmicola f. stilbacea Moreau, Rev. Mycol. 12: 38. 1947 Nom. inval., Art.
39.1 (Shenzhen)
≡Helminthosporiella stilbacea Hern.-Restr., Sarria & Crous, in Crous et al., Persoonia 36:
437. 2016; Nom. inval., Art. 39.1 (Shenzhen)
Helminthosporium stilbaceum Moreau ex S. Hughes, Mycol. Pap.48: 38. 1952; Nom.
inval., Art. 39.1 (Shenzhen).
≡Exosporium stilbaceum Moreau ex M.B. Ellis, Mycol. Pap.82: 38. 1961; Nom. inval.,
Art. 39.1 (Shenzhen).
=Exosporium stilbaceum var. macrosporum Subramon. & V.G. Rao, Journal of the
Annamalai University, part B, Sciences 29: 404. 1971; Nom. inval., Art. 35.1 (Shenzhen).
Saprobic on dead petiole of Cocos nucifera.Sexual morph: Undetermined. Asexualmorph: Colony on natural substrate black, hairy. Mycelium mostly immersed, at the surface forming small stroma-like aggregations of dark brown pseudoparenchymatous cells
(6–)11–15(–25) µm diam (x = 14 µm). Conidiophores (60–)165–270(–310) × (5–)7–9(–12) µm
(x = 200 × 8 µm, n = 30), macronematous, wide at the apex and base, arising singly from
the stroma cells, erect, simple, unbranched, straight or flexuous, thick-walled, cylindrical,
smooth-walled, dark brown, becoming pale brown at the apex, (4–)12–15-septate. Conidiogenous cells terminal and intercalary, polytretic, with well-defined thick, pale brown
pores. Conidia (30–)45–60(–70) × 6–9 µm (x = 50 × 7 µm, n = 30), obpyriform to lageniform,
straight or curved, smooth-walled, subhyaline to light brown, 5–8-distoseptate, with a
thick scar at the base.
Culture characteristics: Culture on MEA, colony yellow-green at the center, turning
dull green, pale yellow next, becoming dull green again, pale yellow, and white toward the
margin. Colony smooth, dense at the middle, zonate, fluffy at the margin (Figure 4P).
Material examined: THAILAND, Prachuap Khiri Khan Province, on dead petiole
of Cocos nucifera L. (Arecaceae), 30 July 2015, Sirinapa Konta PJK04gHB (MFLU 20-0521,
holotype); ex-type living culture, MFLUCC 15-0813.
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Figure 4.
4.Helminthosporiella
Figure
Helminthosporiellastilbacea
stilbacea (MFLU
(MFLU 20-0521,
20-0521, holotype)
holotype) (A)
(A) A
A coconut
coconut plantation
plantation in
in Prachuap
Prachuap Khiri
Khiri Khan
Khan Province.
Province.
(B) Palm
Palm samples.
samples. (C–E)
(C–E) Conidiogenesis.
Conidiogenesis. (F–H)
(F–H) Conidiophores
(at red
red arrow
arrow are
are pores).
pores). (I–M)
(I–M) Conidia.
Conidia. (N,O)
(B)
Conidiophores (at
(N,O) Germinated
Germinated
conidia. (P)
(P) Culture
Culture on
on MEA.
MEA. Scale
Scale bars:
bars: B
B=
= 22 cm,
conidia.
cm, C,
C, I–O
I–O == 20
20 μm,
µm, D–H
D–H == 50
50 μm.
µm.
Life 2021, 11, 454
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Notes: Crous et al. [63] introduced a new genus Helminthosporiella with a new combination of Hel. stilbacea based on fresh collections from oil palm (Elaeis oleifera) in Colombia
and the second collection of Hel. stilbacea was also collected from oil palm (Elaeis guineensis)
in Brazil by Rosado et al. [64]. The full descriptions, illustrations, and sequence data are
provided with interesting information as this species causes elliptical necrotic spots with
a yellowish halo on living leaves of commercial oil palm plantations [63,64]. However,
the type species was invalid because of the basionym lacked a Latin diagnosis [63]. From
these, our fresh collection was collected from dead petiole of coconut (Cocos nucifera) and in
phylogenetic analysis (Figures 1 and 2), three strains of Hel. stilbacea, including our strain,
are grouped together with high bootstrap support. In this study, we therefore provide
a holotype from our specimen, and introduce a new species Helminthosporiella stilbacea,
complete with an English diagnosis, and validated by using the same name while linking
to the valuable information provided from the previous publication of this species.
A BLAST search of the ITS sequence of our isolate showed 90.19% similarity with H.
velutinum (L131), the LSU sequence showed 97.05% similarity with H. aquaticum (MFLUCC
15-0357), the SSU sequence showed 99.15% similarity with H. quercinum (L90), and the
tef1-α sequence showed 92.61% similarity with H. tiliae (L88). These blast results do not
match the results of the phylogenetic analyses.
The comparison between three strains of Hel.stilbacea (see Table 4) from three collections showed that our collection MFLU 20-0521 has several differences when compared
with the other two strains CPHmZC-01 and COAD 2126. Our collection was obtained
from a dead petiole, while the two other strains were isolated from living leaves [63,64].
Therefore, our new collection has been provided as a holotype for Hel. Stilbacea. It is also
the first geographical record from Thailand, and is a new record of the species from a
coconut host (Cocos nucifera).
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Table 4. Comparison of three strains of Helminthosporiella stilbacea.
No.
Herbarium/
Culture No.
Host
(Genus/Family)
Morphology
Locality
Conidiophores
(µm)
Conidiogenous Cells
(µm)
Conidia
(µm)
Mono- or polytretic,
integrated, determinate,
terminal, cylindrical,
31–67 × 4.5–7
Catenate, obclavate,
subcylindrical,
occasionally bifurcate,
medium brown,
26–83 × 7–10,
(1–)3–5(–6)-distoseptate
[63]
[64]
1.
Herbarium: Culture no.:
CPHmZC-01
On leaves of Elaeis
oleifera/Arecaceae
Colombia
Hyaline to pale
brown, smooth,
branched, septate
Erect, brown to red-brown,
synnematous, septate,
compacted, 620–1400 ×
19–54, individual hyphae
3–4 wide
2.
Herbarium: Culture no.:
COAD 2126
On old leaves of
Elaeis guineensis/Arecaceae
Brazil
Hyaline to pale
brown, 2–4
Erect, brown, septate,
synnematous, 66–201(−770)
× 2.5–6(−18)
Mono or polytretic,
cylindrical, terminal,
18–59 × 4–7
Catenate, subcylindrical,
obclavate, brown, 32–83
× 4–11, 2–7-distoseptate
Mostly immersed,
dark brown
Solitarily, erect, unbranched,
straight or flexuous,
cylindrical, bulbous at base,
dark brown, becoming pale
brown at the apex,
(60–)165–270(–310),
(5–)7–9(–12) at the base, 5–8
µm wide at the apex,
(4–)12–15 septate
Terminal and intercalary
with well-defined pores,
pale brown
Obpyriform to
lageniform, straight or
curved, light brown,
(30–)45–60(–70) × 6–9,
5–8-distoseptate
3.
Herbarium: MFLU
20-0521
Culture no.:
MFLUCC 15-0813
On dead petiole of
Cocos
nucifera/Arecaceae
Thailand
References
Mycelia
(µmWide)
This study
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4. Conclusions
In this study, we introduce the new genus Haplohelminthosporium,with Hap. calami as
the type species. In multigene phylogenetic analyses, Hap.calami clustered together with
Helminthosporium endiandrae (CBS 138902) without strong good bootstrap support (other H. endiandrae (AKRM1, CBS 138902 (ex-type), SM61) groups together in Helminthosporium). Moreover, we were unable to synonymize H. endiandrae (CBS 138902) under Haplohelminthosporium
because H. endiandrae has only LSU sequence data available [60]. In the future, H. endiandrae
needs more collections and sequence data to confirm taxonomic placement.
Another newly described isolate clusters together with Helminthosporiella stilbacea.
Helminthosporiella was introduced by Crous et al. [63] but was invalidated as the type species
was not provided with a Latin diagnosis. In this study, we validate Helminthosporiella with
Hel. stilbacea as the type species. Moreover, the newly described strain from this study
is the first saprobic report of Hel. stilbacea, as this was reported in previous studies as a
pathogenic fungus on leaves [63,64]. Moreover, topological nodes in phylogenic analyses
showed conflicting results (Figures 1 and 2). Probably, using only single gene ITS or LSU
analyses will preclude the establishment of taxonomic placements, while combined gene
analyses (including protein coding genes) provide sufficient molecular data to determine
the placements.
Helminthosporium is generally described as a common saprobe found on leaf or twig
litter, and it appears to have a diverse distribution. Occasionally, members of this genus
are also described as pathogens, occurring on a wide range of hosts. Comparison of morphology is important for fungal identification [79]. In this study, we provide a checklist for
Helminthosporium species reported worldwide including details of each species based on
records from Species Fungorum [80] (Table 5). We noted that ten Helminthosporium species
have been found on palm substrates (Arecaceae). Although Helminthosporium conidia
superficially resemble many genera, such as Drechslera, Bipolaris, and Exserohilum, phylogenetic analyses have provided different results [19,33,81–83]. Furthermore, we recommend
revision of the genus Helminthosporium with fresh collections and DNA sequence data
(specifically the ITS region and protein coding genes).
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Table 5. Morphology, host information, locality, sequence data, and related references of Helminthosporium reported worldwide based on the record of Species Fungorum 2021 (bold text
present Helminthosporium reported from Arecaceae).
No.
1
Taxa
H. abietis
Host(Genus/Family)
Abies sp./Pinaceae
Locality
Morphology
Sequence Data
References
U.S.A./Washington
Conidiophores irregularly branched; Conidia 126–150 × 12–16 µm,
fusiform, pointed at both ends, olive-green, 12–15-distoseptate
Absent
[84]
Sierra Leone
Conidiophores 140–280 × 7–11 µm, dense, fasiculate, simple,
straight or flexuous, sometimes swollen at at the tip, septate,
smooth, thick-walled, brown, with well-difinded small pores at
the apex; Conidia 31–(44–)49 × 10–(12–)14 µm in widest part,
narrowing towards the apex to 3–5 µm, obclavate, straight or
flexuous, smooth-walled subhyaline to pale brown,
3–6-distoseptate, with a small dark blackish-brown to black scar
at the base
Absent
[85]
Absent
[86]
H. acaciae
On dead branches of Acacia
farnesiana/Fabaceae
3
H. acalyphae
On leaves of Acalypha
angustifolia/Euphorbiaceae
Dominican Republic
Conidiophores 2.5–4 µm thick, erect, simple, superficial,
brown-blackish, septate; Conidia 9–16 × 4–6 µm, one for each
conidiophore, ovate-ellipsoid, olivaceous-brown or dull-brown,
2–3-distoseptate
4
H. accedens
On living leaves of Dolichos
baumii/Fabaceae
Namibia
Conidiophores 250–300 × 5–9 µm, erect, olive-brown; Conidia
35–57 × 6.5–9 µm, solitary, oblong-fusoid, olive,
3–6-distoseptate
Absent
[87]
Absent
[85]
2
5
H. ahmadii
On dead branches of Quercus
sp./Fagaceae
Pakistan
Conidiophores 220–650 × 12–15 µm, dense, fasiculate, simple,
straight or flexuous, smooth, thick-walled, brown to dark
brown, with small pores at the apex, septate; Conidia
95–(110–)150 × 25–30(–38) µm wide inthe broadest part,
tapering towards the apex to 5–9 µm, obclavate, sometimes
rostrate, straight or flexuous, smooth-walled, brown or dark
brown, 5–15-distoseptate, with a dark blackish-brown to black
scar at base
6
H. aichrysonis
On leaves of Aichryson
dichotomum/Crassulaceae
Spain
No information available
Absent
[88]
7
H. alatum
On dying leaves of Dioscorea
alata/Dioscoreaceae
Dominican Republic
No information available
Absent
[89]
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Table 5. Cont.
No.
8
Taxa
H. albiziae
Host(Genus/Family)
On leaves of Albizia
lebbeck/Fabaceae
Locality
Morphology
Sequence Data
References
Sri Lanka
Conidiophores 70 ×7 µm; Conidia 42–56 × 12 µm, tapering to 4
µm diam. clavate, ends rounded, at the lower end, rough with
minute warts, fuliginous, terminal cell paler, strgight or curved
below, 3–4-distoseptate
Absent
[90]
India
Conidiophores 28–44 × 4.5–6 µm, straight or slightly curved,
one-septate at the base; Conidia 23.5–34 × 8–9 µm, pyriform,
prolongate at the apex, rounded at the base, pale,
cinnamon-brown, 3-distoseptate
Absent
[91]
Absent
[92]
H. albiziicola
Albizzia lebbek/Fabaceae
10
H. allamandae
On living leaves of Allamanda
cathartica/Apocynaceae
Dominican Republic
Conidiophores 100–180 × 8–10 µm, solitary or aggregate, curved,
simple, dark-brown; Conidia 66–110 × 17–20 µm, clavate,
elongate-ellipsoid or subfusoid, erect or curved, gray-brown,
7–10-distoseptate
11
H. alphitoniae
On living leaves of Alphitonia
sp./Rhamnaceae
Malaysia/Mount
Kinabalu
Conidiophores 250–500 ×5–8 µm, erect, dark-brown; Conidia
25–66 × 8–13 µm, obclavate, erect or curved, yellow-brown or
pale olive, 1–6-distoseptate
Absent
[93]
12
H. aneurolepidii
On leaves of Aneurolepidium
ramosum/Poaceae
Russia/West Siberia
No information available
Absent
[94]
13
H. anomalum
From soil
U.S.A./Iowa, Utah
No information available
Present
[17,63]
14
H. anonymicum
In culture: former Soviet Union
Russia
No information available
Absent
[95]
15
H. apiculatum
On dry tree of Betula sp.
