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

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 micro-fungi 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.

1. Introduction

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 [3,4,5,6,7,8,9,10,11]. However, more research on fungal diversity on palms in Thailand is needed.

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 [17,18,19,20,21,22,23]. 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 [20,25,26,27,28,29,30,31,32,33,34,35,36,37]. 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 Helminthosporium-like 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. Details of genes/loci with PCR primers and PCR conditions.

Genes/loci	PCR Primer (Forward/Reverse)	PCR Conditions
LSU	LR0R/LR5	a; 95 °C: 30 s, 55 °C: 50 s, 72 °C: 30 s (35 cycles); b
ITS	ITS5/ITS4
SSU	NS1/NS4
tef1-α	983F/2218R

^a Initiation step of 95 °C: 3 min; ^b Final elongation step of 72 °C: 10 min and final hold at 4 °C.

. 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).

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 [49,50,51]. 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 (Figure 1 and Figure 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).

Table 2. Taxa names, strain numbers and GenBank accession numbers of the sequences used in phylogenetic analyses.

Family	Species	Strain No.	GenBank Accession No.				References
			ITS	LSU	SSU	tef1-α
Corynesporaceae	Corynespora cassiicola	CBS 100,822	-	GU301808	GU296144	GU349052	[54]
Corynesporaceae	Corynespora cassiicola	CCP	KF810854	-	GU296145	-	[54,55]
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]
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]
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]
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]
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]
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.

Table 2. Taxa names, strain numbers and GenBank accession numbers of the sequences used in phylogenetic analyses.

Family	Species	Strain No.	GenBank Accession No.				References
			ITS	LSU	SSU	tef1-α
Corynesporaceae	Corynespora cassiicola	CBS 100,822	-	GU301808	GU296144	GU349052	[54]
Corynesporaceae	Corynespora cassiicola	CCP	KF810854	-	GU296145	-	[54,55]
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]
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]
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]
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]
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]
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.

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 (Figure 1 and Figure 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 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 (Figure 1 and Figure 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 ($\overline{x}$ = 12 μm). Conidiophores (110–)140–175(–215) × (4–)5–7(–8) μm ($\overline{x}$ = 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 ($\overline{x}$ = 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.

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. 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	Strain	LSU
		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 Figure 1B and Figure 2)	CBS 138,902 (MH878637)	C	A	C	C	C	C	T	T	C	T	T	G	C	G	-

. 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.

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 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 ($\overline{x}$ = 14 μm). Conidiophores (60–)165–270(–310) × (5–)7–9(–12) μm ($\overline{x}$ = 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 ($\overline{x}$ = 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.

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 (Figure 1 and Figure 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. Comparison of three strains of Helminthosporiella stilbacea.

No.	Herbarium/ Culture No.	Host (Genus/Family)	Locality	Morphology				References
				Mycelia (μmWide)	Conidiophores (μm)	Conidiogenous Cells (μm)	Conidia (μm)
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	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]
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	[64]
3.	Herbarium: MFLU 20-0521 Culture no.: MFLUCC 15-0813	On dead petiole of Cocos nucifera/Arecaceae	Thailand	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	This study

) 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).

