life Article Multi-Gene Phylogeny and Morphology Reveal Haplohelminthosporium gen. nov. and Helminthosporiella gen. nov. Associated with Palms in Thailand and A Checklist for Helminthosporium Reported Worldwide Sirinapa Konta 1,2,3 , Kevin D. Hyde 1,2 , Samantha C. Karunarathna 1 , Ausana Mapook 2 , Chanokned Senwanna 4 , Lucas A. P. Dauner 1 , Chandrika M. Nanayakkara 5 , Jianchu Xu 1 , Saowaluck Tibpromma 1, * and Saisamorn Lumyong 6,7, * 1



 Citation: Konta, S.; Hyde, K.D.; Karunarathna, S.C.; Mapook, A.; Senwanna, C.; Dauner, L.A.P.; Nanayakkara, C.M.; Xu, J.; 2 3 4 5 6 Tibpromma, S.; Lumyong, S. Multi-Gene Phylogeny and 7 Morphology Reveal * CAS Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China; sirinapakonta@gmail.com (S.K.); kdhyde3@gmail.com (K.D.H.); samantha@mail.kib.ac.cn (S.C.K.); luke.dauner1@gmail.com (L.A.P.D.); jxu@mail.kib.ac.cn (J.X.) Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand; phung.ausana@gmail.com School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200, Thailand; chanokned.swn@gmail.com Department of Plant Sciences, University of Colombo, Colombo 00300, Sri Lanka; chandi@pts.cmb.ac.lk Research Center of Microbial Diversity and Sustainable Utilization, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand Correspondence: saowaluckfai@gmail.com (S.T.); saisamorn.l@cmu.ac.th (S.L.) Haplohelminthosporium gen. nov. and Helminthosporiella gen. nov. Associated with Palms in Thailand and A Checklist for Helminthosporium Reported Worldwide. Life 2021, 11, 454. https://doi.org/10.3390/ life11050454 Academic Editor: Arnold J. M. Driessen Received: 16 March 2021 Accepted: 11 May 2021 Published: 19 May 2021 Publisher’s Note: MDPI stays neutral Abstract: Palms (Arecaceae) are substrates for a highly diverse range of fungi. Many species are known as saprobes and many are important plant pathogens. Over the course of our studies of microfungi from palms in Thailand, two new taxa were discovered. Morphological characteristics and phylogenetic analyses of combined ITS, LSU, SSU, and tef1-α sequence data revealed their taxonomic positions within Massarinaceae. There are currently ten genera identified and accepted in Massarinaceae, with the addition of the two new genera of Haplohelminthosporium and Helminthosporiella, that are introduced in this paper. Each new genus is provided with a full description and notes, and each new taxon is provided with an illustration for the holotype. A list of identified and accepted species of Helminthosporium with morphology, host information, locality, sequence data, and related references of Helminthosporium reported worldwide is provided based on records in Species Fungorum 2021. This work provides a micro-fungi database of Haplohelminthosporium, Helminthosporiella, and Helminthosporium which can be modified and validated as new data come to light. Keywords: 4 new taxa; Massarinaceae; morphology; multi-genes; palm fungi; Thailand with regard to jurisdictional claims in published maps and institutional affiliations. 1. Introduction Copyright: © 2021 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ In Thailand, a large number of novel fungi from a variety of hosts have been recently described, adding to the region’s highly known fungal diversity [1,2]. This diversity is supported by various factors, including host–plant species relationships, geography, seasons, air humidity, and temperature. Many interesting fungi from Thai monocotyledons such as bamboo (Poaceae) and Pandanaceae have been described in previous studies, and some new taxa and records of microfungi on palms have been published, especially from the southern region of Thailand [3–11]. However, more research on fungal diversity on palms in Thailand is needed. 4.0/). Life 2021, 11, 454. https://doi.org/10.3390/life11050454 https://www.mdpi.com/journal/life Life 2021, 11, 454 2 of 53 Pleosporales is the largest order in Dothideomycetes [12] with 566 genera in 91 families accepted, while 48 genera have been placed in Pleosporales genera incertae sedis with an estimated stem age of 205 MYA [12,13]. Massarinaceae is a family within Pleosporales introduced by Munk [14] to accommodate the genus Massarina, with M. eburnea being designated as the type species and described based on the sexual morph [15]. Hongsanan et al. [12] and Wijayawardene et al. [13] accepted nine genera in Massarinaceae (Byssothecium, Helminthosporiella, Helminthosporium, Massarina, Pseudodidymosphaeria, Pseudosplanchnonema, Semifissispora, Stagonospora, and Suttonomyces). Helminthosporium has the asexual morph of H. velutinum as the type species. It is characterized by terminal and intercalary conidiogenous cells as well as solitary conidia with distosepta [16]. The members of this genus are commonly found as saprobes and endophytes, but they are often isolated from dead corticated twigs or wood, living leaves, and soils [17–23]. Most Helminthosporium species have been described based on their asexual morph, and only a few species have been described based on both morphs viz., H. massarinum, H. microsorum, H. oligosporum, H. quercicola, H. quercinum, and H. tiliae [19,21,24]. Several species in the Helminthosporium complex are polyphyletic and have been placed in other genera viz. Bipolaris, Curvularia, and Exserohilum within Pleosporales, other families viz. Corynesporaceae, Massarinaceae, and Mycosphaerellaceae within Dothideomycetes, or other unrelated Ascomycetes groups that were initially based on morphological characteristics and later on molecular data, although some species still remain unresolved [20,25–37]. Wijayawardene et al. [13] approximated the number of taxa in Helminthosporium at 416 species. However, this genus was not updated with the DNA sequencesin the most recent monograph. Few previous studies have investigated the Helminthosporium-like taxa from plants, particularly palms, in Thailand. In this study, we were able to isolate Helminthosporiumlike taxa from palms collected in Thailand. Morphology and multi-gene phylogenetic analyses showed two Helminthosporium-like taxa are novel in Massarinaceae. In addition, we provide a checklist of Helminthosporium and the name for Helminthosporiella stilbacea is also validated. 