(Betulinum)/Betulaceae
Czech Republic
Conidiophores fasiculate, flexuous, simple, hyaline; Conidia long,
37 µm, elliptical-fusiform, with color, multi-septate
Absent
[96]
Absent
[18]
Absent
[96]
9
16
H. appatternae
From leaves of Cynodon
dactylon/Poaceae; from culture
India/Maharashtra
Conidiophores unbranched, of two types; determinate
conidiophores uniform, 182 × 5.2 µm, single, olivaceous, 1–3
septate; indeterminante conidiophores narrower, 208–520 × 7.8
µm, paler and distantly septate at base, gradually broadened
into a darker, close septate; Conidia 20.8–152.0 × 7.8 µm,
6–18-distoseptate
17
H. appendiculatum
On branches of the trees
Czechia
Conidiophores simple, fasciculate; Conidia 65 × 11 µm, clavate,
curved, blunted, whitish, multi-septate
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
China/Yunnan
Conidiophores 410–580 × 13–17 µm, solitary or in groups of 2–4,
erect, flexuous, unbranched, smooth, dark brown paler towards
the apex, bulbous at base, 14–23 septate; Conidia 70–80 × 16–18
µm, single, obclavate, straight or curved, pale brown to brown,
truncate and cicatrized at base, wider than apex, guttulate,
8–10-distoseptate
Present
[20]
Absent
[95,97]
H. aquaticum
On submerged decaying wood
19
H. arcautei
On living leaves Scorpiurus
subvillosa/Fabaceae
Spain
Conidiophores 35–50 × 7–8 µm, erect, simple, cylindrical,
brownish-purple, 2–3 septate; Conidia 48–86 × 10.5–11 µm,
cylindrical-fusoid, straight or slightly curved, light-brown
chestnut, 3–8-distoseptate
20
H. asterinoides
On living leaves of Eugenia
sp./Myrtaceae
Brazil
Conidiophores 5–7 µm thick, fasciculate, rhizoid; Conidia 22–24 ×
5–6 µm, fusoid, curved, colorless at each bottom, 3-distoseptate
Absent
[98]
21
H. asterinum
On Liquidambar sp./
Altingiaceae
U.S.A./Florida
Conidiophores erect, simple, septate; Conidia 500–600 × 80 µm,
clavate, 3–4-distoseptate
Present
[99]
22
H. astragali
On leaves of Astragalus
siversianus/Fabaceae
Kyrgyzstan
No information available
Absent
[100]
India/Maharashtra
Conidiophores 3–7 septate, unbranched, and of two types; shorter
conidiophore uniformly wide, 62.4–72.8 × 7.8 µm, brown;
longer ones narrow at the base and paler, gradually broadening
and darkening towards the apex, 440–680 × 5.2–10 µm; Conidia
yellow to brown, darkening at maturity, of two kinds; normal
ones 23–93.6 × 26 µm, elliptical with hemispherical edges,
widest at the middle, 0–10-distoseptate; a typical conidia
abundant, forked or geniculate, septation forked, brown to dark
brown, 5–8-distoseptate
Absent
[101]
18
23
H. atypicum
On leaves of Triticum
sp./Poaceae
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
24
H. austriacum
On dead corticated twigs of
Fagus sylvatica/Fagaceae
Austria/Döbling,
Kahlenberg, Wien
Conidiophores 275–700(–920) µm long, 11.5–19 µm wide at the
base, tapering to 7–11 µm near the apex, solitarily or fasciculate,
erect, simple, sub-cylindrical, straight or flexuous, thick-walled,
smooth, brown to dark brown, paler near the apex, with
well-defined small pores at the apex, 1–12 septate; Conidia
(30–)35–48(–97) × (10.0–)13.7–16.5(–19.8) µm, tapering to 4.5–6.0
µm at the distal end, obpyriform to lageniform, straight or
curved, smooth, pale brown, (4–)5–7(–10)-distoseptate, with a
blackish-brown 3–6 µm wide scar at the base
25
H. avenae-pratensis
On sheaths of Avena
pratensis/Poaceae
Germany
Conidiophores 300 × 8–11 µm, solitary or fasciculate,
dark-chestnut, septate; Conidia 70–107 × 16–21 µm, cylindrical
or obclavate, light brown, on both sides paler, 5–11-distoseptate
Absent
[102]
26
H. bactridis
On sheaths of Bactris
sp./Arecaceae
Brazil/Pará
Conidiophores 200 × 3–4.5 µm, septate; Conidia 20–30 × 6–8 µm,
fusoid, 6–7-distoseptate
Absent
[103]
27
H. bakeri
On dead stems of Premnavestita
sp./Lamiaceae
Philippines
Conidiophores 500–800 ×12 µm wide at base to below, 10 µm
wide, erect, unbranched, dark; Conidia 80–150 × 17–22 µm,
solitary, oblong, obclavate, 3–6-distoseptate
Absent
[104]
Absent
[105]
Absent
[106]
28
H. bambusicola
On dead culm of Bambusa
sp./Poaceae
China/Sichuan
Conidiophores 55–247 × 4–6 µm, fasciculate or solitary, simple,
cylindrical, straight or flexuous, thick walled, smooth, brown,
paler towards the apex, with well-defined small pores, 1–2
septate; Conidia 36–66 × 6–11 µm narrowing towards the apex
to 2–4.5 µm wide, obclavate, straight or slightly flexuous,
thin-walled 1–1.5 µm thick, smooth, pale brown, paler towards
the apex, 5–8-distoseptate, scar not distinct at the base
29
H. bataticola
On living leaves of Ipomoea
batatas/Convolvulaceae
Caucasus
No information available
Present
[21]
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Sierra Leone
Conidiophores 350–110 × 10–15 µm thick at the apex, 15–20 µm
thick at the base, dense, fasciculate, simple, straight or flexuous,
smooth-walled, dark brown, sometimes paler towards the apex,
with well definded, small pores septate; Conidia 55-(86–)145 ×
16–(17.2–)18 µm thick in broadest part, tapering to 3–4 µm the
apex, obclavate, straight or flexuous, rostrate, smooth-walled,
subhyaline to brown, 7–18-distoseptate, with a dark blackish
brown to black scar ath the base
Absent
[85]
Absent
[107]
H. bauhiniae
On dead twigs of Bauhinia
tomentosa/Fabaceae
31
H. belgaumense
On litter, Calamus
thwaitesii/Arecaceae
India/Karnataka
Conidiophores 140–250 × 6–9 µm, erect, straight to flexuous,
unbranched, smooth, brown; Conidia 10–15 × 6–11 µm, solitary,
dry, sub-spherical, dark brown, truncate at base, roundea at the
apex, 1-distoseptate
32
H. bhawanii
On leaves of Eragrostis
japonica/Poaceae
India/Bihar
No information available
Absent
[108]
33
H. bigenum
Palmae rotten
petiole/Arecaceae
Peru
No information available
Absent
[109]
34
H. bondarzewii
From grains of Triticum sp. and
Secale sp./
Poaceae
Russia, Ukraine
No information available
Present
[60,110]
35
H. cacaliae
Cacalia sonchifolia/Asteraceae
Brazil
No information available
Absent
[111]
36
H. cacaophilum
From unfermented Cacao beans,
Theobroma cacao/Malvaceae
Dominican
Republic/Santo
Domingo
No information available
Absent
[112]
37
H. cactacearum
In young plants of Cereus
species/Cactaceae
Italy
No information available
Absent
[113]
H. caespitiferum
Meliola spec. in leaf spots of
living leafs of Omphalea
pauciflora/
Euphorbiaceae
Dominican
Republic/Santo
Domingo
Conidiophores 150–300 × 6.5–8 µm, simple, dark-brown, septate;
Conidia 18–42 × 8–11 µm, oblong to fusoid, dark-brown,
constrict at septum, (3–)6–7-distoseptate
Absent
[92]
30
38
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
39
H. canephorae
Coffea canephora/Rubiaceae
Democratic Republic of
the Congo/Zaire
No information available
Absent
[114]
40
H. cantareirense
On dead stems
Brazil/São Paulo
Conidiophores 7–12 µm thick, erect, fasciculate; Conidia 50–60 ×
8–12 µm, clavate, brown, constrict at septum, 6–8-distoseptate
Absent
[115]
41
H. cantonense
On decaying culms of Bambusa
vulgaris/Poaceae
China
Conidiophores 80–95 × 6 µm; Conidia 50–62 × 8 µm, obclavate,
7–9-distoseptate
Absent
[116]
42
H. caperoniae
On living leaves of Caperonia
palustris/Euphorbiaceae
Dominican Republic
Conidiophores 100–300 × 3.5–5 µm, 2–5 fasciculate, simple,
olive-brown; Conidia 22–55 × 4–6 µm, oblong-fusoid or
subclavate, rarely cylindrical, yellow or gray-brown
Absent
[92]
Dominican
Republic/Santo
Domingo
Conidiophores 1–4 articulate, 200–350 µm long, very densely
fasciculate, erect to sub-erect, straight or slightly irregularly
curved, almost straight ot curved, dark-brown to blackish, tip
light-colored; Conidia 22–25 ×8–10 µm, 1–4 to each conidiopore,
easily falling, ellipsoid to ovoid, with narrowed ends, or basal
end narrowed-truncate, apical end rounded to acute, not
caudate, central cells from dark-brown to brownish, and cells
light brown to yellowish, 2–5-distoseptate
Absent
[117]
43
H. carpocrinum
Parasite on perithecia of Meliola
funebris on leaves of Omphalea
sp./Euphorbiaceae (O.
pauciflora)
44
H. carposaprum
On Lycopersicon
esculentum/Solanaceae
British Guiana, Haiti,
Mexico
No information available
Absent
[118]
45
H. ceibae
On leaves of Ceiba
pentandra/Malvaceae
Philippines
No information available
Absent
[119]
Present
[85,120]
Absent
[121]
46
H. chlorophorae
On dead twigs of Chlorophora
regia/Moraceae
Sierra Leone
Conidiophores 120–270 × 7–10 µm thick at the base, often swollen
towards the tip up to 12 µm, single or fasciculate, simple,
straight or flexuous, smooth-walled, brown to dark brown, with
1–3 well-definded, small pores, septate; Conidia 52–(73–)102 ×
8–(9.5–)11 µm, thick in the widest part narrowing gradually
towards the apex to 3–5 µm, obclavate, straight or flexuous,
smooth-walled, subhyaline to pale brown, 6–9-distoseptate,
with a tather large dark blackish-brown to black scar at the base
47
H. chrysobalani
On dry leaves of Chrysobalanus
icaco/Chrysobalanaceae
Dominican
Republic/Bonao
Conidiophores up to 6 µm, fasciculate, erect, 2–3 septate; Conidia
25–50 × 3–4 µm, fusoid, 2–4-distoseptate
Life 2021, 11, 454
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Ecuador/Tungurahua
Conidiophores 200–350 × 4–6 µm, dense, erect, fasciculate,
simple, straight or slightly curved, dark-brown or olive, septate;
Conidia 32–50 × 9–11 µm, elongate-fusiform, blunt at both ends,
curved, rarly straight, gray or olive-brown, 3–4-distoseptate
Absent
[122]
48
H. chusqueae
On living and dying leaves of
Chusquea serrulata/Poaceae
49
H. cibotii
On leaves of Cibotium
sp./Cibotiaceae
U.S.A./Hawaii Islands
No information available
Absent
[123]
50
H. ciliare
-
-
No information available
Absent
[124]
51
H. citri
On leaves of Citrus poonensis,
Citrus tankart, Citrus ponki, and
of Citrus sinensis var.
brasiliensis/Rutaceae
China/Taiwan
No information available
Absent
[125]
52
H. claviphorum
Rotten branch
Peru
No information available
Absent
[109]
53
H. cleosmatis
On living leaves of Clematis sp./
Ranunculaceae (in foliisvivis
Cleosmati soctandri)
Dominican Republic
Conidiophores 140–250(–300) µm long, 4–5 µm wide, solitary,
erect, simple, dark-brown, often becoming paler; Conidia 28–52
× 6.5–9 µm, clavate or fusoid, yellow or pale olive-brownish,
(3–)4–5-distoseptate
Absent
[92]
54
H. clusiae
On leaves of Clusiarosa
sp./Clusiaceae
Dominican Republic
Conidiophores 108–128 × 12–16.5 µm effuse, brown-black,
irregular at based, or subbulbose, septate; Conidia 26–32 ×
10–11.5 µm, fusoid, subfusoid or cylindrical, 4–8-distoseptate
Absent
[126]
55
H. coffeae
On leaves of Coffea
liberica/Rubiaceae
Ghana
Conidiophores 300–400 × 7–8 µm, effuse, nigro-olivaceas,
aggregate, erect, cylindrical, rect or flexuous, olives-brown,
septate; Conidia 45–55 × 8–10 µm, obovate, 3–5-distoseptate
Absent
[127]
China/Guangxi
Conidiophores 60–280 × 7.0–8.5 µm, fasciculate, simple,
subcylindrical, straight or flexuous, thick-walled, smooth, dark
brown, paler towards the apex, with 1–3 well-defined small
pores at the apex, 1–2 septate; Conidia 100–147.5 µm long,
9.5–11 µm diam in the widest part, narrowing towards the apex
to 3–4 µm diam, straight or slightly flexuous, smooth-walled,
pale brown, sometimes verruculose at apex, 11–17-distoseptate,
with a large dark blackish-brown scar at the base, 2–3 µm thick
Absent
[128]
56
H. conidiophorellum
On dead branches of tree
Life 2021, 11, 454
28 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
57
H. constrictum
On dead branches of
Trachycarpus fortunei/Arecaceae
China/Guangdong
Conidiophores single, simple, subcylindrical, straight or slightly
flexuous, brown to dark brown, paler towards the apex, 1–3
septate; Conidia 57–120 × 9–12 µm, thick in the widest part,
narrowing toward the apex to 2.5–5 µm, abruptly tapered to a
truncate base, tretic, obclavate, straight or slightly flexuous, pale
brown, paler toward to apex, 9–15-distoseptate, sometimes
constricted at one or two septa
58
H. conviva
On Hyphoderma caliciferum, the
genus of crust fungi in the
family Meruliaceae.
Spain/Archipelago/
Balearic/Baleares
Islands
No information available
Absent
[130]
59
H. corchori
On leaves of Corchorus
capsularis/Malvaceae
China/Taiwan
No information available
Absent
[131]
60
H. crassiseptum
Meliola abrupta
Dominican Republic
Conidiophores 30–50 × 2–3 µm, septate; Conidia 45–55(–65) ×
12–14 µm, ovoid or elliptical, (2–)3-distoseptate
Absent
[86]
61
H. crotalariae
On leavesof Crotalaria
juncea/Fabaceae
India/Assam
No information available
Absent
[132]
62
H. crus-galli
On living leaves of Echinochloa
crus-galli (=Panicum
crista-galli)/Poaceae
Japan
No information available
Absent
[133,134]
63
H. cubense
On rachis of Roystonea
regia/Arecaceae
Cuba
No information available
Absent
[135]
64
H. cucumerinum
On living leaves of Cucumis
sativus/Zingiberaceae
Russia/Krym
No information available
Absent
[136]
65
H. curvulum
On decaying leaves of Zea
mays/Poaceae
Philippines
Conidiophores 160–180 × 7–7.5 µm, fasciculate, filiform, septate;
Conidia 25–35 × 8–9 µm, oblong-fusoid, narrow,
3(–4)-distoseptate
Absent
[137]
66
H. cuspidatum
On decaying branches of Afzelia
rhomboidea/Fabaceae
Philippines
Conidiophores 800–900 × 8–9 µm, fasciculate, filiform,
multiseptate; Conidia 100–130 × 11–12 µm, obclavate,
8–12-distoseptate
Absent
[137]
Absent
[129]
Life 2021, 11, 454
29 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Conidiophores 100–130 × 4–5 µm, subfasciculate, filiform long,
simple, fuliginous up paler, septate; Conidia 14–15 × 2.5 µm,
cylindrical, apex rounded, base acuted, minute, pale fuliginous,
3-distoseptate
Absent
[138]
67
H. cylindricum
On rotten wood
Czech
Republic/Bohemia
68
H. cymmartinii
On leaves of Cymbopogon
martinii/Poaceae
India/Uttar Pradesh
No information available
Absent
[108]
69
H. cyperi
On Cyperus sp./
Cyperaceae
Greece
Conidiophores straight to subflexuous, greenish, paler at apex;
Conidia 78 × 9 µm, fusoid, fuscidull, 5–8-distoseptate
Absent
[139]
70
H. dactylidis
On leaves of Dactylis
glomerata/Poaceae
U.S.A./Pennsylvania
No information available
Absent
[140]
Present
[85]
71
H. dalbergiae
On dead branches of Dalbergia
sissoo/Fabaceae
Pakistan
Conidiophores 300–1300 × 10–12(–15) µm, dense, fasciculate,
simple, flexuous, smooth-walled, brown to dark brown,
sometimes paler towards the apex, with well-definded small
pores, septate; Conidia 58–(93–)125 × 12–(13.2–)14 µm thick in
broadest part, tapering to gradually towards the apex to 3–5 µm,
obclavate, straight or flexuous, smooth-walled, straw-coloured
to pale brownwith, 5–17-distoseptate, large dark
blackish-brown to black scar at the base
72
H. davillae
On leaves of Davilla
rugosa/Dilleniaceae
U.S.A./San Francisco
Conidiophores 4–6 µm, thick filiform, flexuous, unbranched,
elongate, brown, septate; Conidia 40–70 × 4–6 µm,
elongate-obclavate, narrower and paler, (1–)2–4-distoseptate
Absent
[141]
H. decacuminatum
In the dry twigs on Vitis
vinifera/Vitaceae
Italy
Conidiophores 4 µm thick, extremely short-articulated, irregular,
dark reddish-brown; Conidia 40–45 × 10 µm, long clavate,
decacumina to tip, or cut down in pedicellum narrowed, pale
brown-gray, 4–5-distoseptate
Present
[60,142]
74
H. delicatulum
On stems of Umbelliferae or
Apiaceae
UK/Great Britain
Conidiophores slender, subulate, multi-articultate, brown, paler
at the tips; Conidia oblong, nearly colourless, with the apices
very obtuse, consisting of about five swollen articulations, one
or two of which have occasionally a vertical dissepiment
Absent
[143]
75
H. delphinii
On stems of Delphinium
brunonianum/Ranunculaceae
Russia
No information available
Absent
[144]
73
Life 2021, 11, 454
30 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
76
H. dendroideum
On Acer sp./Sapindaceae
U.S.A./South Carolina
Conidiophores 1–2 short branchlets termintated, oblong,
subfusiform, slightly curved, multiarticulate conidia; Conidia
60 µm long, each joint containing a globose nucleus
Absent
[145]
77
H. densum
-
-
No information available
Absent
[146]
78
H. desmodii
On Desmodium buergeri/
Fabaceae
Japan
No information available
Absent
[147]
79
H. diedickei
No information available
No information available
No information available
Absent
[148]
Absent
[149]
80
H. dimorphosporum
On decaying rotting stems of
unknown liana
Cuba
Conidiophores 150–400 µm long, at the apex 9–12 µm, at the base
10–14 µm wide, single or fasciculate 2–10, simple, straight or
flexuous, smooth, dark brown, paler towards the apex, septate;
Conidia of two different types arising through pores a t the apex
(1–4 pores) and late rally beneath the upper septa: (a) 19–24 ×
8–10.5 µm, broadly ellipsoidal, ovoid or broadly fusiform,
thick-walled, smooth, brown to dark brown, 1-distoseptate; (b)
24–65 µm long, 10–15 µm wide in the broadest part, tapering to
3.2–4.8 µm at the apex, obclavate, rostrate, straight or flexuous,
pale brown, smooth, 6–9-distoseptate, with a dark brown scar at
the base
81
H. dolichi
On living leaves of Dolichos
sp./Fabaceae
Namibia
Conidiophores 250–350 × 4–6 µm, erect, olive-brown; Conidia
27–38 × 5.5–8 µm, solitary, oblong-subfusoid, olive,
2–3-distoseptate
Absent
[87]
82
H. dongxingense
Rhododendron sp.