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 (Figure 1 and Figure 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. 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.	Taxa	Host(Genus/Family)	Locality	Morphology	Sequence Data	References
1	H. abietis	Abies sp./Pinaceae	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]
2	H. acaciae	On dead branches of Acacia farnesiana/ Fabaceae	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]
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	Absent	[86]
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]
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	Absent	[85]
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]
8	H. albiziae	On leaves of Albizia lebbeck/Fabaceae	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]
9	H. albiziicola	Albizzia lebbek/Fabaceae	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]
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	Absent	[92]
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]
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	Absent	[18]
17	H. appendiculatum	On branches of the trees	Czechia	Conidiophores simple, fasciculate; Conidia 65 × 11 μm, clavate, curved, blunted, whitish, multi-septate	Absent	[96]
18	H. aquaticum	On submerged decaying wood	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]
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	Absent	[95,97]
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]
23	H. atypicum	On leaves of Triticum sp./Poaceae	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]
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	Present	[21]
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]
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	Absent	[105]
29	H. bataticola	On living leaves of Ipomoea batatas/ Convolvulaceae	Caucasus	No information available	Absent	[106]
30	H. bauhiniae	On dead twigs of Bauhinia tomentosa/ Fabaceae	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]
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	Absent	[107]
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]
38	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]
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]
43	H. carpocrinum	Parasite on perithecia of Meliola funebris on leaves of Omphalea sp./Euphorbiaceae (O. pauciflora)	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]
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]
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	Present	[85,120]
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	Absent	[121]
48	H. chusqueae	On living and dying leaves of Chusquea serrulata/Poaceae	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]
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]
56	H. conidiophorellum	On dead branches of tree	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]
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	Absent	[129]
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]
67	H. cylindricum	On rotten wood	Czech Republic/Bohemia	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]
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]
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	Present	[85]
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]
73	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]
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]
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	Absent	[149]
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]
84	H. endiandrae	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]
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]
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	Present	[22]
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]
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]
102	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	Absent	[128]
103	H. guianense	Meliola guianensis parasitic on mycelium on living leaves of Theobroma cacao/ Malvaceae	Guyana	No information available	Absent	[162]
104	H. heringerianum	Tipuana speciosa/ Fabaceae	Brazil	No information available	Absent	[163]
105	H. hispanicum	On dead corticated twigs of Juglans regia/Juglandaceae	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]
106	H. hispaniolae	On living leaves of Manihot utilissima/ Euphorbiaceae	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]
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	Absent	[67]
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]
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	Absent	[122]
111	H. ipomoeae	On leaves of Ipomoea reptans/ Convolvulaceae	China/Taiwan	No information available	Absent	[130]
112	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]
113	H. italicum	On dead branch of Alnus glutinosa/ Betulaceae	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]
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	Present	[21]
115	H. juglandis	Juglans sp./ Juglandaceae	China, Yunnan	No information available	Absent	[165]
116	H. kakamegense	On dead attached twig of Uvariopsis congensis/ Annonaceae	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]
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]
122	H. leucadendri	On leaves of Leucadendron sp./ Proteaceae	South Africa/Western Cape Province, Helderberg Nature Reserve	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]
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]
124	H. ligustri	On dead branches of Ligustrum quihoui/ Oleaceae	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]
125	H. litseae	Litsea polyantha/ Lauraceae	India/Assam	No information available	Absent	[170]
126	H. livistonae	On leaves of Livistona australis/Arecaceae	Australia/New South Wales, Murramarang National Park	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]
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]
134	H. macilentum	On rotten wood	UK/Great Britain	Conidiophores erect, simple, fusiform, 7–10 septate; Conidia 0.5-0.65 × 0.1 mm	Absent	[176]
135	H. magnisporum	On dead fallen branches of an unknown woody plant	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]
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]
145	H. microsorum	On twigs of Quercus ilex/ Fagaceae	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]
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]
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	Absent	[129]
149	H. nadsonii	On fibers of Gossypium sp./ Malvaceae	Russia	No information available	Absent	[186]
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	Present	[187]
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]
154	H. novae-zelandiae	On dead wood and bark of Vitex lucens/ Lamiaceae	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]
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	Absent	[128]
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]
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]
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	Absent	[128]
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]
165	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]
166	H. palmigenum	On rotten fruit of Cocos nucifera/Arecaceae; On petriole and rachis from reference specimen	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]
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]
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]
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	Absent	[128]
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]
183	H. quercicola	On dead corticated branches of Quercus cf. reticulata/Fagaceae	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]
184	H. quercinum	On dead corticated twigs of Quercus petraea/Fagaceae	Austria/Niederösterreich, Spitzerberg	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, brown to dark brown, with well-defined small pores at the apex, 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	Present	[21]
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]
193	H. solani	On stem of Solanum nigrum/Solanaceae (type); Citrus linella; Leucaena glauac; Solanum dulcamara; S. nigrum; Solanum tuberosum	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]
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	Absent	[213]
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]
198	H. subhyalinum	On living leaves of Phoenix hanceana/Arecaceae	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]
199	H. submersum	On submerged decaying wood	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]
200	H. subsimile	On withered and dead leaves of Bruguiera sexangula (=Bruguiera eriopetala)/Rhizophoraceae	Singapore	Conidiophores 200–250 × 8–9 μm, filiform-fasciculate, brown, septate; Conidia (38–)45–50 × 11–12(–14) μm, brown, 3-distoseptate	Absent	[216]
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	Present	[22]
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]
209	H. varium	On decaying leaves of unidentified plants	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]
210	H. varroniae	On leaves of Varronia sp./Boraginaceae	Puerto Rico	Conidiophores 160–200 × 4 μm; Conidia 27–44 × 6–7 μm, 3-distoseptate	Absent	[223]
211	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]

). 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 [19,33,81,82,83]. Furthermore, we recommend revision of the genus Helminthosporium with fresh collections and DNA sequence data (specifically the ITS region and protein coding genes).