2. Materials and Methods 2.1. Collection, Isolation, and Identification The plant materials containing the fungal structures were collected from Krabi and Prachuap Khiri Khan Provinces, Thailand, from living and dead parts of palm trees (Calamus sp. and Cocos nucifera). Samples were taken to the laboratory for morphological study following the methods provided by Konta et al. [9]. Single spore isolates were obtained following the method of Senanayake et al. [38]. Measurements were taken using an Image Framework program. Illustrations were made in Adobe Photoshop CS6. Specimens and cultures were deposited in the herbarium of Mae Fah Luang University (MFLU) and Mae Fah Luang Culture Collection (MFLUCC). Faces of Fungi and Index Fungorum numbers were registered as outlined in Jayasiri et al. [39] and Index Fungorum [40], respectively. 2.2. DNA Extraction and Amplification (PCR) DNA extraction was performed using the Biospin Fungus genomic DNA extraction kit-BSC14S1 (Bioflux, P.R. China) according to Dissanayake et al. [41]. Partial nucleotide genes were subjected to PCR amplification and sequencing of the large subunit (28S, LSU) [42], the internal transcribed spacer (ITS) [43], the small subunit (18S, SSU) [43], and the translation elongation factor 1-alpha (tef1-α) was performed [44,45]. For primers and conditions, see Table 1. PCR amplification and sequencing were carried out following Konta et al. [9]. The resulting fragments were sequenced in both forward and reverse directions, the generated DNA sequences were analysed, and the consensus sequences were computed using SeqMan software. New sequences generated in this study were deposited in GenBank (Table 2). Life 2021, 11, 454 3 of 53 Table 1. Details of genes/loci with PCR primers and PCR conditions. Genes/loci LSU ITS SSU tef1-α a PCR Primer (Forward/Reverse) LR0R/LR5 ITS5/ITS4 NS1/NS4 983F/2218R PCR Conditions a; 95 ◦ C: 30 s, 55 ◦ C: 50 s, 72 ◦ C: 30 s (35 cycles); b Initiation step of 95 ◦ C: 3 min; b Final elongation step of 72 ◦ C: 10 min and final hold at 4 ◦ C. 2.3. Phylogenetic Analyses The sequences generated in this study were subjected to a BLAST search in GenBank to identify closely related sequences. Sequence data retrieved from GenBank and recent publications were used as references [24]. Sequence data for the ITS, LSU, SSU, and tef1-α regions were analysed both individually and in combination. A total of 93 taxa were used for the combined phylogenetic analyses (ITS, LSU, SSU, and tef1-α) in order to find a natural classification placement. In addition, 103 taxa of ITS and 113 taxa of LSU were used for phylogenetic analyses. For both the individual and combined phylogenetic analyses, Cyclothyriella rubronotata (Cyclothyriellaceae) was selected as the outgroup taxon. Absent sequence data (i.e., ITS, LSU, SSU, tef1-α sequence data) in the alignments were treated with gaps as missing data. Sequence alignments were carried out with MAFFT v.6.864b [46] and were manually improved where necessary. The single gene datasets were combined using Mega7 [47]. Data were converted from fasta to nexus and PHYLIP format with Alignment Transformation Environment online, https://sing.ei.uvigo.es/ALTER/ (accessed on 15 July 2020) [48]. The tree topologies obtained from single gene sequence data were compared prior to the combined gene analysis in order to check for incongruence in the overall topology of the phylogenetic tree. Maximum likelihood (ML) analysis was accomplished using RAxML-HPC2 (v.8.2.12) on XSEDE in the CIPRES Science Gateway platform (http://www.phylo.org) (accessed on 12 May 2020) [49] with GTRGAMMA model and set as 1000 bootstrap replicates. Bayesian analysis was performed at CIPRES using Bayesian analysis on XSEDE (v.3.2.7) as part of the “MrBayes on XSEDE” tool [49–51]. GTR+I+G model was selected by using MrModelTest 2.2 [52] under the Akaike information criterion (AIC) as the best-fit models of the combined dataset for maximum likelihood and Bayesian analysis [52]. Bayesian posterior probabilities (BYPP) were determined by Markov Chain Monte Carlo sampling (MCMC) in MrBayes on XSEDE v.3.2.7. Six simultaneous Markov chains were run for 5,000,000 generations and trees were sampled every 1000th generation. An MCMC heated chain was set with a “temperature” value of 0.20. All sampled topologies beneath the asymptote (25%) were discarded as part of a burn-in procedure; the remaining trees (7502) were used for calculating posterior probabilities in the majority rule consensus tree. Bootstrap support values for ML and BYPP are given near to each node (Figures 1 and 2). The phylogenetic trees were configured in FigTree v1.4.0 [53] and edited using Microsoft Office PowerPoint 2016 and Adobe Photoshop CS6 (Adobe Systems, San Jose, CA, USA). Life 2021, 11, x FOR PEER REVIEW Life 2021, 11, 454 4 of 53 4 of 57 Figure 1. Comparison of the topology of Maximum likelihood majority rule consensus tree for the analyses of some selected Figure 1. Comparison of the topology of Maximum likelihood majority rule consensus tree for the analyses of some Corynesporaceae, Massarinaceae, Massarinaceae, and Perioconiaceae (A) Phylogenetic tree of the dataset selected Corynesporaceae, and isolates. Perioconiaceae isolates. (A) Phylogenetic treefor of ITS the sequence dataset fordata. ITS (B) Phylogenetic tree of the dataset for LSU sequence data. Bootstrap support values for maximum likelihood equal sequence data. (B) Phylogenetic tree of the dataset for LSU sequence data. Bootstrap support values for(ML) maximum to or higher than (ML) 50%, and Posterior Probabilities (BYPP) equal to or greater than 0.90equal are given above each likelihood equalBayesian to or higher than 50%, and Bayesian Posterior Probabilities (BYPP) to or greater than are given above each branch. blue.The Ex-type are bold. The tree is rooted to Cyclothybranch.0.90 Novel taxa are in blue. Ex-typeNovel strainstaxa areare in in bold. tree isstrains rooted toinCyclothyriella rubronotata strains TR, riella rubronotata strains TR, TR9 (Cyclothyriellaceae). TR9 (Cyclothyriellaceae). Life 2021, 11, x FOR PEER REVIEW Life 2021, 11, 454 5 of 53 5 of 57 Figure 2. Maximum likelihood majority rule consensus for the tree analyses of Massarinaceae and sister family PeriocoFigure 2. Maximum likelihood majority rule tree consensus for the analyses of Massarinaceae and sister family sequence data. Bootstrap support values for niaceae isolates based on a dataset of combined ITS, LSU, SSU, and tef1-α Perioconiaceae isolates based on a dataset of combined ITS, LSU, SSU, and tef1-α sequence data. Bootstrap support maximum likelihood equal to or higher(ML) than 50%, Bayesian probabilities (BYPP) equal to or greater than values for(ML) maximum likelihood equaland to or higherposterior than 50%, and Bayesian posterior probabilities (BYPP) equal to oreach greater than Novel 0.90 are given each branch. Novel taxa are inThe blue. Ex-type strains are in bold. The 0.90 are given above branch. taxa are above in blue. Ex-type strains are in bold. tree is rooted to Cyclothyriella is rooted to Cyclothyriella rubronotata strains TR, TR9 (Cyclothyriellaceae). rubronotata tree strains TR, TR9 (Cyclothyriellaceae). Life 2021, 11, 454 6 of 53 Table 2. Taxa names, strain numbers and GenBank accession numbers of the sequences used in phylogenetic analyses. Family GenBank Accession No. Species Strain No. References ITS LSU SSU tef1-α - GU301808 GU296144 GU349052 [54] KF810854 - GU296145 - [54,55] Corynesporaceae Corynespora cassiicola CBS 100,822 Corynesporaceae Corynespora cassiicola CCP Corynesporaceae Corynespora smithii CBS 139,925 KY984299 KY984299 - - [21] Corynesporaceae Corynespora smithii L120 KY984297 KY984297 - KY984435 [21] Corynesporaceae Corynespora smithii L130 KY984298 KY984298 KY984419 KY984436 [21] Corynesporaceae Corynespora smithii L139 KY984300 KY984300 - - [21] Cyclothyriellaceae Cyclothyriella rubronotata TR KX650541 KX650541 - KX650516 [56] Cyclothyriellaceae Cyclothyriella rubronotata TR9 * KX650544 KX650544 KX650507 KX650519 [56] Massariaceae Byssothecium circinans CBS 675.92 - GU205217 GU205235 GU349061 [54] Massarinaceae Byssothecium circinans CBS 675.92 - AY016357 AY016339 - [57,58] Massarinaceae Haplohelminthosporium calami MFLUCC 18-0074 * MT928158 MT928156 MT928160 - This study Massarinaceae Helminthosporium aquaticum MFLUCC 15-0357 KU697302 KU697306 KU697310 - [20] Massarinaceae Helminthosporium aquaticum DLUCC 0758 MG098779 MG098786 MG098795 MG98585 [24] Massarinaceae Helminthosporium austriacum L132 * KY984301 KY984301 KY984420 KY984437 [21] Massarinaceae Helminthosporium austriacum L169 KY984303 KY984303 - KY984439 [21] Massarinaceae Helminthosporium austriacum L137 KY984302 KY984302 - KY984438 [21] Massarinaceae Helminthosporium caespitosum L99 * JQ044429 JQ044448 KY984421 KY984440 [21] Massarinaceae Helminthosporium caespitosum L141 KY984305 KY984305 - - [21] Massarinaceae Helminthosporium caespitosum L151 KY984306 KY984306 - - [21] Massarinaceae Helminthosporium dalbergiae H 4628 LC014555 AB807521 AB797231 AB808497 [19] Massarinaceae Helminthosporium endiandrae CBS 138902 * KP004450 KP004478 - - [59] Massarinaceae Helminthosporium endiandrae CBS 138,902 - MH878637 - - [60] Life 2021, 11, 454 7 of 53 Table 2. Cont. Family GenBank Accession No. Species Strain No. ITS LSU SSU tef1-α References Massarinaceae Helminthosporium endiandrae SM64 MT279335 - - - Unpublished Massarinaceae Helminthosporium endiandrae SM61 MT279339 - - - Unpublished Massarinaceae Helminthosporium endiandrae SM64 MT279340 - - - Unpublished Massarinaceae Helminthosporium endiandrae SM61 MT279336 - - - Unpublished Massarinaceae Helminthosporium endiandrae AKRM1 MN880136 - - - Unpublished Massarinaceae Helminthosporium erythrinicola CBS 145,569 MK876391 MK876432 - - [22] Massarinaceae Helminthosporium genistae L128 KY984308 KY984308 KY984422 - [21] Massarinaceae Helminthosporium genistae L129 KY984309 KY984309 KY984423 - [21] Massarinaceae Helminthosporium genistae L142 * KY984310 KY984310 - - [21] Massarinaceae Helminthosporium hispanicum L109 * KY984318 KY984318 KY984424 KY984441 [21] Massarinaceae Helminthosporium italicum MFLUCC 17-0241 KY797638 KY815015 - KY815021 [61] Massarinaceae Helminthosporium juglandinum L97 KY984322 KY984322 KY984425 KY984445 [21] Massarinaceae Helminthosporium juglandinum L118 * KY984321 KY984321 - KY984444 [21] Massarinaceae Helminthosporium leucadendri CBS 135133 * KF251150 KF251654 - KF253110 [62] Massarinaceae Helminthosporium magnisporum H 4627 * AB811452 AB807522 AB797232 AB808498 [19] Massarinaceae Helminthosporium massarinum KT 1564 * AB809629 AB807524 AB797234 AB808500 [19] Massarinaceae Helminthosporium massarinum KT 838 AB809628 AB807523 AB797233 AB808499 [19] Massarinaceae Helminthosporium microsorum L94 KY984327 KY984327 KY984426 KY984446 [21] Massarinaceae Helminthosporium microsorum L95 KY984328 KY984328 - KY984447 [21] Massarinaceae Helminthosporium microsorum L96 * KY984329 KY984329 KY984427 KY984448 [21] Massarinaceae Helminthosporium oligosporum L92 KY984332 KY984332 KY984428 KY984450 [21] Massarinaceae Helminthosporium oligosporum L93 * KY984333 KY984333 - KY984451 [21] Life 2021, 11, 454 8 of 53 Table 2. Cont. Family GenBank Accession No. Species Strain No. ITS LSU SSU tef1-α References Massarinaceae Helminthosporium oligosporum L106 KY984330 KY984330 - KY984449 [21] Massarinaceae Helminthosporium quercinum L90 * KY984339 KY984339 KY984429 KY984453 [21] Massarinaceae Helminthosporium quercinum L91 KY984340 KY984340 - KY984454 [21] Massarinaceae Helminthosporium solani CBS 365.75 KY984341 KY984341 KY984430 KY984455 [21] Massarinaceae Helminthosporium solani CBS 640.85 KY984342 KY984342 - - [21] Massarinaceae Helminthosporiella stilbacea CPHmZC-01 KX228298 KX228355 - - [63] Massarinaceae Helminthosporiella stilbacea COAD 2126 MG668862 - - - [64] Massarinaceae Helminthosporiella stilbacea MFLUCC 15-0813 * MT928159 MT928157 MT928161 MT928151 This study Massarinaceae Helminthosporium submersum MFLUCC 16-1360 * - MG098787 MG098796 MG098586 [24] Massarinaceae Helminthosporium submersum MFLUCC 16-1290 MG098780 MG098788 MG098797 MG098587 [24] Massarinaceae Helminthosporium submersum DLUCC 0805 MG098781 MG098789 MG098798 - [24] Massarinaceae Helminthosporium syzygii CBS 145,570 * MK876392 MK876433 - - [22] Massarinaceae Helminthosporium tiliae L88 * KY984345 KY984345 KY984431 KY984457 [21] Massarinaceae Helminthosporium tiliae L89 KY984346 KY984346 - - [21] Massarinaceae Helminthosporium tiliae L171 KY984343 KY984343 - KY984456 [21] Massarinaceae Helminthosporium velutinum yone 38 - AB807527 AB797237 AB808502 [19] Massarinaceae Helminthosporium velutinum yone 63 - AB807528 AB797238 AB808503 [19] Massarinaceae Helminthosporium velutinum MFLUCC 15-0423 KU697300 KU697304 KU697308 - [20] Massarinaceae Helminthosporium velutinum MFLUCC 15-0428 KU697299 KU697303 KU697307 - [20] Massarinaceae Helminthosporium velutinum H 4626 LC014556 AB807530 AB797240 AB808505 [19] Massarinaceae Helminthosporium velutinum L117 KY984349 KY984349 - KY984460 [21] Massarinaceae Helminthosporium velutinum L126 KY984350 KY984350 - KY984461 [21] Massarinaceae Helminthosporium velutinum L131 * KY984352 KY984352 KY984432 KY984463 [21] Life 2021, 11, 454 9 of 53 Table 2. Cont. Family GenBank Accession No. Species Strain No. ITS LSU SSU tef1-α References Massarinaceae