China
No information available
Absent
[150]
83
H. elasticae
-
-
No information available
Absent
[151]
On leaves of Endiandra
introrsa/Lauraceae
Australia/New South
Wales, Nightcap
National Park
Conidiophores 200–300 × 5–7 µm, solitary, erect, subcylindrical,
straight to flexuous, unbranched, thick-walled, base bulbous,
lacking rhizoids, brown, 8–16 septate; Conidia (35–)37–45(–57) ×
(7–)8(–9) µm, solitary or in short chains (2–3), obclavate,
thick-walled, finely roughened, brown, 3(–4)-distoseptate
Present
[21,59]
84
H. endiandrae
Life 2021, 11, 454
31 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
85
H. eragrostiellae
On inflorescence and leaves of
Eragrostis bifida/Poaceae
India/Uttar Pradesh
No information available
Absent
[108]
86
H. erythrinae
On leaves of Erythrina
suberosa/Leguminosae
India/Karnataka
Conidiophores 32–42 × 4–5 µm, simple, brownish-yellow; Conidia
39–62 µm at base, straight or vermiform, rounded at the apex
and flat at the base, pale cinnamon-brown, 4–8-distoseptate
Absent
[91]
Present
[22]
87
H. erythrinicola
On leaves of Erythrina
humeana/Fabaceae
South Africa/Eastern
Cape
Conidiophores 500–1200 × 6–10 mm, fasciculate, subcylindrical,
unbranched, brown, becoming pale brown at apex, multiseptate;
Conidia (70–)80–90(–110) × (9–)10–11(–12) mm, obclavate,
straight to curved, apex subobtuse, smooth, medium brown,
(6–)7–8(–12)-distoseptate
88
H. exasperatum
On Dianthus barbatus/
Caryophyllaceae
UK/Great Britain
Conidiophores flexuous, knotted above, each knot bearing oblong
conidia; Conidia 30–45 × 10–12 µm
Absent
[152]
89
H. feijoae
On leaves of Acca
sellowiana/Myrtaceae (syn:
Feijoa sellowiana)
North
America/Hispaniola
island
No information available
Absent
[153]
90
H. ferrugineum
On leaves of Hiraea sp. and
Heteropterys sp./Malpighiaceae
U.S.A./San Francisco
Conidiophores 8–9 µm thick, filiform, yellow, septate; Conidia
50–62 × 11–14 µm, obclavate, subhyaline, last 2 septate
hyaline-yellow to yellow
Absent
[141]
91
H. fici
On leaves of Ficus
retusa/Moraceae
Philippines, Thailand
Conidiophores fusciculate, long, nodulosis, septate; Conidia 18–20
× 5–6 µm, cylindrical, reddish-brown, 3-distoseptate
Absent
[137,154]
92
H. ficinum
On leaves of Ficus
ulmifolia/Moraceae
Philippines
Conidiophores 250 × 6 µm, filiform, septate; Conidia 50–60 × 6–8
µm, obclavate, 4–5-distoseptate
Absent
[137]
93
H. filicicola
On leaves of Lygodium
sp./Lygodiaceae and of
Selaginella sp./Selaginellaceae
Peru
Conidiophores 400 × 3–5 µm thick, erect, simple, filiform, septate;
Conidia 30–40 × 6–10 µm, cylindrical-fusoid or clavate, both
side blunt, 3–5-distoseptate
Absent
[155]
94
H. flagellatum
On mycelium of Meliola, in
leaves of Ardisia
disticha/Myrsinaceae
Philippines
Conidiophores 2.5–4 µm thick, erect, sub-hylaline
Absent
[156]
95
H. flumeanum
On leaves of Bambusa
sp./Bambuseae
Philippines
Conidiophores 90–100 × 6–7 µm, dense, fasticulate, filiform;
Conidia 35–40 × 9–12 µm, obclavate, 3-distoseptate
Absent
[157]
Life 2021, 11, 454
32 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
96
H. fumagineum
On leaves of ficusulmifolia/
Moraceae
Philippines
Conidiophores 240–300 × 7 µm, filiform, septate; Conidia 35 ×
9–10 µm, oblong-obclavate, 3-distoseptate
Absent
[137]
97
H. gibberosporum
Musa cavendishii/Musaceae
Somalia
No information available
Absent Present
[158]
98
H. glabroides
On Meliola glabroides, on Piper
aduncum/
Piperaceae
Puerto Rico
Conidiophores 100–140 × 7 µm; Conidia 40–81 × 6–7 µm,
3–6-distoseptate
Absent
[159]
99
H. gleicheniae
On leaves of Dicranopteris
linearis (=Gleichenia dichotoma)/
Gleicheniaceae
U.S.A./Hawaii Islands
No information available
Absent
[123]
100
H. gossypii
On living leaves and bracts of
Gossypium sp./
Malvaceae
North America
Conidiophores 40–185 × 6.5–8.5 µm, singly or in groups of three
to six, straight cylindrical to nodose or bent, brown, 5 septate;
Conidia 35–118 × 11.7–18.4 µm, elliptical, curved, rarely straight,
light to dark fuliginous, thick walled, rounded at the ends,
1–8-distoseptate
Absent
[160]
101
H. grewiae
On leaves of Grewia
sp./Malvaceae
Democratic Republic of
the Congo
Conidiophores 80–120 × 5–8 µm, fasciculate, septate; Conidia
35–45 × 8–10 µm, fusoid, 2–4-distoseptate
Absent
[161]
Absent
[128]
Absent
[162]
102
103
H. guangxiense
On dead branches of
unidentified tree
China/Guangxi,
Shanglin
Conidiophores 330–850 µm long, 15–20 µm wide just above the
base and 8–13 µmwide toward the apex, fasciculate, simple,
straight or flexuous, sub-cylindrical, thick-walled, smooth,
brown, with 1–3 well-defined small pores at the apex, 1–4
septate; Conidia 76–110 µm long, 16–22 µm wide in the widest
part, narrowing towards the apex to 3–6µm wide, straight or
curved, obclavate, smooth, middle brown, paler towards the
apex, 9–17-distoseptate, with a large dark blackish-brown scar
at the base, 1.5–3.5 µm thick
H. guianense
Meliola guianensis parasitic on
mycelium on living leaves of
Theobroma cacao/
Malvaceae
Guyana
No information available
Life 2021, 11, 454
33 of 53
Table 5. Cont.
No.
104
105
106
Taxa
H. heringerianum
Host(Genus/Family)
Tipuana speciosa/
Fabaceae
H. hispanicum
On dead corticated twigs of
Juglans regia/Juglandaceae
H. hispaniolae
On living leaves of Manihot
utilissima/
Euphorbiaceae
Locality
Morphology
Sequence Data
References
Brazil
No information available
Absent
[163]
Asturias, Selviella, Spain
Conidiophores 130–540 µm long, 13–22.5 µm wide at the base,
tapering to 8–15 µm near the apex, solitarily or in small groups,
erect, simple, straight or flexuous, thick-walled, subcylindrical,
smooth, dark to blackish brown, paler near the apex, with
well-defined small pores at the apex, 1–2 septate; Conidia
69–99(–130) × (17–)18–21(–24) µm, obclavate, straight or
flexuous, thin-walled, smooth, pale brown,
(4–)6–11(–14)-distoseptate, with a blackish-brown 4–6 µm wide
scar at the base
Present
[21]
Dominican
Republic/Haiti
Conidiophores sub-hyaline to light-grey, when old, with an
almost hyaline tip; Conidia 14.8–(53.5–)81.4 × 7.4–(11–)14.8 µm,
sub-hyaline to smoky, irregular, cylindric-elongate to ellipsoidal,
straight or slightly curved, with the basal end applanate,
1–8-distoseptate
Absent
[112]
Absent
[67]
107
H. hunanense
On dead branches of
unidentified tree
China/Zhangjiajie,
Hunan
Conidiophores 70–226 × 5–7 above, 8.5–14 µm base, solitary or
fasciculate, simple, cylindrical, straight or flexuous,
thick-walled, smooth, brown, well-defined small pores at the
apex, 1–3 septate; Conidia 56–127 × 10–14 base, apex 2–4 µm,
obclavate, straight or curved, smooth, middle brown, paler
towards the apex, 4–12-distoseptate, blackish-brown scar at the
base, 1.5 µm thick
108
H. hygrophilae
On leaves of Hygrophila
brasiliensis/Acanthaceae
Dominican Republic
No information available
Absent
[89]
109
H. insigne
On leaves of Mallotus
philippensis/
Euphorbiaceae
Philippines
Conidiophores 600–800 × 50 µm, fasciculate, filiform, blackish,
septate; Conidia 45–55 × 7–8 µm, obclavate, often curved,
4–5-distoseptate
Absent
[137]
Life 2021, 11, 454
34 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
110
H. insuetum
On living leaves of Philodendron
sodiroi (=Piplocarpha
sodiroi)/Araceae
Ecuador/Pichincha
Conidiophores 2.5–5 µm thick, olive brown or dark brown;
Conidia 17–38 × 7–12 µm, oblong, ellipsoid or oblong-ellipsoid
fusiform and often subclavate, rarely cylindrical, often straigtly,
rarely curved, olive brown or dark-brown,
(3–)5–7(–9)-distoseptate, scared or a little more often in the
middle constricted
111
H. ipomoeae
On leaves of Ipomoea reptans/
Convolvulaceae
China/Taiwan
No information available
Absent
[130]
H. iranicum
On living leaves of Indigofera
sp./Fabaceae
Iran/Bandar Abbas
Conidiophores 40–75 × 6-9 µm, dense, curved, rarely straight,
dark-brown, septate; Conidia 36(–42) × 7–11 µm, oblong,
narrowly ellipsoid or curved, obtuse at both ends, straight or
curved, sometimes irregular, olive, 1–3-distoseptate
Absent
[164]
Italy
Conidiophores (190–)330–600 × (12–)16–18(−20) µm, aggregated,
erect, straight or slightly flexuous, unbranched, cylindrical, dark
brown, 13–25 septate; Conidia 58–78 × 15–19(−23) µm,
obclavate, straight or curved, pale brown to brown, slightly
truncate and black at base, rounded, narrowed,
6–11-distoseptate
Absent
[61]
Present
[21]
Absent
[165]
112
113
H. italicum
On dead branch of Alnus
glutinosa/Betulaceae
114
H. juglandinum
On dead corticated twigs of
Juglans regia/Juglandaceae
Austria/ Niederösterreich/Gießhübl,
Italy
Conidiophores (175–)215–325(–455) µm long, 11–23 µm wide at
the base, 8.5–14 µm wide near the slightly inflated apex,
fasciculate, erect, simple, straight or flexuous, thick-walled,
sub-cylindrical, smooth, brown to dark brown, darker to black
at the apex, the latter with a well-defined apical pore; Conidia
(69–)89–145(–205) × (15.0–)16.5–20.0(–25.0) µm, rostrate,
straight or flexuous, thin-walled, smooth, pale brown,
(5–)9–17(–20)-distoseptate, blackish-brown scar at the base
115
H. juglandis
Juglans sp./Juglandaceae
China, Yunnan
No information available
Absent
[122]
Life 2021, 11, 454
35 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Kenya
Conidiophores 250–550 × 8–12 µm, solitary, unbranched; Conidia
30–90 × 8–10 µm, in the broadest part, uniformly tapering to
2–4 µm wide at at the apex, solitary, simple straight or
somewhat curved, obclavate, rostrate, subhyaline, smooth,
4–15-distoseptate
Absent
[166]
116
H. kakamegense
On dead attached twig of
Uvariopsis congensis/
Annonaceae
117
H. kalakadense
On dead unidentified twig
India/Tamil Nadu
Conidia 13–15 µm
Absent
[21]
118
H. kalopanacis
On dead wood of Kalopanax
septemlobus/
Araliaceae
Russia/Primorye
No information available
Absent
[167]
119
H. kok-saghyz
In seeds of Taraxacum
kok-saghyz/Asteraceae
Russia
No information available
Absent
[168]
120
H. kyllingae
Kyllinga sp./
Cyperaceae
Uganda
No information available
Absent
[169]
121
H. lablab
On leaves of Dolichos
lablab/Fabaceae
China/Taiwan
No information available
Absent
[130]
On MEA and PDA
Conidiophores 100–300 × 4–6(–7) µm, erect, subcylindrical,
thick-walled, medium brown, multiseptate; Conidia
(35–)70–110(–170) × (6–)7–8(–11) µm, obclavate to
subcylindrical, straight to slightly curved, thick-walled,
medium brown, (3–)4–6(–10)-distoseptate
Present
[21]
122
H. leucadendri
On leaves of Leucadendron sp./
Proteaceae
South Africa/Western
Cape Province,
Helderberg Nature
Reserve
123
H. leucosykes
On Meliola, on leaves of
Leucosyke capitellata/
Urticaceae
Philippines
Conidiophores 300 × 7–8 µm, erect, brown, septate; Conidia 30 ×
8 µm, 3-distoseptate
Absent
[156]
China/Guangxi,
Nanning
Conidiophores 127–700 µm long, 9.5–18 µm diam just above the
base and 8.5–10 µm diam towards the apex, solitary, simple,
straight or flexuous, smooth or verruculose, thickwalled, dark
brown, with 1–3 well-defined small pores at the apex, 1–4
septate; Conidia 24–38.5 × 9.5–13 µm, obclavate, straight or
slightly curved, rostrate or pseudorostrate, smoothwalled, pale
brown, subhyaline towards the apex, 4–6-distoseptate, with a
large dark blackish-brown scar at the base, 1–2 µm thick
Absent
[128]
124
H. ligustri
On dead branches of Ligustrum
quihoui/
Oleaceae
Life 2021, 11, 454
36 of 53
Table 5. Cont.