Helminthosporium velutinum CPC 26297= CBS 141,504 KX306757 KX306785 - - [65] Massarinaceae Helminthosporium velutinum yone 96 LC014558 AB807529 AB797239 AB808504 [19] Massarinaceae Helminthosporium velutinum H 4739 LC014557 AB807525 AB797235 AB808501 [19] Massarinaceae Helminthosporium velutinum L115 KY984347 KY984347 - KY984458 [21] Massarinaceae Helminthosporium velutinum L116 KY984348 KY984348 - KY984459 [21] Massarinaceae Helminthosporium velutinum L127 KY984351 KY984351 - KY984462 [21] Massarinaceae Helminthosporium velutinum L98 KY984359 KY984359 KY984433 KY984466 [21] Massarinaceae Helminthosporium velutinum H 4743 - AB807526 AB797236 - [19] Massarinaceae Helminthosporium velutinum MFLUCC 16-1096 MG098783 MG098791 MG098799 MG098588 [24] Massarinaceae Helminthosporium velutinum MFLUCC 16-1282 MG098784 MG098792 MG098800 MG098589 [24] Massarinaceae Helminthosporium velutinum MFLUCC 17-1707 MG098785 MG098793 MG098801 MG098590 [24] Massarinaceae Helminthosporium velutinum MFLUCC 17-1321 - MG098794 MG098802 MG098591 [24] Massarinaceae Helminthosporium velutinum S-076 KU697301 KU697305 KU697309 - [20] Massarinaceae Helminthosporium velutinum MFLUCC 15-0243 KU697301 KU697305 KU697309 - [20] Massarinaceae Helminthosporium velutinum MFLUCC 16-1300 MG098782 MG098790 - - [24] Massarinaceae Massarina albocarnis CBS119345 LC194503 LC194379 LC194337 LC194416 [66] Massarinaceae Massarina cisti CBS 266.62 * LC014568 AB807539 AB797249 AB808514 [19] Massarinaceae Massarina cisti CBS 266.62 - FJ795447 FJ795490 - [67] Massarinaceae Massarina eburnea CBS 473.64 AF383959 GU301840 AF164367 - [60,68] Massarinaceae Massarina eburnea JCM 14422 LC014569 AB521735 AB521718 AB808517 [19] Massarinaceae Massarina igniaria CBS 845.96 - FJ795452 FJ795494 - [67] Massarinaceae Massarina pandanicola MFLUCC 17-0596 MG646958 MG646947 MG646979 MG646986 [4] Massarinaceae Massarina phragmiticola CBS 110,446 - DQ813510 DQ813512 - [69] Life 2021, 11, 454 10 of 53 Table 2. Cont. Family GenBank Accession No. Species Strain No. ITS LSU SSU tef1-α References Massarinaceae Neottiosporina paspali CBS 331.37 - EU754172 EU754073 - [70] Massarinaceae Pseudodidymosphaeria spartii CBS 183.58 - GU205225 GU205250 - [71] Massarinaceae Pseudodidymosphaeria spartii MFLUCC 13-0273 KP325434 KP325436 KP325438 - [72] Massarinaceae Pseudodidymosphaeria spartii MFLUCC 14-1212 KP325435 KP325437 KP325439 - [72] Massarinaceae Pseudosplanchnonema phorcioides MFLUCC 14-0618 KP683372 KP683373 KP683374 - [72] Massarinaceae Pseudosplanchnonema phorcioides MFLUCC 13-0533 - KM875454 KM875455 - [73] Massarinaceae Pseudosplanchnonema phorcioides L16 KY984360 - KY984434 KY984467 [21] Massarinaceae Pseudosplanchnonema phorcioides MFLUCC 13-0611 KP683375 KP683376 KP683377 - [21] Massarinaceae Semifissispora natalis CPC 25383 KT950846 KT950858 - KT950878 [21] Massarinaceae Semifissispora natalis CBS 140659 - MH878157 - - [21] Massarinaceae Semifissispora rotundata CPC 549 KT950847 KT950859 - - [21] Massarinaceae Semifissispora tooloomensis CBS143431 MG38607 MG386124 - - [21] Massarinaceae Stagonospora perfecta KT 1726A AB809642 AB807579 AB797289 AB808555 [19] Massarinaceae Stagonospora cf. paludosa CBS 130,005 KF251254 KF251757 - - [62] Massarinaceae Stagonospora duoseptata CBS 135,093 KF251255 KF251758 - - [62] Massarinaceae Stagonospora imperaticola MFLUCC 15-0026 KY706143 KY706133 KY706138 KY706146 [74] Massarinaceae Stagonospora multiseptata MFLUCC 15-0449 KX965735 KX954404 - - [74] Massarinaceae Stagonospora paludosa CBS 135088 * KF251257 KF251760 - KF253207 [62] Massarinaceae Stagonospora perfecta CBS 135,099 KF251258 KF251761 - - [62] Massarinaceae Stagonospora perfecta KT 1726A AB809642 AB807579 AB797289 AB808555 [19] Massarinaceae Stagonospora pseudocaricis CBS 135,132 KF251259 KF251763 - - [62] Massarinaceae Stagonospora pseudopaludosa CPC 22,654 KF777188 KF777239 - - [62] Massarinaceae Stagonospora pseudoperfecta KT 889 * AB809641 AB807577 AB797287 AB808553 [19] Massarinaceae Stagonospora sp. CBS 135,096 KF251263 KF251766 - - [62] Life 2021, 11, 454 11 of 53 Table 2. Cont. Family GenBank Accession No. Species Strain No. ITS LSU SSU tef1-α References Massarinaceae Stagonospora tainanensis KT 1866 AB809643 AB807580 AB797290 AB808556 [19] Massarinaceae Stagonospora trichophoricola CBS 136,764 KJ869110 KJ869168 - - [75] Massarinaceae Stagonospora uniseptata CPC 22,150 KF251266 KF251769 - - [62] Massarinaceae Stagonospora uniseptata CBS 135,090 KF251264 KF251767 - - [62] Massarinaceae Suttonomyces clematidis MFLUCC 14-0240 - KP842917 KP842920 - [76] Massarinaceae Suttonomyces rosae MFLUCC 15-0051 MG828973 MG829085 MG829185 - [77] Periconiaceae Periconia byssoides H 4600 LC014581 AB807570 AB797280 AB808546 [19] Periconiaceae Periconia digitata CBS 510.77 LC014584 AB807561 AB797271 AB808537 [19] Periconiaceae Periconia macrospinosa CBS 135,663 KP183999 KP184038 KP184080 - [78] Periconiaceae Periconia pseudodigitata KT 1395 * LC014591 AB807564 AB797274 AB808540 [19] * = The asterisks after the strain number represent the ex-type strains from the holotype specimens. Life 2021, 11, 454 12 of 53 3. Results and Discussion 3.1. Phylogenetic Analyses The individual datasets for ITS and LSU regions comprised selected isolates from closely related families (Figure 1). The RAxML analyses of the ITS dataset yielded the best-scoring trees with a final ML optimization likelihood value of -9830.778478 (Figure 1A). The matrix had 531 distinct alignment patterns with 51.80% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.227770, C = 0.273565, G = 0.243931, T = 0.254733; substitution rates AC = 2.172295, AG = 3.427213, AT = 2.029849, CG = 0.957843, CT = 5.859679, GT = 1.000000; and gamma distribution shape parameter α = 0.350193. In Figure 1A, the novel taxon Haplohelminthosporium calami grouped within Massarinaceae and was well separated from other genera but without good bootstrap support. Helminthosporiella stilbacea (MFLUCC 15-0813) is closely related to Hel.stilbacea (strains CPHmZC-01 and COAD 2126) with 100% ML/1.00 BYPP. The RAxML analyses of the LSU dataset yielded the best-scoring trees with a final ML optimization likelihood value of −4283.882978 (Figure 1B). The matrix had 307 distinct alignment patterns with 12.16% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.246483, C = 0.214075, G = 0.309890, T = 0.229553; substitution rates AC = 1.828869, AG = 4.019496, AT = 3.119987, CG = 0.662100, CT = 12.098644, GT = 1.000000; and gamma distribution shape parameter α = 0.159335. In Figure 1B, the