No.
125
Taxa
H. litseae
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Litsea polyantha/
Lauraceae
India/Assam
No information available
Absent
[170]
Conidiophores 500 × 4–6 µm, erect, flexuous, cylindrical, smooth
to rough-walled, medium brown, multiseptate; Conidia
(25–)40–55(–65) × (7–)8–9 µm, subcylindrical, straight, smooth,
medium brown, apex obutuse, base somewhat obconic,
(3–)4–6(–7)-distoseptate
Present
[171]
126
H. livistonae
On leaves of Livistona
australis/Arecaceae
Australia/New South
Wales, Murramarang
National Park
127
H. longisinuatum
Palmae rotten trunk
Peru
Conidiophores 20–75 × 3.5–5 µm; Conidia 65–220(–1000) × 8–10.5
µm, solitary, long, narrowly obclavate, 9–22-distoseptate
Absent
[109]
128
H. lonicerae
On Lonicera sp./
Caprifoliaceae
Brazil
No information available
Absent
[111]
129
H. lophirae
On leaves of Lophiraalata
sp./Ochnaceae
Sierra Leone
Conidiophores 110–200 × 3–4 µm thick, simple, bluntly rounded
ends; Conidia solitary 15–29 × 3.5–4.5 µm, oblong or
oblong-cylindrical, hook or curved, smooth, olive- brown, 1–2
guttulate, 1–3-distoseptate
Absent
[172]
130
H. lunzinense
No information available
No information available
No information available
Absent
[173]
131
H. lusitanicum
On Alnus glutinosa/
Betulaceae
Portugal
No information available
Absent
[174]
132
H. lycopersici
On Solanum
lycopersicum/Solanaceae
Guinea
No information available
Absent
[175]
133
H. machaerii
On Machaerium sp./ Fabaceae
Brazil
No information available
Absent
[111]
UK/Great Britain
Conidiophores erect, simple, fusiform, 7–10 septate; Conidia
0.5-0.65 × 0.1 mm
Absent
[176]
Japan
Conidiophores 150–270 µm long, 9.5–13 µm thick at the apex,
8.5–13.5 µm thick at the base, single or fasciculate, straight or
flexuous, smooth walled, brown to dark brown, sometimes
paler toward the apex, septate; Conidia 100–203 × 12.5–22.5 µm
tapering gradually to 2.5–5 µm thick near the apex, solitary,
obclavate or rostrate, straight or flexuous, pale olive-brown to
pale brown, paler toward the apex, 7–18-distoseptate, with a
blackish-brown to black scar, 4–7 µm thick
Present
[177,178]
134
135
H. macilentum
H. magnisporum
On rotten wood
On dead fallen branches of an
unknown woody plant
Life 2021, 11, 454
37 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
136
H. makilingense
On dead branches of Paramignya
monophylla/Rutaceae
Philippines
Conidiophores 400–600 × 7–9 µm, dense, erect, curved, brown,
septate; Conidia 100–300 × 10–12 µm, obclavate,
12–18-distoseptate
Absent
[179]
137
H. manihotis
on living leaves of Manihot sp./
Euphorbiaceae
Brazil
Conidiophores 50–95 × 4–6 µm, 4–6 septate; Conidia 40–50 × 6–8
µm, vermiform, clavate to subfusoid, olives, 4–7-distoseptate
Absent
[180]
138
H. marantae
On leaves of Maranta
arundinacea/Marantaceae
China/Taiwan
No information available
Absent
[130]
139
H. massarinum
Berchemia racemose/
Rhamnaceae
Japan
Conidiophores 380–810 × 7–9 wide at the apex, 13.5–21 wide at
the base µm, 15–25 septate; Conidia 17–56.5 × 5–9 µm, tretic,
solitary or in short chains (5–6), obclavate, rostrate, pale brown,
smooth, with or without guttules, 1–8-distoseptate
Present
[19]
140
H. mattiroloi
On branches of Sideroxylon
oxyacantha/Sapotaceae
Etiopia
No information available
Absent
[181]
141
H. mayaguezense
On culms and leaves of
Paspalum conjugatum/
Poaceae
Puerto Rico
Conidiophores 300–500 × 18–22 µm; Conidia 135–155 × 35–45 µm,
fusoid to clavate, 3–4-distoseptate
Absent
[182]
142
H. melastomacearum
On Meliolamelastomacearum, on
Miconiaracemose/
Melastomataceae
Puerto Rico
Conidiophores 280 × 3 µm; Conidia 14–21 × 3.5–6 µm, ellipsoid,
3-distoseptate
Absent
[159]
143
H. meliae
On leaves of Melia
azedarach/Meliaceae
Dominican Republic
Conidiophores 250–350 × 15–22 µm, simple, aggregated,
branched, olive-brown to black, septate; Conidia 70–100 × 12–15
µm, elongate, fusoid, or clavate
Absent
[183]
144
H. melioloides
On leaves of Uvaria
sp./Annonaceae
Philippines
Conidiophores 250–300 × 6–8 µm; Conidia 35–45 × 9–10 µm,
obclavate, 3-distoseptate
Absent
[137]
England, Italy
Conidiophores 100–550 × 8–14 µm, fasciculate, simple, flexuous,
cylindrical, smooth-walled, dark brown, with a pore at the apex
and often 1–2, septate; Conidia 60–(114–)160 × 12–(17–)22 µm
thick in broadest part, tapering to 4–10 µm near the apex,
obclavate, smooth-walled, pale to mid golden-brown,
9–17-distoseptate, with 5–7 µm wide at the scar
Present
[184]
145
H. microsorum
On twigs of Quercus ilex/
Fagaceae
Life 2021, 11, 454
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
146
H. microsporum
From soil
India/Maharashtra
Conidiophores 234–468 × 10.8 µm, pale brown, 10–16 septate;
Conidia 26–41 × 22 µm, fusoid, widest at the middle, brown,
2–7-distoseptate
Absent
[18]
147
H. minimum
On dead decorticatd branches
UK/Great
Britain/England
Conidiophores erect, simple, septate; Conidia 12–14 × 3–4 µm,
fusiform, obtuse at the ends, triseptate, scarcely
constricted, hyaline
Absent
[185]
Absent
[129]
148
H. multiseptatum
On dead branches
China/Guangdong
Conidiophores 390–650 × 10–14 µm wide at the base, 7–9 µm at
apex, simple, subcylindrical, straight or slightly flexuous,
smooth-walled, brown to dark brown, paler towards the apex,
with 1–3 well definded, small pores, 1–3 septate; Conidia
78–190 µm long, 11–16 µm thick in the widest part, narrowing
toward the apex to 3–6 µm, tretic, straight or slightly flexuous,
obclavate or whip-like, smooth-walled, pale brown paler
toward the apex, 13–25-distoseptate, with a dark
blackish-brown scar at the base
149
H. nadsonii
On fibers of
Gossypium sp./Malvaceae
Russia
No information available
Absent
[186]
Present
[187]
150
H. nanjingense
On dead branches of an
unidentified tree
China/Jiangsu, Nanjing
Conidiophores 250–470 × 6.9–7.7 µm, solitary or fasciculate,
simple, straight or flexuous, thick-walled, sub-cylindrical,
smooth, brown to dark brown, with well-defined small pores at
the apex,1–4 septate; Conidia 64.5–170.5 µm long, 7.3–10.3 µm
wide in the widest part, narrowing towards the apex to
5.0–6.8 µm wide, subulate or nearly whip-like, straight or
curved, thin-walled, smooth, pale brown, 6–17-distoseptate,
with a blackish-brown scar at the base, 1.4–2.7 µm thick
151
H. naviculare
On leaves of Euphorbia sp./
Euphorbiaceae
Brazil/Tubarão
Conidiophores 6–8 µm thick, branched, often curved, yellow,
septate; Conidia 50–84 × 11–16 µmnaviculiform, hyaline at
length, very pale with brown
Absent
[188]
152
H. naviculatum
On dead herbaceous stems of
Solidago sp./Asteraceae
U.S.A./New York
No information available
Absent
[189]
153
H. newbouldiae
On leaves of Newbouldialaevis/
Bignoniaceae
Guinea
No information available
Absent
[190]
Life 2021, 11, 454
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Table 5. Cont.
No.
154
Taxa
H. novae-zelandiae
Host(Genus/Family)
On dead wood and bark of Vitex
lucens/
Lamiaceae
Locality
Morphology
Sequence Data
References
New Zealand
Conidiophores 165 µm long, 4.8–7(–9) µm, erect, single or in
groups, simple, or once-branched at the base, straight or
flexuous, subcylindrical, brown to dark brown below, very pale
brown to subhyaline above, 15 septate; Conidia 13.5–16.2 ×
7.2–9.0 µm, solitary, obovoid, sometimes slightly, smooth, the 2
lower cells being brown and the distal cell paler with a dark
band of wall overlying each septum, 2-distoseptate
Absent
[191]
Absent
[128]
155
H. obpyriforme
On dead branches of
unidentified tree
China/Guangxi
Conidiophores 225–460 µm long, 9.5–13 µm diam just above the
base and 6–8.5 µm diam towards the apex, arising singly from
the upper cells of the stromata, simple, subcylindrical, straight
or flexuous, dark brown, paler towards the apex, with
well-defined small pores at the apex,1–3 septate; Conidia
47–74 µm long, 14–19 µm diam in the widest part, narrowing in
diameter towards the apex to 2.5–5 µm, straight or slightly
curved, obpyriform, smooth-walled, middle brown, paler
towards the apex, 5–9-distoseptate, with a large dark
blackish-brown scar at conidium base, 1–2 µm thick
156
H. ocoteae
On Meliola ocoteae, on
Guareatrichilioides
Puerto Rico
Conidiphores 135–200 × 4 µm, septate; Conidia 20–28 × 4–6 µm,
3-distoseptate
Absent
[159]
157
H. oligosporum
Holotype of Sporidesmium
olivaceum: on rotten branches of
Tilia sp.
Lectotype of
Coryneumoligosporum, here
designated: on rotten branches
of Corylus sp.
Epitype of
Sporidesmiumolivaceum and of
Coryneumoligosporum: on dead
corticated twigs of Tiliacordata
sp.
Austria, Czech Republic,
Germany
From Epitype specimen [21]
Conidiophores (17–)22–35(–46) × (8.0–)8.5–10.5(–11.5) µm,
densely crowded, erect, simple, straight, cylindrical to slightly
swollen at the apex, brown to dark brown, darker at the apex,
smooth, 0–2 septate; Conidia (37–)59–80(–124) ×
(14.8–)15.8–18.0(–20.0) µm, tapering to 4–10.5 µm at the distal
end, with 4–8 µm wide, dark brown to black scar at the base,
obclavate, sometimes rostrate, straight or curved, smooth but
occasionally wrinkled with age, pale brown to brown, paler
toward the apex, 6–12(–16)-distoseptate
Present
[21,124]
158
H. olisipponense
Culture from the perithecia
stage of Pyrenophora polytricha
-
No information available
Absent
[192]
Life 2021, 11, 454
40 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
159
H. oplismeni
On leaves of Oplismenus
cotnpositus/Poaceae
China/Taiwan
No information available
Absent
[130]
160
H. orchidacearum
On leaves of Neottia ovata
(=Listera ovata)/Orchidaceae
France
No information available
Absent
[193]
161
H. orthospermum
On rotten wood
U.S.A./New York
Conidiophores 50–60 × 5 µm, erect, simple, fasciculate, straight,
dark, 3–4 septate; Conidia 60–80(–110) × 10–12 µm, cylindrical,
straight, apex rounded, tuncated at base, 12–14-distoseptate
Absent
[194]
162
H.
oryzae-microsporae
On Oryza sativa/Poaceae
Japan
No information available
Absent
[195]
Absent
[128]
163
H. ovoideum
On dead branches of tree
China/Jilin
Conidiophores 380–510 × 15–25 µm diam just above the base,
7.5–10 µm diam towards the apex, arising singly from the upper
cells of the stromata, simple, subcylindrical, straight or flexuous,
thick-walled, smooth, brown to dark brown, paler towards the
apex, with 1–3 well-defined small pores at the apex, 1–6 septate;
Conidia 27–61 × 13–21 µm diam in the widest part, narrowing
towards the apex to 4.5–8.5 µm, straight, ovoid, to ellipsoidal,
smooth-walled, moderately brown, paler towards the apex,
3–8-distoseptate, with a large dark blackish-brown scar at the
base, 1.5–2.5 µm thick
164
H. pachystelae
On living leaves of Synsepalumm
solo(=Pachystelam solo)/
Sapotaceae
Tanzania
Conidiophores 300–350 × 6–8 µm, erect, simple, septate; Conidia
35–50 × 10–13 µm, fusoid or oblong clavate or lanceolate,
3-5-distoseptate
Absent
[196]
H. palaestinum
On stems and flowers of
Dianthus sp./
Caryophyllaceae
Israel
Conidiophores 30–160 × 6–8 µm, fasciculate, 8–16 aggregate,
simple, bent, thick-walled, coffin terminal obtuse, thin, yellow
or colorless, 5–7 septate; Conidia 60–120 × 9–12 µm, solitary,
obclavate, rectiusculis or curved, pale-olive, minute-granule,
thick-walled, towards colorless above, 5–7-distoseptate
Absent
[197]
165
Life 2021, 11, 454
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Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
Brasil/Pará, Papua New
Guinea
From reference specimen [190]; Conidiophores 132.5–195 × 5–6
µm, solitary, erect, simple, cylindrical, straight or flexuous,
smooth, brown, light brown at the apex, 7–10 septate; Conidia
38–53 × 8–11 base, 3–4 µm apex, solitary, in small chains,
obclavate or cylindrical, straight or slightly curved, simple,
smooth, brown with light brown at apical cell, 6–10-distoseptate
Absent
[198,199]
166
H. palmigenum
On rotten fruit of Cocos
nucifera/Arecaceae; On petriole
and rachis from reference
specimen
167
H. panici
On leaves of Panicum
maximum/Poaceae
Indonesia/Java
Conidiophores 115–180 × 8–10 µm; Conidia (35–)50–75 ×
(7–)10–13 µm, ellipsoidal-truncate, ellipsoidal-elongate,
dull-brown, (1–)3(–4)-distoseptate
Absent
[159]
168
H. papulosum
On bark of Malus sylvestris or
Pyrus communis/Rosaceae
West Virginia
No information available
Absent
[200]
169
H. parathesicola
On Meliola parathesicola, on
Parathesis serrulate/Primulaceae
Puerto Rico
Conidiophores 120 × 4 µm, solitary; Conidia 17–20 × 4–6 µm, base
truncate, apex beaked, beak often 7 µm long, 1–3-distoseptate
Absent
[159]
170
H. paulense
On leaves of Myrtaceae
Brazil/São Paulo
Conidiophores 3–4.5 µm thick, brown, septate; Conidia 15–24 × 4
µm, fusoid, brown, 3-distoseptate
Absent
[115]
171
H. penniseti
On leaves of Pennisetum glaucum
(=Pennisetum typhoides)/Poaceae
India/Uttar Pradesh
No information available
Absent
[108]
172
H. philippinum
On decaying leaves of Arenga
mindorensis/
Arecaceae
Philippines
Conidiophores 300–400 × 6–7 µm, fasciculate, filiform, curved,
septate; Conidia 33–35 × 8–9 µm, obclavate, 4-distoseptate
Absent
[137]
173
H. philodendri
On Meliola philodendri, on
Philodendron krebsii/Araceae
Puerto Rico
Conidiphores 400 × 3–4 µm; Conidia 24–35 × 5–9 µm, clavate,
3-distoseptate
Absent
[159]
174
H. phomatae
On bark of Acer pennsylvanicum/
Sapindaceae
U.S.A./New York
No information available
Absent
[189]
175
H. phyllantheum
On dead branches hanging
down of Phyllanthus
sp./Phyllanthaceae
Philippines
Conidiophores 180–200 × 4.7–6 µm, fillifrom, blackedned, septate;
Conidia 80–90 × 9–10 µm, obclavate, long, 9–11-distoseptate
Absent
[137]
Life 2021, 11, 454
42 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
176
H. piperis
On leaves of Piper
betle/Piperaceae
China/Taiwan
No information available
Absent
[130]
177
H. portoricense
On dead branches hanging
down of Phyllanthus
sp./Phyllanthaceae
Philippines
Conidiophores 25–250 × 2–5 µm; Conidia 30–60 × 6–10 µm,
elongate-fusoid, olive-brown or brown, (2–)4-distoseptate
Absent
[86,201]
178
H. proliferatum
On grain of Triticum
sp./Poaceae
India/Maharashtra
Colony on PDA; Conidiophores 292–510 × 7–13.8 µm,
unbranched, pale, olivaceous, 5–20 septate; Conidia 23–126 ×
11.5–13.8 µm; cylindrical, olivaceous, 3–13-distoseptate
Absent
[101]
Absent
[128]
179
H.
pseudomicrosorum
On dead branches of
unidentified tree
China/Changbaishan,
Jilin
Conidiophores 155–288 × 11–15 µm, fasciculate, simple,
cylindrical, straight or flexuous, smooth, dark brown, paler
towards the apex, with 1–3 well-defined small, 1–4 septate;
Conidia 82–142 × 17–27 µm in the widest part, narrowing
towards the apex to 3–6 µm diam, tretic, straight or slightly
flexuous, obclavate, smooth-walled, brown, paler towards the
apex, 7–16-distoseptate, with a large dark blackish-brown scar
at the base, 2–4 µm thick
180
H. pseudotsugae
On bark and resin exudations of
Pseudotsuga taxifolia var.
glauca/Pinaceae
U.S.A.