novel taxon Haplohelminthosporium calami was also well separated within Massarinaceae and clustered with Helminthosporium and Helminthosporiella. Helminthosporiella stilbacea (MFLUCC 15-0813) is closely related to Hel. stilbacea (strain CPHmZC-01) with 100% ML/1.00 BYPP. The RAxML analysis of the combined (ITS, LSU, SSU, and tef1-α) dataset yielded a best scoring tree with a final ML optimization likelihood value of -22122.846454 (Figure 2). The matrix had 1363 distinct alignment patterns, with 41.38% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.241467, C = 0.241603, G = 0.271551, T = 0.245380; substitution rates AC = 1.860804, AG = 3.064520, AT = 1.916442, CG = 1.009390, CT = 7.530432, GT = 1.000000; and gamma distribution shape parameter α = 0.183588. In the phylogenetic analyses (Figure 2), twelve genera are included in the tree. The novel taxon of Haplohelminthosporium calami grouped within Massarinaceae without strong bootstrap support. Haplohelminthosporium calami is closely related to H. endiandrae (CBS 138902, MH878637), but this is statistically unsupported. Helminthosporiella stilbacea (MFLUCC 15-0813) constitutes a sister phylogenetic affiliation to Hel. stilbacea (strains CPHmZC-01 and COAD 2126) with 100% ML/1.00 BYPP statistical support. The phylogenetic analyses (Figures 1 and 2) showed several topologies of the tree had generally rather low support (ML ≤50% and BYPP ≤0.90). This reflects the relatively high amount of homoplasy in the data. Most Helminthosporium-like taxa did not have SSU and tef1-α sequence data for the phylogenetic analyses. In the future, divergent time estimations will be needed for Helminthosporium-like taxa to resolve taxonomic confusion and placement. 3.2. Taxonomy 3.2.1. Haplohelminthosporium Konta & K.D. Hyde, gen. nov Index Fungorum number: IF557873; Facesoffungi number: FoF09169 Etymology—Haplo in Greek means single, which refers to the single conidium in each conidiophore. It is a close relative of Helminthosporium. Saprobic on living leaves and petioles of Calamus sp. On living leaves, small spots, circular to irregular, yellow in the beginning, later becoming red brown surrounded by yellow. Colonies on natural substrate forming black patches on the upper leaf, petiole surfaces. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate forming black patches on the upper leaf, petiole surfaces. Mycelium mostly immersed, partly on the surface forming small stroma-like aggregations of red brown pseudoparenchymatous cells. Conidiophores arising singly or fasciculate from stroma Life 2021, 11, 454 13 of 53 cells, erect, simple, unbranched, straight, curved and swollen at apex, septate, thick-walled, cylindrical, smooth, bulbous at base, hyaline in the middle, brown to yellow-brown at 1–2-cells above the base, pale brown to yellow-brown at apical cell. Conidiogenous cells monotretic, terminal, determinate, cylindrical, wide and yellow-brown with a well-defined, small, noncicatrized pore at the apex. Conidia one for each conidiophore, obpyriform to lageniform, straight or curved, smooth, olive-brown, distoseptate, with a dark scar at the base. Type species—Haplohelminthosporium calami Konta & K.D. Hyde Notes: Haplohelminthosporium is established as a monotypic genus with Hap. calami as the type species. ITS phylogenetic analyses separated this genus from other genera, while in the LSU and multigene analyses it clustered with Helminthosporium and Helminthosporiella, but both without good statistical support (Figures 1 and 2). Haplohelminthosporium is presented herein as an asexual morph (hyphomycete) similar to Helminthosporium and Helminthosporiella in that it is hyphomycete with an erect conidiophore, monotretic conidiogenous cell and distoseptate conidia [19,22,63]. The type species of Helminthosporium has pale to dark brown, septate conidiophores, with terminal and intercalary polytretic conidiogenous cells, noncicatrized pores at the apex and upper 3–4 cells, solitary or short catenate conidia that are subhyaline to brown, distoseptate, and is dark brown to black scar at the base [19]. Helminthosporiella has brown to red-brown conidiophores with terminal, polytretic conidiogenous cells, with catenate and easily disarticulating chains of conidia that are medium brown, striated at surface and distoseptate [63]. However, Haplohelminthosporium is distinguished by its unbranched conidiophores arising solitarily or fasciculate from the stroma-like bulbous basal cells that are hyaline in the middle, brown to red-brown at 1–2-cells above the base, pale brown to red-brown and curved at the apical cell with well-defined non-cicatrized small pores and with a single olive-brown conidium arising from each conidiophore (Figure 3). In the BLAST search of GenBank, the closest match of the LSU, ITS, and SSU sequence data were identical to Helminthosporium spp. Based on distinguishing morphological characteristics together with single/multigene phylogenetic analyses we introduce the newly described strain as a new genus Haplohelminthosporium in Massarinaceae. Haplohelminthosporium calami Konta & K.D. Hyde, sp. nov. Index Fungorum number: IF557874, Facesoffungi number: FoF09170, Figure 3 Etymology: Referring to the genus of palm trees Calamus L. Holotype: MFLU 20-0520. Saprobic on living leaves and petioles of Calamus sp. On living leaves, small spots, circular to irregular, yellow in the beginning, later becoming red-brown surrounded by yellow. Colonies on natural substrate forming black patches on the upper leaf, petiole surfaces. Sexual morph: Undetermined. Asexual morph: Mycelium mostly immersed, on the surface forming small stroma-like aggregations of red brown pseudoparenchymatous stromal cells (7–)10–14(–20) µm (x = 12 µm). Conidiophores (110–)140–175(–215) × (4–)5–7(–8) µm (x = 160 × 6 µm, n = 50), wide at the base and apex, macronematous, mononematous, arising singly or fasciculate from the stroma cells, erect, simple, unbranched, straight, curved and swollen at the apex, thick-walled, cylindrical, smooth, bulbous at base, hyaline in the middle, brown to red-brown at 1–2-cells above the base, pale brown to red brown at the last cell of the apex, (3–)4–5(–6) septa. Conidiogenous cells monotretic, terminal, determinate, cylindrical, with well-defined small noncicatrized pores at the apex, wide and yellow-brown at the apex. Conidia (55–)70–100(–120) × (13–)17–20(–23) µm (x = 80 × 20 µm, n = 60), one on each conidiophore, obpyriform to lageniform, straight or curved, smooth, olive-brown, (3–)4–6(–7)-distoseptate, with a dark scar at the base. Life 2021, 2021, 11, 11, 454 x FOR PEER REVIEW Life 15 57 14 of of 53 Figure 3. holotype) (A)(A) The forest in Krabi Province. (B–E) Fresh and herFigure 3. Haplohelminthosporium Haplohelminthosporiumcalami calami(MFLU (MFLU20-0520, 20-0520, holotype) The forest in Krabi Province. (B–E) Fresh and barium palm samples. (F,G) Colonies on living leaf.leaf. (H–L) Conidiophores. (M–U) Conidia. (V,W) Germinated conidia. (X) herbarium palm samples. (F,G) Colonies on living (H–L) Conidiophores. (M–U) Conidia. (V,W) Germinated conidia. Culture on on PDA. (Y)(Y) Conidiophore and conidia ononculture. (X) Culture PDA. Conidiophore and conidia culture.