Conidiophores scattered on aerial hyphae with usually one at
each cell; Conidia 65–105 × 14–15 µm, opaque, black or greenish
black, smooth walls, with 8–14-distoseptate
Absent
[202]
181
H. purpurascens
On leaves of Panicum
purpurascens/Poaceae
U.S.A./Florida
No information available
Absent
[203]
182
H. pyracanthae
Pyracantha sp./Rosaceae
Portugal
No information available
Absent
[204]
U.S.A.
Conidiophores (115–)133–226(–300) µm long, 14–20 µm wide at
the base, tapering to 10–15 µm near the apex, solitarily or
fasciculate, simple, straight or flexuous, cylindrical,
thick-walled, smooth, brown to dark brown, with well-defined
small pores at the apex; Conidia 60–100 × 15–22 µm, straight or
flexuous, obclavate, smooth-walled, brown, 8–10-distoseptate,
with blackish-brown to black scar at the base
Absent
[21]
183
H. quercicola
On dead corticated branches of
Quercus cf. reticulata/Fagaceae
Life 2021, 11, 454
43 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
Locality
Morphology
Sequence Data
References
184
H. quercinum
On dead corticated twigs of
Quercus petraea/Fagaceae
Conidiophores (40–)74–199(–332) µm long, 11–18 µm wide at the
base, tapering to 8.5–13.5 µm near the apex, solitarily or
fasciculate, simple, straight or flexuous, cylindrical, smooth,
Austria/Niederösterreich, brown to dark brown, with well-defined small pores at the apex,
Spitzerberg
1–5 septa; Conidia (47–)78–130(–201) × (13.2–)15.3–18.0(–20.5)
µm, straight or flexuous, rostrate, smooth-walled, brown,
8–13(–20)-distoseptate, with blackish-brown to black scar at
the base
185
H. repens
On bark of dead Acer
grandidentatum/
Sapindaceae
U.S.A./Utah, Red Butte
Canyon
Conidia 40–45(–60) × 8–9 µm, sub-oblong, 5–12-distosepate
Absent
[205]
186
H. reyesii
On dead branch of Guioa sp./
Sapindaceae
Philippines
Conidiophores 130 × 8–10 µm, erect, brown, septate; Conidia
34–112 × 8–13 µm, tereti-fusoid, brown, ends hyaline,
5–14-distosepate
Absent
[137]
187
H. rhodomyrti
On leaves of Rhodomyrtus
tomentosa/Myrtaceae
China/Guangdong
Conidia 42–60 × 17–20 µm, fusoid, brown, 5–7-distosepate
Absent
[206]
188
H. rhopaloides
On decraying stem of Brassica
oleracea/Brassicaceae
Britain, France, Germany,
Italy, Portugal
Conidiophores short, dark brown-black, 12–14 septate; Conidia
0.04–0.1 mm long, 0.08 mm wide, straight or slightly curved
Absent
[207,208]
189
H. schelkownikowii
On branches
Armenia, Azerbaijan,
Georgia, Russia
No information available
Absent
[209]
190
H. scolecoides
On dry woood
Germany/Reichenberg
Conidiophores simple, branched; Conidia 80 × 7.5 µm, torulus,
fusciculate, septate, yellow
Absent
[96]
191
H. sechiicola
On Sechium edule/
Cucurbitaceae
Puerto Rico
No information available
Absent Present
[210]
192
H. sichuanense
On dead branches of tree
China/Sichuan
No information available
Absent Present
[211]
Present
[21]
Life 2021, 11, 454
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Table 5. Cont.
No.
193
Taxa
H. solani
Host(Genus/Family)
On stem of Solanum
nigrum/Solanaceae (type);
Citrus linella; Leucaena glauac;
Solanum dulcamara; S. nigrum;
Solanum tuberosum
Locality
Morphology
Sequence Data
References
England, Guernsey, New
Zealand, New Guinea,
Sierra Leone, Wales
Conidiophores 120–600 × 9–15 µm thick near base, 6–9 µm thick
near the apex, erect, simple, straight or flexuous, smooth or
occasionally, brown to very dark brown, paler near apex,
septate, with small pores at apex, 1–8 septate; Conidia (24–)39–85
× (7–)9–11 µm, straight or curved, obclavated, smooth-walled,
subhyaline to brown, 2–8-distosepate, with a welll-defind dark
brown to black scar at base
Present
[212]
Absent
[213]
194
H. solitarium
On leaves of Iris sp./Iridaceae
U.S.A./Minnesota
Conidiophores 60–150 × 6 µm, solitary, slightly fasciculate, erect,
swollen at the base, lighter colored at the apex, dark brown,
septate; Conidia 24–30 × 8–9 µm, oblong-elliptical, sometimes
slightly curved, dark brown, at first 2–4 guttulate,
3–5-distosepate
195
H. spirotrichum
On withered leaves of
Cyrtophyllum
fragrans/Gentianaceae
Singapore
Conidiophores 190–220 × 6 µm, fasciculate, filiform, brown,
septate; Conidia 23–25 × 9 µm, oblong-obclavate, gently curving,
brown, 3-distosepate
Absent
[214]
196
H. spurirostrum
On dead branches of tree
China/Sichuan
No information available
Absent
[211]
197
H. subapiculatum
On dead wood of Sambucus
callicarpa/Adoxaceae
U.S.A./Washington
Conidiophores 8–10 µm thick; Conidia 35–80 × 12–16 µm, oblong
or subfusiform, 6–7-distosepate
Absent
[215]
China/Guangdong
Conidiophores 120–200 × 10–12 (basal), above 6–8.5 µm thick,
simple or fasciculate, erect, subcylindrical, brown, pores 1–3 µm,
septate; Conidia 72–125 × 9–11.5 µm, obclavate, straight or
flexuous, subhyaline, apex 2.5–5 µm, black at tip,
6–9-distosepate, dark blackish-brown scar
Absent
[129]
China/Yunnan
Conidiophores 239–423 × 8.5–15.5 µm, solitary or in group of 2–4,
unbranched, straight or curved, smooth, dark brown, paler
towards to the apex, bulbous at base 9–14 septate; Conidia 41–55
× 14.5–18.5 µm, straight or curved, wider below than apex,
truncate and dark at base, apically rostrate and pale, smooth,
pale brown to mid-brown, guttulate, 6–10-distosepate
Present
[24]
198
199
H. subhyalinum
H. submersum
On living leaves of Phoenix
hanceana/Arecaceae
On submerged decaying wood
Life 2021, 11, 454
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Table 5. Cont.
No.
200
Taxa
H. subsimile
Host(Genus/Family)
On withered and dead leaves of
Bruguiera sexangula (=Bruguiera
eriopetala)/Rhizophoraceae
Locality
Morphology
Sequence Data
References
Singapore
Conidiophores 200–250 × 8–9 µm, filiform-fasciculate, brown,
septate; Conidia (38–)45–50 × 11–12(–14) µm, brown,
3-distoseptate
Absent
[216]
Present
[22]
201
H. syzygii
On bark canker of Syzygium
sp./Myrtaceae
South Africa/Eastern
Cape Province
Conidiophores 150–400 × 10–15 mm, fasciculate, unbranched,
clavate at apex, dark brown, multiseptate; Conidia
(70–)80–100(–150) × (19–)22–23(–25) mm, obclavate, curved,
apex subobtuse, warty, inner surface striate, medium brown,
(7–)9–12-distoseptate
202
H. theobromae
On leaves of Theobroma
cacao/Malvaceae
Italy
Conidiophores 500–1000 µm, erect, 6–10 septate; Conidia 60–160 ×
12–20 µm, obclavate to tereti-obclavate
Absent
[217]
203
H. theobromicola
On rotten leaves of Theobroma
cacao/Malvaceae
Dominican
Republic/Moca
Conidiophores 20–33 × 3.5–5 µm olives-brown, septate; Conidia
46–58 × 10–13.5 µm, elliptic-oblong or subfusoid, irregular,
3–5-distoseptate
Absent
[218]
204
H. tritici
On seedhead of Triticum
vulgare/Poaceae
Tanzania
Conidiophores 3.5–5 µm, thick fasciculate, erect, sepate; Conidia
12–25 × 4–7 µm, subcylindrical-oblong, clavate or fusoid,
2–4-distoseptate, constrict at septum
Absent
[219]
205
H. tritikernelis
On kernels of Triticum
aestivum/Poaceae
India/Bihar
No information available
Absent
[108]
206
H. turbinatum
On unidentified wood
Great Britain
Conidiophores simple, slender; Conidia elongated, turbinatis,
tuncatus, apiculate, brown, 4–7-distoseptate
Absent
[220]
207
H. ubangiense
On leaves of Coffea
sp./Rubiaceae
Democratic Republic of
the Congo/Ubangi River
Conidiophores (2–)3–6 µm, fasciculate, erect, branched, septate;
Conidia 30–60 × 5–8 µm, fusoid, 3–4-distoseptate
Absent
[221]
208
H. ustilaginoideum
On flowers of Panicum
spicatum/Poaceae
Democratic Republic of
the Congo
Conidiophores 3–3.5 µm thick, fasciculate; Conidia 10–50 × 3.5–4.5
µm, cylindrical or subfusoid, blunted, 1–5-distoseptate
Absent
[121]
Brazil/Pernambuco
Conidiophores 150–200 × 10–14 µm, erect, unbranched, straight
or flexuous, cylindrical, slightly inflated at the apex, smooth,
brown, 5–7 septate; Conidia 29–58 × 4–7 µm,
cylindrical-obclavate, subcylindrical, oblong or navicular, dry,
trick-walled, with wall verrucose or verruculose, gray-brown,
lumina pale yellow, (0–)1–4-distoseptate
Absent
[222]
209
H. varium
On decaying leaves of
unidentified plants
Life 2021, 11, 454
46 of 53
Table 5. Cont.
No.
Taxa
Host(Genus/Family)
H. varroniae
On leaves of Varronia
sp./Boraginaceae
211
Locality
Morphology
Sequence Data
References
Puerto Rico
Conidiophores 160–200 × 4 µm; Conidia 27–44 × 6–7 µm,
3-distoseptate
Absent
[223]
H. velutinum
Fagus sylvatica dead corticated
twigs/saprobic on decaying
wood submerged in stream
Austria, Wien, Döbling,
Kahlenberg/China
From reference specimen [20]; Conidiophores 530–655 × 16–18
µm, erect or flexuousk, unbranched, dark brown, 17–23 septate;
Conidia 67–79 × 15–19 µm, single, obclavate, straight or curved,
smooth, pale brown to brown, 7–9-distoseptate, rounded at
apex, guttulate when young, non-guttulate at maturity
Present
[16,20]
212
H. viticis
On leaves of Vitis sp./Vitaceae
Brazil/Pará
Conidiophores 80 × 2–3 µm, fasciculate, septate; Conidia 12–20 ×
2.5–3.5 µm cylindrical, 1–3-distoseptate
Absent
[223]
213
H. wagateae
On leaves of Moullava spicata
(=Wagatea spicata)/Fabaceae
India/Karnataka
Conidiophores 81–125 × 1.5 –2.5 µm, yellowish-brown,
multiseptate; Conidia 15.5–28 × 3–4 µm, clavate-cylindric,
cinnamon-yellow, rounded at both ends, 2–4-distoseptate
Absent
[91]
214
H. warpuriae
On stem of Warpuria
clandestina/Acanthaceae
Great Britain/England
Conidiophores 300–500 × 6–8 µm; Conidia 115–190 × 12–14 µm,
obclavate, 8–11-distoseptate
Absent
[224]
215
H. xanthosomatis
On leaves of Xanthosoma
violaceum/Araceae
Dominican
Republic/Moca
Conidiophores 35–90 µm long, septate; Conidia 185 × 24 µm,
fusoid, subfusoid to subclavate, 3–7(-10)-distoseptate
Absent
[225]
216
H. xylopiifolii
On Asterina, on Xylopia
sericea/Annonaceae
Brazil/Pernambuco
Conidiophores 85–305 × 5.5–8 µm, erect, 3–5 septate; Conidia
38–62 × 8–13.5 µm, cylindrical or clavate, 3–6-distoseptate
Absent
[226]
210
Life 2021, 11, 454
47 of 53
Author Contributions: Conceptualization, S.K., K.D.H., S.C.K., C.S. and S.T.; Data curation, S.K.;
Methodology, S.K., A.M. and S.T.; Resources, K.D.H., S.C.K., J.X. and S.L.; Supervision, K.D.H.
and S.T.; Writing—original draft, S.K., K.D.H., S.C.K., A.M., C.S., L.A.P.D., C.M.N., J.X. and S.T.;
Writing—review & editing, S.K., K.D.H., S.C.K., S.T. and S.L. All authors have read and agreed to the
published version of the manuscript.
Funding: Saowaluck Tibpromma would like to thank the International Postdoctoral Exchange
Fellowship Program (number Y9180822S1), CAS President’s International Fellowship Initiative (PIFI)
(number 2020PC0009), China Postdoctoral Science Foundation and the Yunnan Human Resources,
and Social Security Department Foundation for funding her postdoctoral research. Kevin D. Hyde
thanks the Thailand Research Funds for the grant “Impact of climate change on fungal diversity and
biogeography in the Greater Mekong Subregion (RDG6130001)”. Samantha Karunarathna thanks
CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research
(number 2018PC0006) and the National Science Foundation of China (NSFC) for funding this research
work under project code 31750110478. This work was partly supported by Chiang Mai University.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Acknowledgments: Sirinapa Konta is grateful to Paul Kirk, Shaun Pennycook, Jayarama Bhat, and
Sirilak Radbouchoom, for their valuable suggestions and comments.