(Z) (Z)Conidiogenesis. Conidiogenesis.(AA) (AA)Conidiophores. Conidiophores. (AB,AC) (AB,AC) Conidia. Conidia. Scale bars: C, E =2 cm, H–W, Y–AC = 50 μm. Scale bars: C, E =2 cm, H–W, Y–AC = 50 µm. Life 2021, 11, 454 15 of 53 Culture characteristics: Culture on PDA, colony yellow-gray-brown at the center, turning dull creamy white toward to margin, smooth, dense, zonate at the margin (Figure 3X). Material examined: THAILAND, Krabi Province, on living leaves and petioles of Calamus sp. (Arecaceae), 14 December 2015, Sirinapa Konta, KHNPR-2 (MFLU 20-0520, holotype); ex-type living culture, MFLUCC 18-0074. Notes: BLAST search of the ITS sequence of the newly described strain (Haplohelminthosporium calami) shows 88.89% similarity with Helminthosporium juglandinum (L118), the LSU sequence shows 98.75% similarity with H. aquaticum (MFLUCC 15-0357), and the SSU sequence shows 99.52% similarity with H. quercinum (L90). Based on ITS phylogenetic analysis, Haplohelminthosporium calami formed a single branch at the basal clades of Helminthosporiella and Helminthosporium (Figure 1A), while based on LSU analysis, Hap. calami clustered together with H. juglandinum (L97), H. endiandrae (CBS 138902, MH878637), and Hel. stilbacea with no strong statistical support for both analyses. The phylogenetic results of the combined dataset indicated that Hap. calami clustered with H. endiandrae (CBS 138902, MH878637) without strong bootstrap support (Figure 2). Comparison of base pair differences between LSU loci for isolates of Hap. calami strains MFLUCC 18-0074 and H. endiandrae strains CBS 138,902 (KP004478; Ex-type from the holotype, and MH878637; sister strain) including gaps showed 1.74% (15/861 bp) differences, and the position of each base pair difference is shown in Table 3. Other H. endiandrae strains (AKMR1, CBS 138902; ex-type from the holotype, and SM61) grouped together in Helminthosporium, as the other strains have an ITS region, but the H. endiandrae (CBS 138902, MH878637) strain that grouped with our new collection lacks the ITS region. Therefore, we compared the morphology of these two species and found that Hap. calami differs from H. endiandrae with respect to its smaller conidiophores ((110–)140–175(–215) × (4–)5–7(–8) vs. 200–300 × 5–7 µm), number of conidiophore septa ((3–)4–5(–6) vs. 8–16 septa), larger conidia ((55–)70–100(–120) × (13–)17–20(–23) vs. (35–)37–45(–57) × (7–)8(–9) µm), solitary conidium per conidiophore, and higher number of distoseptate ((3–)4–6(–7)-distoseptate vs. 3(–4)-distoseptate). The results show the placement of Haplohelminthosporium calami within Massarinaceae, and that this species is distinct from other known species. Therefore, we introduce Hap. Calami as a new species based on both morphological and phylogenetic data. Table 3. Polymorphic nucleotides from sequence data of the LSU loci for isolates of Haplohelminthosporium calami MFLUCC 18-0074 and Helminthosporium endiandrae CBS 138,902 (KP004478, MH878637). Species LSU Strain 6 34 74 270 400 412 419 427 480 484 490 491 524 644 843 Haplohelminthosporium calami (this study) MFLUCC 18-0074 - A A T T T C C A C A T T T G Helminthosporium endiandrae (Ex-type from the holotype) CBS 138,902 (KP004478) - C C C C C T T C T T G C G G H. endiandrae (sister strain in Figures 1B and 2) CBS 138,902 (MH878637) C A C C C C T T C T T G C G - 3.2.2. Helminthosporiella Konta & K.D. Hyde, gen. nov. Index Fungorum number: IF558311, Facesoffungi number: FoF09171 Helminthosporiella Hern.-Restr., Sarria & Crous, in Crous et al., Persoonia 36: 437 (2016), MycoBank MB816988, Nom. inval., Art. 40.3 (Shenzhen) Saprobic on dead petiole of Cocos nucifera.Sexual morph: Undetermined. Asexualmorph:Colony on natural substrate black, hairy. Mycelium mostly immersed, at the surface forming small stroma-like aggregations of dark brown pseudoparenchymatous cells. Conidiophores macronematous, wide at the apex and base, arising singly from the stroma cells, erect, simple, unbranched, straight or flexuous, thick-walled, cylindrical, smooth-walled, dark brown, becoming pale brown at the apex, septate. Conidiogenous cells terminal and intercalary, polytretic, with well-defined thick, pale brown pores. Conidia obpyriform to Life 2021, 11, 454 16 of 53 lageniform, straight or curved, smooth-walled, subhyaline to light brown, distoseptate, with a thick scar at the base. Type species—Helminthosporiella stilbacea Konta & K.D. Hyde Notes: Helminthosporiella was introduced by Crous et al. [63] to accommodate a new combination of Hel. stilbacea Hern.-Restr., Sarria & Crous, in Massarinaceae, the basionym of the type species was not provided a Latin diagnosis [63]. In this paper we accept Helminthosporiella as a distinct genus, presently with a single species Helminthosporiella stilbacea. Since a Latin diagnosis is no longer required, we provide an English diagnosis and priority was given to the previous genus and species names. Furthermore, this study provides the holotype to validate the genus and species, and reports the first host record of Hel. stilbacea associated with coconut tree (Arecaceae) in Thailand. In particular, based on the present morphology and DNA sequence data, Helminthosporiella is identified as a monotypic genus, with Hel. stilbacea as the type species. The members of Helminthosporiella were found associated with leaf spots on oil palm (Arecaceae) [64]. Helminthosporiella stilbacea Konta & K.D. Hyde, sp. nov. Index Fungorum number: IF558312, Facesoffungi number: FoF09172, Figure 4. =Cercospora palmicola f. stilbacea Moreau, Rev. Mycol. 12: 38. 1947 Nom. inval., Art. 39.1 (Shenzhen) ≡Helminthosporiella stilbacea Hern.-Restr., Sarria & Crous, in Crous et al., Persoonia 36: 437. 2016; Nom. inval., Art. 39.1 (Shenzhen) Helminthosporium stilbaceum Moreau ex S. Hughes, Mycol. Pap.48: 38. 