Conflicts of Interest: The authors declare no conflict of interest.
References
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
Hyde, K.D.; Norphanphoun, C.; Chen, J.; Dissanayake, A.J.; Doilom, M.; Hongsanan, S.; Jayawardena, R.S.; Jeewon, R.; Perera,
R.H.; Thongbai, B.; et al. Thailand’s amazing diversity: Up to 96% of fungi in northern Thailand may be novel. Fungal Divers.
2018, 93, 215–239. [CrossRef]
Hyde, K.D.; de Silva, N.I.; Jeewon, R.; Bhat, D.J.; Phookamsak, R.; Doilom, M.; Boonmee, S.; Jayawardena, R.S.; Maharachchikumbura, S.S.N.; Senanayake, I.C.; et al. AJOM new records and collections of fungi: 1–100. AJOM 2020, 3, 22–294. [CrossRef]
Dai, D.Q.; Phookamsak, R.; Wijayawardene, N.N.; Li, W.J.; Bhat, D.J.; Xu, J.C.; Taylor, J.E.; Hyde, K.D.; Chukeatirote, E.
Bambusicolous fungi. Fungal Divers. 2017, 82, 1–105. [CrossRef]
Tibpromma, S.; Hyde, K.D.; McKenzie, E.H.; Bhat, D.J.; Phillips, A.J.; Wanasinghe, D.N.; Samarakoon, M.C.; Jayawardena, R.S.;
Dissanayake, A.J.; Tennakoon, D.S.; et al. Fungal diversity notes 840–928: Micro-fungi associated with Pandanaceae. Fungal
Divers. 2018, 93, 1–160. [CrossRef]
Pinruan, U.; Hyde, K.D.; Lumyong, S.; McKenzie, E.H.C.; Jones, E.G. Occurrence of fungi on tissues of the peat swamp palm
Licuala longicalycata. Fungal Divers. 2007, 25, 157–173.
Pinnoi, A.; Phongpaichit, S.; Hyde, K.D.; Jones, E.G. Biodiversity of fungi on Calamus (Palmae) in Thailand. Cryptogamie 2009, 30,
181–190.
Konta, S.; Hongsanan, S.; Tibpromma, S.; Thongbai, B.; Maharachchikumbura, S.S.N.; Bahkali, A.H.; Hyde, K.D.; Boonmee, S. An
advance in the endophyte story: Oxydothidaceae fam. nov. with six new species of Oxydothis. Mycosphere 2016, 7, 1425–1446.
[CrossRef]
Konta, S.; Hongsanan, S.; Eungwanichayapant, P.D.; Liu, J.K.; Jeewon, R.; Hyde, K.D.; Maharachchikumbura, S.S.N.; Boonmee, S.
Leptosporella (Leptosporellaceae fam. nov.) and Linocarpon and Neolinocarpon (Linocarpaceae fam. nov.) are accommodated in
Chaetosphaeriales. Mycosphere 2017, 8, 1943–1974. [CrossRef]
Konta, S.; Hyde, K.D.; Eungwanichayapant, P.D.; Doilom, M.; Tennakoon, D.S.; Senwanna, C.; Boonmee, S. Fissuroma (Aigialaceae:
Pleosporales) appears to be hyperdiverse on Arecaceae: Evidence from two new species from southern Thailand. Acta Bot. Bras.
2020, 34, 384–393. [CrossRef]
Konta, S.; Maharachchikumbura, S.S.N.; Senanayake, I.C.; McKenzie, E.H.C.; Stadler, M.; Boonmee, S.; Phookamsak, R.;
Jayawardena, R.S.; Senwanna, C.; Hyde, K.D.; et al. A new genus Allodiatrype, five new species and a new host record of
diatrypaceous fungi from palms (Arecaceae). Mycosphere 2020, 11, 239–268. [CrossRef]
Zhang, S.N.; Hyde, K.D.; Jones, E.B.G.; Cheewangkoon, R.; Liu, J.K. Additions to Fissuroma and Neoastrosphaeriella (Aigialaceae,
Pleosporales) from palms. Mycosphere 2020, 11, 269–284. [CrossRef]
Hongsanan, S.; Hyde, K.D.; Phookamsak, R.; Wanasinghe, D.N.; McKenzie, E.H.C.; Sarma, V.V.; Lücking, R.; Boonmee, S.; Bhat,
J.D.; Liu, N.G.; et al. Refined families of Dothideomycetes: Orders and families incertae sedis in Dothideomycetes. Fungal Divers.
2020, 105, 17–318. [CrossRef]
Wijayawardene, N.N.; Hyde, K.D.; Al-Ani, L.K.T.; Tedersoo, L.; Haelewaters, D.; Rajeshkumar, K.C.; Zhao, R.L.; Aptroot, A.;
Leontyev, D.V.; Saxena, R.K.; et al. Outline of Fungi and fungus-like taxa. Mycosphere 2020, 11, 1060–1456. [CrossRef]
Munk, A. On Metasphaeria coccodes (Karst.) Sacc. (Massarinaceae n. fam.). Friesia 1956, 5, 303–308.
Life 2021, 11, 454
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
48 of 53
Saccardo, P.A. Sylloge Fungorum; Johnson Reprint Corporation: New York, NY, USA, 1883; Volume 2.
Link, H.F. Observationes in ordines plantarum naturales. Dissertatio I. Mag. der Ges. Nat. Freunde Berl. 1809, 3, 3–42.
Gilman, J.C.; Abbott, E.V. A summary of the soil fungi. Iowa State Coll. J. Sci. 1927, 1, 225–343.
Deshpande, K.S.; Deshpande, K.B. Contribution to the taxonomy of the genus Helminthosporium II. Sydowia 1969, 23, 69–76.
Tanaka, K.; Hirayama, K.; Yonezawa, H.; Sato, G.; Toriyabe, A.; Kudo, H.; Hashimoto, A.; Matsumura, M.; Harada, Y.; Kurihara,
Y.; et al. Revision of the Massarineae (Pleosporales, Dothideomycetes). Stud. Mycol. 2015, 82, 75–136. [CrossRef]
Zhu, D.; Luo, Z.L.; Baht, D.J.; Mckenzie, E.H.; Bahkali, A.H.; Zhou, D.Q.; Su, H.Y.; Hyde, K.D. Helminthosporium velutinum and H.
aquaticum sp. nov. from aquatic habitats in Yunnan Province, China. Phytotaxa 2016, 253, 179–190. [CrossRef]
Voglmayr, H.; Jaklitsch, W.M. Corynespora, Exosporium and Helminthosporium revisited new species and generic reclassification.
Stud. Mycol. 2017, 87, 43–76. [CrossRef]
Crous, P.W.; Carnegie, A.J.; Wingfield, M.J.; Sharma, R.; Mughini, G.; Noordeloos, M.E.; Santini, A.; Shouche, Y.S.; Bezerra, J.D.P.;
Dima, B.; et al. Fungal Planet description sheets: 868–950. Pers. Mol. Phylogeny Evol. Fungi 2019, 42, 291–473. [CrossRef] [PubMed]
Rashmi, M.; Kushveer, J.S.; Sarma, V.V. A worldwide list of endophytic fungi with notes on ecology and diversity. Mycosphere
2019, 10, 798–1079. [CrossRef]
Zhao, N.; Luo, Z.L.; Hyde, K.D.; Su, H.Y.; Bhat, D.J.; Liu, J.K.; Bao, D.F.; Hao, Y.E. Helminthosporium submersum sp. nov.
(Massarinaceae) from submerged wood in north-western Yunnan Province, China. Phytotaxa 2018, 348, 269–278. [CrossRef]
Drechsler, C. Some graminicolous species of Helminthosporium. J. Agric. Res. 1923, 24, 641–739.
Drechsler, C. Phytopathological and taxonomical aspects of Ophilobolus, Pyrenophora, Helminthosporium and a new genus Cochliobolus. Phytopathology 1934, 24, 953–985.
Shoemaker, R.A. Nomenclature of Drechslera and Bipolaris, grass parasites segregated from ‘Helminthosporium’. Can. J. Bot. 1959,
37, 879–887. [CrossRef]
Misra, A.P.; Prakash, O. Helminthosporium species occurring on graminaceous hosts in India. Indian J. Mycol. Plant Pathol. 1972, 2,
95–97.
Misra, A.P. Helminthosporium Species Occurring in Cereals and Other Gramineae; U.S.P.L. 480 Project No.A7-CR 133, Grant No.
FG-IN-223Tirhut college of Agriculture, Dholi, Muzaffarpur, Bihar, India; Catholic Press: Ranchi, India, 1973; p. 289.
Shenoy, B.D.; Jeewon, R.; Wu, W.P.; Bhat, D.J.; Hyde, K.D. Ribosomal and RPB2 DNA sequence analyses suggest that Sporidesmium
and morphologically similar genera are polyphyletic. Mycol. Res. 2006, 110, 916–928. [CrossRef]
Bärlocher, F. Molecular approaches promise a deeper and broader understanding of the evolutionary ecology of aquatic
hyphomycetes. J. N. Am. Benthol. Soc. 2010, 29, 1027–1041. [CrossRef]
Manamgoda, D.S.; Cai, L.; McKenzie, E.H.; Crous, P.W.; Madrid, H.; Chukeatirote, E.; Shivas, R.G.; Tan, Y.P.; Hyde, K.D. A
phylogenetic and taxonomic re-evaluation of the Bipolaris-Cochliobolus-Curvularia complex. Fungal Divers. 2012, 56, 131–144.
[CrossRef]
Manamgoda, D.S.; Rossman, A.Y.; Castlebury, L.A.; Crous, P.W.; Madrid, H.; Chukeatirote, E.; Hyde, K.D. The genus Bipolaris.
Stud. Mycol. 2014, 79, 221–288. [CrossRef] [PubMed]
Zhang, Y.; Crous, P.W.; Schoch, C.L.; Hyde, K.D. Pleosporales. Fungal Divers. 2012, 53, 1–221. [CrossRef] [PubMed]
Baschien, C.; Tsui, C.K.M.; Gulis, V.; Szewzyk, U.; Marvanová, L. The molecular phylogeny of aquatic hyphomycetes with affinity
to the Leotiomycetes. Fungal Biol. 2013, 117, 660–672. [CrossRef] [PubMed]
Woudenberg, J.H.; Groenewald, J.Z.; Binder, M.; Crous, P.W. Alternaria redefined. Stud. Mycol. 2013, 75, 171–212. [CrossRef]
Ariyawansa, H.A.; Thambugala, K.M.; Manamgoda, D.S.; Jayawardena, R.; Camporesi, E.; Boonmee, S.; Wanasinghe, D.N.;
Phookamsak, R.; Hongsanan, S.; Singtripop, C.; et al. Towards a natural classification and backbone tree for Pleosporaceae.
Fungal Divers. 2015, 71, 85–139. [CrossRef]
Senanayake, I.C.; Rathnayaka, A.R.; Marasinghe, D.S.; Calabon, M.S.; Gentekaki, E.; Lee, H.B.; Hurdeal, V.G.; Pem, D.; Dissanayake, L.S.; Wijesinghe, S.N.; et al. Morphological approaches in studying fungi: Collection, examination, isolation, sporulation
and preservation. Mycosphere 2020, 11, 2678–2754. [CrossRef]
Jayasiri, S.C.; Hyde, K.D.; Ariyawansa, H.A.; Bhat, J.; Buyck, B.; Cai, L.; Dai, Y.C.; Abd-Elsalam, K.A.; Ertz, D.; Hidayat, I.; et al.
The Faces of Fungi database: Fungal names linked with morphology, phylogeny and human impacts. Fungal Divers. 2015, 74,
3–18. [CrossRef]
Index Fungorum. 2021. Available online: http://www.indexfungorum.org/names/Names.asp (accessed on 30 April 2020).
Dissanayake, A.J.; Bhunjun, C.S.; Maharachchikumbura, S.S.N.; Liu, J.K. Applied aspects of methods to infer phylogenetic
relationships amongst fungi. Mycosphere 2020, 11, 2652–2676. [CrossRef]
Vilgalys, R.; Hester, M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several
Cryptococcus species. J. Bacteriol. 1990, 172, 4238–4246. [CrossRef]
White, T.J.; Bruns, T.; Lee, S.J.W.T.; Taylor, J.W. Amplification and Direct Sequencing of Fungal Ribosomal RNA Genes for Phylogenetics;
Innis, M.A., Gelfand, D.H., Sninsky, J.J., White, T.J., Eds.; PCR Protocols: A Guide to Methods and Applications; Academic Press:
New York, NY, USA, 1990; pp. 315–322. [CrossRef]
Rehner, S. Primers for Elongation Factor 1-α (EF1-α). 2001. Available online: http://ocid.NACSE.ORG/research/deephyphae/
EF1primer.pdf (accessed on 1 November 2019).
Rehner, S.A.; Buckley, E. A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: Evidence for cryptic diversification and links to Cordyceps teleomorphs. Mycologia 2005, 97, 84–98. [CrossRef]
Life 2021, 11, 454
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
49 of 53
Katoh, K.; Standley, D.M. MAFFT multiple sequence alignment software version 7: Improvements in performance and usability.
Mol. Biol. Evol. 2013, 30, 772–780. [CrossRef] [PubMed]
Kumar, S.; Stecher, G.; Tamura, K. MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol. Biol.
Evol. 2016, 33, 1870–1874. [CrossRef] [PubMed]
Glez-Peña, D.; Gómez-Blanco, D.; Reboiro-Jato, M.; Fdez-Riverola, F.; Posada, D. ALTER: Program-oriented conversion of DNA
and protein alignments. Nucleic Acids Res. 2010, 38, 14–18. [CrossRef] [PubMed]
Miller, M.A.; Pfeiffer, W.; Schwartz, T. The CIPRES science gateway: A community resource for phylogenetic analyses. In
Proceedings of the 2011 TeraGrid Conference: Extreme Digital Discovery, Association for Computing Machinery, New York, NY,
USA, 18–21 July 2011; pp. 1–8. [CrossRef]
Huelsenbeck, J.P.; Ronquist, F. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 2001, 17, 754–755. [CrossRef]
Ronquist, F.; Huelsenbeck, J.P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 2003, 19,
1572–1574. [CrossRef]
Nylander, J.A.A. MrModeltest 2.0.; Program distributed by the author; Evolutionary Biology Centre, Uppsala University: Uppsala,
Sweden, 2004.
Rambaut, A. FigTree version 1.4.0. 2012. Available online: http://tree.bio.ed.ac.uk/software/figtree/ (accessed on 1 November
2019).
Schoch, C.L.; Crous, P.W.; Groenewald, J.Z.; Boehm, E.W.A.; Burgess, T.I.; De Gruyter, J.; De Hoog, G.S.; Dixon, L.J.; Grube, M.;
Gueidan, C.; et al. A class-wide phylogenetic assessment of Dothideomycetes. Stud. Mycol. 2009, 64, 1–15. [CrossRef]
Déon, M.; Fumanal, B.; Gimenez, S.; Bieysse, D.; Oliveira, R.R.; Shuib, S.S.; Breton, F.; Elumalai, S.; Vida, J.B.; Seguin, M.; et al.
Diversity of the cassiicolin gene in Corynespora cassiicola and relation with the pathogenicity in Hevea brasiliensis. Fungal Biol. 2014,
118, 32–47. [CrossRef]
Jaklitsch, W.M.; Voglmayr, H. Hidden diversity in Thyridaria and a new circumscription of the Thyridariaceae. Stud. Mycol. 2016,
85, 35–64. [CrossRef]
Lumbsch, H.T.; Lindemuth, R. Major lineages of Dothideomycetes (Ascomycota) inferred from SSU and LSU rDNA sequences.