1952; Nom. inval., Art. 39.1 (Shenzhen). ≡Exosporium stilbaceum Moreau ex M.B. Ellis, Mycol. Pap.82: 38. 1961; Nom. inval., Art. 39.1 (Shenzhen). =Exosporium stilbaceum var. macrosporum Subramon. & V.G. Rao, Journal of the Annamalai University, part B, Sciences 29: 404. 1971; Nom. inval., Art. 35.1 (Shenzhen). Saprobic on dead petiole of Cocos nucifera.Sexual morph: Undetermined. Asexualmorph: Colony on natural substrate black, hairy. Mycelium mostly immersed, at the surface forming small stroma-like aggregations of dark brown pseudoparenchymatous cells (6–)11–15(–25) µm diam (x = 14 µm). Conidiophores (60–)165–270(–310) × (5–)7–9(–12) µm (x = 200 × 8 µm, n = 30), macronematous, wide at the apex and base, arising singly from the stroma cells, erect, simple, unbranched, straight or flexuous, thick-walled, cylindrical, smooth-walled, dark brown, becoming pale brown at the apex, (4–)12–15-septate. Conidiogenous cells terminal and intercalary, polytretic, with well-defined thick, pale brown pores. Conidia (30–)45–60(–70) × 6–9 µm (x = 50 × 7 µm, n = 30), obpyriform to lageniform, straight or curved, smooth-walled, subhyaline to light brown, 5–8-distoseptate, with a thick scar at the base. Culture characteristics: Culture on MEA, colony yellow-green at the center, turning dull green, pale yellow next, becoming dull green again, pale yellow, and white toward the margin. Colony smooth, dense at the middle, zonate, fluffy at the margin (Figure 4P). Material examined: THAILAND, Prachuap Khiri Khan Province, on dead petiole of Cocos nucifera L. (Arecaceae), 30 July 2015, Sirinapa Konta PJK04gHB (MFLU 20-0521, holotype); ex-type living culture, MFLUCC 15-0813. Life 2021, 11, 454 Life 2021, 11, x FOR PEER REVIEW 17 of 53 17 of 57 Figure 4. 4.Helminthosporiella Figure Helminthosporiellastilbacea stilbacea (MFLU (MFLU 20-0521, 20-0521, holotype) holotype) (A) (A) A A coconut coconut plantation plantation in in Prachuap Prachuap Khiri Khiri Khan Khan Province. Province. (B) Palm Palm samples. samples. (C–E) (C–E) Conidiogenesis. Conidiogenesis. (F–H) (F–H) Conidiophores (at red red arrow arrow are are pores). pores). (I–M) (I–M) Conidia. Conidia. (N,O) (B) Conidiophores (at (N,O) Germinated Germinated conidia. (P) (P) Culture Culture on on MEA. MEA. Scale Scale bars: bars: B B= = 22 cm, conidia. cm, C, C, I–O I–O == 20 20 μm, µm, D–H D–H == 50 50 μm. µm. Life 2021, 11, 454 18 of 53 Notes: Crous et al. [63] introduced a new genus Helminthosporiella with a new combination of Hel. stilbacea based on fresh collections from oil palm (Elaeis oleifera) in Colombia and the second collection of Hel. stilbacea was also collected from oil palm (Elaeis guineensis) in Brazil by Rosado et al. [64]. The full descriptions, illustrations, and sequence data are provided with interesting information as this species causes elliptical necrotic spots with a yellowish halo on living leaves of commercial oil palm plantations [63,64]. However, the type species was invalid because of the basionym lacked a Latin diagnosis [63]. From these, our fresh collection was collected from dead petiole of coconut (Cocos nucifera) and in phylogenetic analysis (Figures 1 and 2), three strains of Hel. stilbacea, including our strain, are grouped together with high bootstrap support. In this study, we therefore provide a holotype from our specimen, and introduce a new species Helminthosporiella stilbacea, complete with an English diagnosis, and validated by using the same name while linking to the valuable information provided from the previous publication of this species. A BLAST search of the ITS sequence of our isolate showed 90.19% similarity with H. velutinum (L131), the LSU sequence showed 97.05% similarity with H. aquaticum (MFLUCC 15-0357), the SSU sequence showed 99.15% similarity with H. quercinum (L90), and the tef1-α sequence showed 92.61% similarity with H. tiliae (L88). These blast results do not match the results of the phylogenetic analyses. The comparison between three strains of Hel.stilbacea (see Table 4) from three collections showed that our collection MFLU 20-0521 has several differences when compared with the other two strains CPHmZC-01 and COAD 2126. Our collection was obtained from a dead petiole, while the two other strains were isolated from living leaves [63,64]. Therefore, our new collection has been provided as a holotype for Hel. Stilbacea. It is also the first geographical record from Thailand, and is a new record of the species from a coconut host (Cocos nucifera). Life 2021, 11, 454 19 of 53 Table 4. Comparison of three strains of Helminthosporiella stilbacea. No. Herbarium/ Culture No. Host (Genus/Family) Morphology Locality Conidiophores (µm) Conidiogenous Cells (µm) Conidia (µm) Mono- or polytretic, integrated, determinate, terminal, cylindrical, 31–67 × 4.5–7 Catenate, obclavate, subcylindrical, occasionally bifurcate, medium brown, 26–83 × 7–10, (1–)3–5(–6)-distoseptate [63] [64] 1. Herbarium: Culture no.: CPHmZC-01 On leaves of Elaeis oleifera/Arecaceae Colombia Hyaline to pale brown, smooth, branched, septate Erect, brown to red-brown, synnematous, septate, compacted, 620–1400 × 19–54, individual hyphae 3–4 wide 2. Herbarium: Culture no.: COAD 2126 On old leaves of Elaeis guineensis/Arecaceae Brazil Hyaline to pale brown, 2–4 Erect, brown, septate, synnematous, 66–201(−770) × 2.5–6(−18) Mono or polytretic, cylindrical, terminal, 18–59 × 4–7 Catenate, subcylindrical, obclavate, brown, 32–83 × 4–11, 2–7-distoseptate Mostly immersed, dark brown Solitarily, erect, unbranched, straight or flexuous, cylindrical, bulbous at base, dark brown, becoming pale brown at the apex, (60–)165–270(–310), (5–)7–9(–12) at the base, 5–8 µm wide at the apex, (4–)12–15 septate Terminal and intercalary with well-defined pores, pale brown Obpyriform to lageniform, straight or curved, light brown, (30–)45–60(–70) × 6–9, 5–8-distoseptate 3. Herbarium: MFLU 20-0521 Culture no.: MFLUCC 15-0813 On dead petiole of Cocos nucifera/Arecaceae Thailand References Mycelia (µmWide) This study Life 2021, 11, 454 20 of 53 4. Conclusions In this study, we introduce the new genus Haplohelminthosporium,with Hap. calami as the type species. In multigene phylogenetic analyses, Hap.calami clustered together with Helminthosporium endiandrae (CBS 138902) without strong good bootstrap support (other H. endiandrae (AKRM1, CBS 138902 (ex-type), SM61) groups together in Helminthosporium). Moreover, we were unable to synonymize H. endiandrae (CBS 138902) under Haplohelminthosporium because H. endiandrae has only LSU sequence