Mycol. Res. 2001, 105, 901–908. [CrossRef]
Schoch, C.L.; Shoemaker, R.A.; Seifert, K.A.; Hambleton, S.; Spatafora, J.W.; Crous, P.W. A multigene phylogeny of the Dothideomycetes using four nuclear loci. Mycologia 2006, 98, 1041–1052. [CrossRef]
Crous, P.W.; Wingfield, M.J.; Schumacher, R.K.; Summerell, B.A.; Giraldo, A.; Gené, J.; Guarro, J.; Wanasinghe, D.N.; Hyde, K.D.;
Camporesi, E.; et al. Fungal Planet description sheets: 281–319. Pers. Mol. Phylogeny Evol. Fungi. 2014, 33, 212–319. [CrossRef]
[PubMed]
Vu, D.; Groenewald, M.; De Vries, M.; Gehrmann, T.; Stielow, B.; Eberhardt, U.; Al-Hatmi, A.; Groenewald, J.Z.; Cardinali, G.;
Houbraken, J.; et al. Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi
and reveals thresholds for fungal species and higher taxon delimitation. Stud. Mycol. 2019, 92, 135–154. [CrossRef]
Tian, Q.; Li, W.J.; Hyde, K.D.; Camporesi, E.; Bhat, D.J.; Chomnunti, P.; Xu, J.C. Molecular taxonomy of five species of microfungi
on Alnus spp. from Italy. Mycol. Prog. 2018, 17, 255–274. [CrossRef]
Quaedvlieg, W.; Verkley, G.J.M.; Shin, H.D.; Barreto, R.W.; Alfenas, A.C.; Swart, W.J.; Groenewald, J.Z.; Crous, P.W. Sizing up
septoria. Stud. Mycol. 2013, 75, 307–390. [CrossRef] [PubMed]
Crous, P.W.; Wingfield, M.J.; Richardson, D.M.; Le Roux, J.J.; Strasberg, D.; Edwards, J.; Roets, F.; Hubka, V.; Taylor, P.W.J.;
Heykoop, M.; et al. Fungal Planet description sheets: 400–468. Pers. Mol. Phylogeny Evol. Fungi. 2016, 36, 316–458. [CrossRef]
Rosado, A.W.C.; de Jesus Boari, A.; Custódio, F.A.; Quadros, A.F.F.; Batista, I.C.A.; Pereira, O.L. Helminthosporiella stilbacea
associated with African oil palm (Elaeis guineensis) in Brazil. For. Pathol. 2019, 49, e12529. [CrossRef]
Hernández-Restrepo, M.; Schumacher, R.K.; Wingfield, M.J.; Ahmad, I.; Cai, L.; Duong, T.A.; Edwards, J.; Gené, J.; Groenewald,
J.Z.; Jabeen, S.; et al. Fungal systematics and evolution: FUSE 2. Sydowia 2016, 68, 193–230. [CrossRef]
Hashimoto, A.; Matsumura, M.; Hirayama, K.; Tanaka, K. Revision of Lophiotremataceae (Pleosporales, Dothideomycetes):
Aquasubmersaceae, Cryptocoryneaceae, and Hermatomycetaceae fam. nov. Pers. Mol. Phylogeny Evol. Fungi 2017, 39, 51–73.
[CrossRef]
Zhang, Y.; Wang, H.K.; Fournier, J.; Crous, P.W.; Jeewon, R.; Pointing, S.B.; Hyde, K.D. Towards a phylogenetic clarification of
Lophiostoma/Massarina and morphologically similar genera in the Pleosporales. Fungal Divers. 2009, 38, 225–251.
Liew, E.C.; Aptroot, A.; Hyde, K.D. An evaluation of the monophyly of Massarina based on ribosomal DNA sequences. Mycologia
2002, 94, 803–813. [CrossRef]
Kodsueb, R.; Lumyong, S.; Ho, W.H.; Hyde, K.D.; Mckenzie, E.H.; Jeewon, R. Morphological and molecular characterization of
Aquaticheirospora and phylogenetics of Massarinaceae (Pleosporales). Bot. J. Linn. Soc. 2007, 155, 283–296. [CrossRef]
De Gruyter, J.; Aveskamp, M.M.; Woudenberg, J.H.; Verkley, G.J.; Groenewald, J.Z.; Crous, P.W. Molecular phylogeny of Phoma
and allied anamorph genera: Towards a reclassification of the Phoma complex. Mycol. Res. 2009, 113, 508–519. [CrossRef]
Hu, F.J.; Jeewon, R.; Hyde, K.D. Relationships among Astrosphaeriella, Caryospora and Trematosphaeria. Ph.D. Thesis, The University
of Hong Kong, Hong Kong, China, 2009.
Thambugala, K.M.; Hyde, K.D.; Tanaka, K.; Tian, Q.; Wanasinghe, D.N.; Ariyawansa, H.A.; Jayasiri, S.C.; Boonmee, S.; Camporesi, E.; Hashimoto, A.; et al. Towards a natural classification and backbone tree for Lophiostomataceae, Floricolaceae, and
Amorosiaceae fam. nov. Fungal Divers. 2015, 74, 199–266. [CrossRef]
Life 2021, 11, 454
73.
74.
75.
76.
77.
78.
79.
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
91.
92.
93.
94.
95.
96.
97.
98.
99.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
110.
111.
50 of 53
Chethana, T.; Liu, M.; Ariyawansa, H.A.; Konta, S.; Wanasinghe, D.N.; Zhou, Y.; Yan, J.; Camporesi, E.; Bulgakov, T.M.;
Chukeatirote, E.; et al. Splanchnonema-like species in Pleosporales: Introducing Pseudosplanchnonema gen. nov. in Massarinaceae. Phytotaxa 2015, 231, 133–144. [CrossRef]
Thambugala, K.M.; Wanasinghe, D.N.; Phillips, A.J.L.; Camporesi, E.; Bulgakov, T.S.; Phukhamsakda, C.; Dissanayake, A.;
Tennakoon, D.S.; Tibpromma, S.; Chen, Y.Y.; et al. Mycosphere notes 1-50: Grass (Poaceae) inhabiting Dothideomycetes.
Mycosphere 2017, 8, 697–796. [CrossRef]
Crous, P.W.; Shivas, R.G.; Quaedvlieg, W.V.; van der Bank, M.; Zhang, Y.; Summerell, B.A.; Guarro, J.; Wingfield, M.J.; Wood, A.R.;
Alfenas, A.C. Fungal Planet description sheets: 214–280. Pers. Mol. Phylogeny Evol. Fungi 2014, 32, 184–306. [CrossRef]
Wijayawardene, N.N.; Hyde, K.D.; Bhat, D.J.; Goonasekara, I.D.; Nadeeshan, D.; Camporesi, E.; Schumacher, R.K.; Wang, Y.
Additions to brown spored coelomycetous taxa in Massarinae, Pleosporales: Introducing Phragmocamarosporium gen. nov. and
Suttonomyces gen. nov. Cryptogam. Mycol. 2015, 36, 213–224. [CrossRef]
Wanasinghe, D.N.; Phukhamsakda, C.; Hyde, K.D.; Jeewon, R.; Lee, H.B.; Jones, E.G.; Tibpromma, S.; Tennakoon, D.S.;
Dissanayake, A.J.; Jayasiri, S.C.; et al. Fungal diversity notes 709–839: Taxonomic and phylogenetic contributions to fungal taxa
with an emphasis on fungi on Rosaceae. Fungal Divers. 2018, 89, 1–236. [CrossRef]
Knapp, D.G.; Kovács, G.M.; Zajta, E.; Groenewald, J.Z.; Crous, P.W. Dark septate endophytic pleosporalean genera from semiarid
areas. Pers. Mol. Phylogeny Evol. Fungi 2015, 35, 87–100. [CrossRef]
Raja, H.A.; Miller, A.N.; Pearce, C.J.; Oberlies, N.H. Fungal identification using molecular tools: A primer for the natural products
research community. J. Nat. Prod. 2017, 80, 756–770. [CrossRef]
Species Fungorum. 2021. Available online: http://www.speciesfungorum.org/Names/Names.asp (accessed on 30 April 2021).
Goh, T.K.; Hyde, K.D.; Lee, D.K. Generic distinction in the Helminthosporium-complex based on restriction analysis of the nuclear
ribosomal RNA gene. Fungal Divers. 1998, 1, 85–107.
Mukerji, K.G. Current Concepts in Botany; IK International Publishing House Pvt Ltd.: New Delhi, India, 2006; pp. 49–78.
Hernández-Restrepo, M.; Madrid, H.; Tan, Y.P.; da Cunha, K.C.; Gene, J.; Guarro, J.; Crous, P.W. Multi-locus phylogeny and
taxonomy of Exserohilum. Pers. Mol. Phylogeny Evol. Fungi 2018, 41, 71–108. [CrossRef] [PubMed]
Cooke, W.B.; Shaw, C.G. Western fungi. III. Mycologia 1952, 44, 795–812. [CrossRef]
Ellis, M.B. Dematiaceous hyphomycetes. III. Mycol. Pap. 1961, 82, 1–55.
Ciferri, R. Observations on meliolicolous Hyphales from Santo Domingo. Sydowia 1955, 9, 296–335.
Sydow, H.; Sydow, P. Novae fungorum species. Ann. Mycol. 1904, 2, 162–174.
Jørstad, I. Parasitic fungi from the Canaries chiefly collected by J. Lid, with a note on Schizophyllum commune. Blyttia 1966, 24,
222–231.
Ciferri, R. Mycoflora Domingensis Integrata. Quaderno del Laboratorio Crittogamico del Istituto Botanicodell’ Università di Pavia 1961,
19, 1–539.
Petch, T. New Ceylon fungi. Ann. R. Bot. Gard. Perad. 1909, 4, 299–307.
Thirumalachar, M.J. Some new or interesting fungi II. Sydowia 1950, 4, 66–73.
Petrak, F.; Ciferri, R. Fungi Dominicani. II. Ann. Mycol. 1932, 30, 149–353.
Petrak, F. Beiträgezur Pilzflora von Britisch Nord-Borneo. Sydowia 1954, 8, 12–26.
Lavrov, N.P. Trud. tomsk. gos. Univ. Kuibysheva 1951, 110, 254.
Jaczewski, A.L.A. Type species—Jaczewskiella altajensis. Mater. Mikol. Fitopat. Ross. 1915, 1, 41.
Corda, A.C.J. Gliostroma. Icon. Fung. 1837, 1, 13.
Unamuno, L.M. NotasMicológicas. II. Adiciones a los Hifales de la flora española. Boletín de la Real Soc. Española de Hist. Nat.
1932, 32, 161–169.
Saccardo, P.A. Sylloge Fungorum. Mem. Reale Ist. Veneto Sci. 1902, 13, (reprint).
Cooke, M.C. Ravenel’s American fungi. Grevillea 1878, 6, 129–146.
Zaprometov, N.G. Fungal flora of the Kyrgyz SSR. Frunze Acad. Sci. Kyrg. SSR 1957, 1, 98.
Deshpande, K.S.; Deshpande, K.B. Contribution to the taxonomy of genus Helminthosporium. I. Sydowia 1966, 20, 39–45.
Diedicke, H. Aufzählungen der in der Umgebung Erfurts Beobachteten Micromyceten; Academische Loge Sincera Concordia: Erfurt,
Germany, 1910; Volume 36, p. 221.
Hennings, P. Fungi paraënses III. Hedwigia 1908, 48, 101–117.
Sydow, H.; Sydow, P. Weitere Diagnosen neuer philippinischer Pilze. Ann. Mycol. 1916, 14, 353–375.
Zhang, M.; Wu, H.Y.; Wang, Z.Y. Taxonomic studies of Helminthosporium from China 5. Two new species from Hunan and Sichuan
Province. Mycotaxon 2010, 113, 95–99. [CrossRef]
Dounine, M.S.; Yakimovitch, E.D. Sweet Potato Diseases and Their Control; Pan-Soviet Science Research Institute Cultural Soybean
and Spice Crops: Moscow, Russia, 1934; p. 247.
Subramanian, C.V.; Bhat, D.J. Hyphomycetes from South India I. Some new taxa. Kavaka 1987, 15, 41–74.
Misra, A.P. Helminthosporium Species Occurring on Cereals and Other Gramineae; Tirhut College of Agriculture: Dholi, India, 1976;
pp. 1–289.
Matsushima, T. Matsushima Mycological Memoirs 7; Matsushima Fungus Collection: Kobe, Japan, 1993; Volume 7, pp. 1–141.
Pidoplichko, N.M. New fungus species on coarse fodders. Mikrobiol. Zh. 1950, 12, 38.
Viégas, A.P. Algunsfungos do Brasil: XIII—Hifomicetos. Bragantia 1946, 6, 353–442. [CrossRef]
Life 2021, 11, 454
112.
113.
114.
115.
116.
117.
118.
119.
120.
121.
122.
123.
124.
125.
126.
127.
128.
129.
130.
131.
132.
133.
134.
135.
136.
137.
138.
139.
140.
141.
142.
143.
144.
145.
146.
147.
148.
149.
150.
151.
152.
153.
154.
155.
156.
157.
158.
159.
160.
51 of 53
Ciferri, R. Mycoflora domingensis exsiccata (Cent. I, no. 1–100). Ann. Mycol. 1931, 29, 283–299.
Bongini, V. Sur una malattia delle Cactacee. Difesa delle Piante. 1932, 9, 38.
Steyaert, R.L. Contribution à l’étude des parasites des végétaux du Congo. Bull. Soc. R. Bot. Belg. 1948, 80, 11–58.
Hennings, P.; Fungi, S. Paulensis IV a cl. Puttemanscollecti. Hedwigia 1909, 48, 1–20.
Saccardo, P.A. Notae mycologicae. Ser. XXVII. Fungi sinenses aliquot a cl. Prof. Otto A. Reinking collecti et communicati. Philipp
J. Sci. 1921, 18, 595–605.
Ciferri, R. Mycoflora domingensis exsiccata. Ann. Mycol. 1938, 36, 198–245.
McColloch; Pollack. Phytopathology 1946, 36, 991.
Orillo, F.T. An undescribed species of Helminthosporium on kapok in the Philippines. Philipp. Agric. 1955, 38, 548–550.
Olivier, C.; Berbee, M.L.; Shoemaker, R.A.; Loria, R. Molecular phylogenetic support from ribosomal DNA sequences for origin of
Helminthosporium from Leptosphaeria-like loculoascomycete ancestors. Mycologia 2000, 92, 736–746. [CrossRef]
Hennings, P. Fungi. Ann. Musée Congo Belge Bot. 1907, 2, 85–106.
Petrak, F. Beiträgezur Pilzflora von Ekuador. Sydowia 1950, 4, 450–587.
Stevens, F.L. Hawaiian fungi. Bull. Bernice Bishop Mus. 1925, 19, 1–189.
Hughes, S.J. Revision es hyphomycetum aliquot cum appendice de nominibusrejiciendis. Can. J. Bot. 1958, 36, 727–836. [CrossRef]
Sawada, K. Descriptive Catalogue of the Formosan Fungi V.; Report of the Department of Agriculture Government Research Institute
of Formosa; Department of Agriculture Government Research Institute of Formosa: Formosa, Japan, 1931; Volume 51, pp. 1–131.
Ciferri, R.; Fragoso, G. Bulletin of the Royal Spanish Society of Natural History; National Museum of Natural Sciences: Madrid, Spain,
1926; Volume 26, p. 340.
Massee, G.E. Fungi exotici, III. Bull. Misc. Inf. R. Bot. Gard. Kew. 1901, 150–169.
Zhang, M.; Zhang, T.Y. Taxonomic studies of Helminthosporium from China 4. Six new species and a key to Helminthosporium from
China. Mycotaxon 2009, 109, 399–413. [CrossRef]
Zhang, M.; Zhang, T.; Wu, W. Taxonomic studies of Helminthosporium from China III. Three new species in Guangdong Province.
Mycotaxon 2007, 99, 137–142.
Malençon, G.; Bertault, R. Champignons de la Péninsule Ibérique, IV. Les Iles Baleares. Acta Phytotax. Barcinon. 1972, 11, 1–64.
Sawada, K. Descriptive catalogue of Taiwan (Formosan) fungi. Part XI. Spec. Publ. Coll. Agric. Natl. Taiwan Univ. 1959, 8, 1–268.
Chowdhury, S. Notes on fungi from Assam. II. Lloydia 1957, 20, 133–138.
Petrak, F. Petrak’s Lists 5; Commonwealth Agricultural Bureaux: Wallingford, UK, 1930; p. 214.
Nisikado, Y. Ber. Ohara Inst. Landw. Forsch. Kurashiki. 1925, 2, 597–612.
Matsushima, T. Matsushima Mycological Memoirs 5; Matsushima Fungus Collection: Kobe, Japan, 1987; Volume 5, pp. 1–100.
Garbowski, L. Bull. Acad. Polon. Sci. Lett. Cracoviae Cl. Math. Nat. Ser. B. 1923 1924, 15.
Saccardo, P.A. Notae mycologicae series XXIII. Fungi Philippinenses. Attidella Accad. Sci. Veneto-Trentino-Istriana 1917, 10, 57–94.
Sturm, J.W. Deutschl. Fl. 3 Abt. (Pilze Deutschl.) 1831, 3, 21.
Baccarini, P. Funghidell’Eritrea. Ann. Bot. 1906, 4, 269–277.
Nishihara, N. Ann. Phytopath. Soc. Japan 1969, 35, 89.
Sydow, H.; Sydow, P. Fungi novibrasilienses a cl. Ulelecti. Bull. del´Herb. Boissier 1901, 1, 77–85.
Von Thümen, F. Die Pilze des Weinstockes; W. Braumüller: Wien, Austria, 1878; pp. 1–225.
Berkeley, M.J. Notices of British fungi [208–256]. Ann. Mag. Nat. Hist. 1841, 6, 430–439. [CrossRef]
Golovin, P.N. Novye vidy gribov Srednej Azii. Trudy Sredneaz. Univ. 1950, 14, 1–47.
Berkeley, M.J.; Broome, C.E. Notices of British fungi (901–951). Ann. Mag. Nat. Hist. 1861, 7, 373–382.
Roumeguère, C.; Saccardo, P.A. Fungi Algerienses Trabutiani—Sertulum II. Rev. Mycol. Toulouse 1881, 3, 26–32.
Katsuki, S. Notes on parasitic fungi of Yaku island. J. Jap. Bot. 1953, 28, 279–288.
Magnus, P. Ein neues Helminthosporium. Hedwigia 1903, 42, 222–225.
Holubová-Jechová, V. Studies on hyphomycetes from Cuba VI. New and rare species with tretic and phialidic conidiogenous
cells. Ceská Mykol. 1987, 41, 107–114.
Zhang, M.; Zhang, T.Y. Flora Fungorum Sinicorum; Science Press: Beijing, China, 2009; Volume 30, pp. 1–272.
Koorders, S.H. Botanische Untersuchungen. Verh. K. Ned. Akad. van Wet. Afd. Nat. 1907, 13, 1–264.
Berkeley, M.J.; Broome, C.E. Notices of British fungi (1335–1401). Ann. Mag. Nat. Hist. 1873, 11, 339–349. [CrossRef]
Ciferri, R. Notae mycologicae et phytopathologicae Serie II, Nr. 1-14. Riv. Patol. Veg. 1927, 17, 35–40.
Rostrup, E. Flora of Koh Chang. Contributions to the knowledge of the vegetation of the gulf of Siam. Fungi. Bot. Tidsskr. 1902,
24, 355–363.
Hennings, P. Fungi Amazonici IV. a cl. Ernesto Ulecollecti. Hedwigia 1905, 44, 57–71.
Yates, H.S. Some recently collected Philippine fungi, II. Philipp. J. Sci. C. Botany 1918, 13, 361–384.
Trotter, A. Supplementum Universale, Pars X. Myxomycetae, Myxobacteriaceae, Deuteromycetae, Mycelia sterilia. Sylloge
Fungorum. 1931, 25, 1–1093.
Curzi, M. Helminthosporium gibberosporum. C. R. Accad. Lincei. 1931, 6, 146.
Stevens, F.L. Some meliolicolous parasites from Porto Rico. Bot. Gaz. Crawfordsville 1918, 65, 227–249. [CrossRef]
Tucker, C.M. J. Agric. Res. 1926, 32, 391.
Life 2021, 11, 454
52 of 53
161. Wildeman, E.de. Etude de Systématique et de Géographie Botanique sur la Flore du Bas- et du Moyen-Congo. Ann. Musée Congo
Belge Bot.Sér. 5 1907, 2, 85–106.
162. Stevens, F.L.; Dowell, R.I. A Meliola disease of cacao. Phytopathology 1923, 13, 247–250.
163. Viégas, A.P. Algunsmicetos Brasileiros. Bragantia 1947, 7, 25–48. [CrossRef]
164. Errampalli, D.; Saunders, J.M.; Holley, J.D. Emergence of silver scurf (Helminthosporium solani) as an economically important
disease of potato. Plant Pathol. 2001, 50, 141–153. [CrossRef]
165. Zhao, G.C.; Zhao, R.L. The Higher Microfungi from Forests of Yunnan Province; Yunnan Science and Technology Press: Kunming,
China, 2012; p. 564.
166. Siboe, G.M.; Kirk, P.M.; Cannon, P.F. New dematiaceous hyphomycetes from Kenyan rare plants. Mycotaxon 1999, 73, 283–302.
167. Gornostai, V.I. Mikol. Fitopatol. 1972, 6, 154.
168. Cheremisinov. Chaetomium subaffine Sergeeva. Notulae Syst. Sect. Crypt. Inst. Bot. Acad. Sci. U.S.S.R. 1951, 7, 158.
169. Hansford, C.G. Contribution towards the fungus flora of Uganda. V. Fungi Imperfecti. Proc. Linn. Soc. Lond. 1943, 155, 34–67.
[CrossRef]
170. Chowdhury, S. Notes on fungi of Assam. Lloydia 1955, 18, 82–87.
171. Crous, P.W.; Wingfield, M.J.; Burgess, T.I.; Hardy, G.S.J.; Gené, J.; Guarro, J.; Baseia, I.G.; García, D.; Gusmão, L.F.P.; Souza-Motta,
C.M.; et al. Fungal Planet description sheets: 716–784. Pers. Mol. Phylogeny Evol. Fungi 2018, 40, 240–393. [CrossRef]
172. Sydow, H. Novae fungorum species. XXV. Ann. Mycol. 1937, 35, 244–286.
173. Von Szilvinyi, A. Mikrobiologische Boden untersuchungenim Lunzer Gebiet. Zent. Bakteriol. und Parasitenkd. Abt. 2 1941, 103,
133–189.
174. Sousa da Câmara, M. Mycetes aliquot Lusitaniae IX. Agron. Lusit. 1949, 11, 39–73.
175. Roldan, E.F. Philipp. J. Sci. 1936, 60, 121.
176. Cooke, M.C. New British fungi [cont.]. Grevillea 1877, 6, 71–76.
177. Shirouzu, T.; Harada, Y. Lignicolous dematiaceous hyphomycetes in Japan: Five new records for Japanese mycoflora, and
proposals of a new name, Helminthosporium magnisporum, and a new combination, Solicorynespora foveolata. Mycoscience 2008, 49,
126–131. [CrossRef]
178. Shirouzu, T.; Harada, Y. Notes on species of Helminthosporium and its allied genera in Japan. Mycoscience 2004, 45, 17–23.
[CrossRef]
179. Sydow, P.; Sydow, H. Weitere neue Micromyceten der Philippinen-Inseln. Ann. Mycol. 1920, 18, 98–104.
180. Rangel, E.S. Arch. Jard. Bot. Rio de Janeiro. 1902, 2, 71.
181. Castellani, E.; Ciferri, R. Scissioni di generi di licheni sulla base delle caratteristiche del fungo. Atti. Ist. Bot. Univ. Pavia Suppl.
Agli. Ser. 5. 1950, 37.
182. Miles, L.E. Trans. Ill. St. Acad. Sci. 1917, 10, 253.
183. Ciferri, R.; González Fragoso, R. Hongosparásitos y saprofitos de la República Dominicana (4a serie). Boletín de la Real Soc.
Española de Hist. Nat. 1926, 26, 192–202.
184. Saccardo, D. Contribuzione alla micologia veneta e modenense. Malpighia 1898, 12, 201–228.
185. Cooke, M.C. New British fungi. Grevillea 1888, 16, 77–81.
186. Jaczewski, A.L.A. Alternative: Transactions of the Illinois Academy of Science. Microbiol. J. 1929, 9, 166.
187. Wang, X.; Wu, H.; Zhang, M. A new species of Helminthosporium from Jiangsu, China. Mycotaxon 2014, 127, 1–4. [CrossRef]
188. Sydow, H.; Sydow, P. Beitragzur Pilzflora Süd-Amerikas. Hedwigia 1903, 42, 105–106.
189. Dearness, J.; House, H.D. New or noteworthy species of fungi. IV. Bull. N. Y. State Mus. 1925, 266, 57–98.
190. Viennot-Bourgin, G. Champignons nouveaux de la Guinée. Bull. Soc. Mycol. Fr. 1959, 75, 33–37.
191. Hughes, S.J. New Zealand Fungi 27. New species of Guedea, Hadrosporium, and Helminthosporium. N. Z. J. Bot. 1980, 18, 65–72.
[CrossRef]
192. Da Câmara, S.M. Contribution esadmyco floram Lusitaniae. Centuria XI. Bol. Agric. Lisboa 1936, 2, 1–80.
193. Chevassut, G. Sur cinq espècesnouvelles et quelquesespècesrares de Micromycètes parasites du groupe des Adelomycetes
(régions du Languedoc et de Franche-Comté) (Five new species and some rare species of parasitic Micromycetes from the
Adelomycetes group (Languedoc and Franche-Comté) (en)). Bull. Trimest. Société Mycol. France 1992, 108, 101–106.
194. Saccardo, P.A. Micromycetes Americani novi. J. Mycol. 1906, 12, 47–52.
195. Hiroë, I.M. Trans. Tottori Soc. Agric. Sci. 1935, 5, 175.
196. Hennings, P. Fungi Africae orientalis III. Bot. Jahrbücher für Syst. Pflanzengesch. und Pflanzengeogr. 1904, 34, 39–57.
197. Savulescu, T.; Rayss, T. Contribution à l’étude de la mycoflore de Palestine. Ann. Crypt. Exot. 1935, 8, 49–87.
198. Matsushima, T. Microfungi of the Solomon Islands and Papua-New Guinea; Matsushima Fungus Collection: Kobe, Japan, 1971;
pp. 1–78.
199. Castro, C.C.D.; Gutiérrez, A.H.; Sotão, H.M.P. Fungosconidiaisem Euterpe oleracea Mart.(açaizeiro) naIlha do Combu, Pará-Brasil.
Acta. Bot. Bras. 2012, 26, 761–771. [CrossRef]
200. Berg, A. Pamietn. Towarz. Nauk Sci. Paryzu. 1934, 260, 14.
201. Liu, L.J. A New Species of Helminthosporium causing leaf spot disease of sugarcane in Puerto Rico. J. Agr. U. Puerto. Rico. 1971, 55,
12–22. [CrossRef]
202. Cooke, W.B. Western fungi. II. Mycologia 1952, 44, 245–261. [CrossRef]
Life 2021, 11, 454
203.
204.
205.
206.
207.
208.
209.
210.
211.
212.
213.
214.
215.
216.
217.
218.
219.
220.
221.
222.
223.
224.
225.
226.
53 of 53
Bourne, B.A. Mém. Mus. Hist. Nat. 1956, 1087.
Lucas, M.T.; Sousa da Câmara, M. Fungi Lusitaniae. V. Agron. Lusit. 1953, 15, 153–182.
Dearness, J. New or noteworthy North American fungi. Mycologia 1917, 9, 345–364. [CrossRef]
Sydow, H.; Sydow, P. Aufzählungeiniger in den Provinzen Kwangtung und Kwangsi (Süd-China) gesammelter Pilze. Ann. Mycol.
1919, 17, 140–143.
Fresenius, G. Beitr. Mykol. 1863, 3, 50. Available online: ia800500.us.archive.org/33/items/beitrgezurmyko00fres/
beitrgezurmyko00fres.pdf (accessed on 12 May 2021).
Berlese, A.N. Sur le développement de quelques champignons nouveaux ou critiques. Bull. Soc. Mycol. Fr. 1892, 8, 94–110.
Woronichin, N.N. Contribution à la floremycologique du Caucase. Trav. du Musée Bot. de l’Académie des Sci. de Russ. 1927, 21,
87–243.
Stevenson, J.A. Rep. P. Ricoinsul. Agric. Exp. Station. 1919, 1917-18, 137.
Zhang, M.; Zhang, T.; Wu, W. Taxonomic studies of Helminthosporium from China II. Two new species in Sichuan Province.
Mycosystema 2004, 23, 179–182. [CrossRef]
Durieu de Maisonneuve, M.C. Expl. Sci. Alg. Fl. Algér. 1 (Livr. 9) 1848, 1, 321–360.
Patterson, F.W. New species of fungi. Bull. Torrey Bot. Club. 1900, 27, 282–286. [CrossRef]
Saccardo, P.A. Notae mycologicae. Series XXIV. I. Fungi Singaporenses Barkesiani. Bolletino dell’Orto Bot. Regia dell’Universita de
Napoli. 1921, 6, 39–73.
Peck, C.H. Report of the state botanist. Bull. N. Y. State Mus. 1911, 150, 5–100.
Saccardo, P.A. Bulletino dell’orto Botanico della R.; Universitá di Napoli: Napoli, Italy, 1918; Volume 6, p. 23.
Turconi, M. Sopra una nuovamalattia del cacao (Theobroma cacao L.). Atti dell’Istituto Bot. Univ. e Lab. Crittogam. di Pavia. 1920,
17, 1–8.
Ciferri, R.; González Fragoso, R. Hongosparásitos y saprofitos de la RepúblicaDominicana (10a serie). Boletín de la Real Soc.
Española de Hist. Nat. 1927, 27, 165–177.
Hennings, P. Schädliche Pilze auf Kulturpflanzenaus Deutsch-Ostafrika. Notizbl. des Bot. Gart. und Mus. Berl. 1903, 3, 239–243.
Berkeley, M.J.; Broome, C.E. Notices of British fungi (502–537). Ann. Mag. Nat. Hist. 1851, 7, 95–102.
Hennings, P.C. Mission. E. Lauren. 1906, 3, 318.
Alves-Barbosa, M.; Costa, P.M.; Malosso, E.; Castañeda-Ruiz, R.F. Two new species of Dictyosporium and Helminthosporium
(Ascomycota) from the Brazilian Atlantic forest. Nova Hedwig 2017, 105, 65–73. [CrossRef]
Sydow, H.; Sydow, P. Fungi Paraenses. Hedwigia 1910, 49, 78–84.
Wakefield, E.M. New and rare British fungi. Bull. Misc. Inf. R. Bot. Gard. Kew. 1918, 1918, 229–233. [CrossRef]
Ciferri, R.; González Fragoso, R. Parasitic and saprophytic fungi of the Dominican Republic, (11th Series.). Bol. de la Real Soc. Esp.
de Hist. Nat. Madr. 1927, 27, 267–280.
Batista, A.C.; Maia, H.S.; Lima, J.A.; Matta, E.A.F. Moniliales–descrição e revisão de algumasespécies. Atas do Inst. de Micol. Univ.
de Pernamb. Recife 1960, 1, 247–274.