data available [60]. In the future, H. endiandrae needs more collections and sequence data to confirm taxonomic placement. Another newly described isolate clusters together with Helminthosporiella stilbacea. Helminthosporiella was introduced by Crous et al. [63] but was invalidated as the type species was not provided with a Latin diagnosis. In this study, we validate Helminthosporiella with Hel. stilbacea as the type species. Moreover, the newly described strain from this study is the first saprobic report of Hel. stilbacea, as this was reported in previous studies as a pathogenic fungus on leaves [63,64]. Moreover, topological nodes in phylogenic analyses showed conflicting results (Figures 1 and 2). Probably, using only single gene ITS or LSU analyses will preclude the establishment of taxonomic placements, while combined gene analyses (including protein coding genes) provide sufficient molecular data to determine the placements. Helminthosporium is generally described as a common saprobe found on leaf or twig litter, and it appears to have a diverse distribution. Occasionally, members of this genus are also described as pathogens, occurring on a wide range of hosts. Comparison of morphology is important for fungal identification [79]. In this study, we provide a checklist for Helminthosporium species reported worldwide including details of each species based on records from Species Fungorum [80] (Table 5). We noted that ten Helminthosporium species have been found on palm substrates (Arecaceae). Although Helminthosporium conidia superficially resemble many genera, such as Drechslera, Bipolaris, and Exserohilum, phylogenetic analyses have provided different results [19,33,81–83]. Furthermore, we recommend revision of the genus Helminthosporium with fresh collections and DNA sequence data (specifically the ITS region and protein coding genes). Life 2021, 11, 454 21 of 53 Table 5. Morphology, host information, locality, sequence data, and related references of Helminthosporium reported worldwide based on the record of Species Fungorum 2021 (bold text present Helminthosporium reported from Arecaceae). No. 1 Taxa H. abietis Host(Genus/Family) Abies sp./Pinaceae Locality Morphology Sequence Data References U.S.A./Washington Conidiophores irregularly branched; Conidia 126–150 × 12–16 µm, fusiform, pointed at both ends, olive-green, 12–15-distoseptate Absent [84] Sierra Leone Conidiophores 140–280 × 7–11 µm, dense, fasiculate, simple, straight or flexuous, sometimes swollen at at the tip, septate, smooth, thick-walled, brown, with well-difinded small pores at the apex; Conidia 31–(44–)49 × 10–(12–)14 µm in widest part, narrowing towards the apex to 3–5 µm, obclavate, straight or flexuous, smooth-walled subhyaline to pale brown, 3–6-distoseptate, with a small dark blackish-brown to black scar at the base Absent [85] Absent [86] H. acaciae On dead branches of Acacia farnesiana/Fabaceae 3 H. acalyphae On leaves of Acalypha angustifolia/Euphorbiaceae Dominican Republic Conidiophores 2.5–4 µm thick, erect, simple, superficial, brown-blackish, septate; Conidia 9–16 × 4–6 µm, one for each conidiophore, ovate-ellipsoid, olivaceous-brown or dull-brown, 2–3-distoseptate 4 H. accedens On living leaves of Dolichos baumii/Fabaceae Namibia Conidiophores 250–300 × 5–9 µm, erect, olive-brown; Conidia 35–57 × 6.5–9 µm, solitary, oblong-fusoid, olive, 3–6-distoseptate Absent [87] Absent [85] 2 5 H. ahmadii On dead branches of Quercus sp./Fagaceae Pakistan Conidiophores 220–650 × 12–15 µm, dense, fasiculate, simple, straight or flexuous, smooth, thick-walled, brown to dark brown, with small pores at the apex, septate; Conidia 95–(110–)150 × 25–30(–38) µm wide inthe broadest part, tapering towards the apex to 5–9 µm, obclavate, sometimes rostrate, straight or flexuous, smooth-walled, brown or dark brown, 5–15-distoseptate, with a dark blackish-brown to black scar at base 6 H. aichrysonis On leaves of Aichryson dichotomum/Crassulaceae Spain No information available Absent [88] 7 H. alatum On dying leaves of Dioscorea alata/Dioscoreaceae Dominican Republic No information available Absent [89] Life 2021, 11, 454 22 of 53 Table 5. Cont. No. 8 Taxa H. albiziae Host(Genus/Family) On leaves of Albizia lebbeck/Fabaceae Locality Morphology Sequence Data References Sri Lanka Conidiophores 70 ×7 µm; Conidia 42–56 × 12 µm, tapering to 4 µm diam. clavate, ends rounded, at the lower end, rough with minute warts, fuliginous, terminal cell paler, strgight or curved below, 3–4-distoseptate Absent [90] India Conidiophores 28–44 × 4.5–6 µm, straight or slightly curved, one-septate at the base; Conidia 23.5–34 × 8–9 µm, pyriform, prolongate at the apex, rounded at the base, pale, cinnamon-brown, 3-distoseptate Absent [91] Absent [92] H. albiziicola Albizzia lebbek/Fabaceae 10 H. allamandae On living leaves of Allamanda cathartica/Apocynaceae Dominican Republic Conidiophores 100–180 × 8–10 µm, solitary or aggregate, curved, simple, dark-brown; Conidia 66–110 × 17–20 µm, clavate, elongate-ellipsoid or subfusoid, erect or curved, gray-brown, 7–10-distoseptate 11 H. alphitoniae On living leaves of Alphitonia sp./Rhamnaceae Malaysia/Mount Kinabalu Conidiophores 250–500 ×5–8 µm, erect, dark-brown; Conidia 25–66 × 8–13 µm, obclavate, erect or curved, yellow-brown or pale olive, 1–6-distoseptate Absent [93] 12 H. aneurolepidii On leaves of Aneurolepidium ramosum/Poaceae Russia/West Siberia No information available Absent [94] 13 H. anomalum From soil U.S.A./Iowa, Utah No information available Present [17,63] 14 H. anonymicum In culture: former Soviet Union Russia No information available Absent [95] 15 H. apiculatum On dry tree of Betula sp. (Betulinum)/Betulaceae Czech Republic Conidiophores fasiculate, flexuous, simple, hyaline; Conidia long, 37 µm, elliptical-fusiform, with color, multi-septate Absent [96] Absent [18] Absent [96] 9 16 H. appatternae From leaves of Cynodon dactylon/Poaceae; from culture India/Maharashtra Conidiophores unbranched, of two types; determinate conidiophores uniform, 182 × 5.2 µm, single, olivaceous, 1–3 septate; indeterminante conidiophores narrower, 208–520 × 7.8 µm, paler and distantly septate at base, gradually broadened into a darker, close septate; Conidia 20.8–152.0 × 7.8 µm, 6–18-distoseptate 17 H. appendiculatum On branches of the trees Czechia Conidiophores simple, fasciculate; Conidia 65 × 11 µm, clavate, curved, blunted, whitish, multi-septate Life 2021, 11, 454 23 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References China/Yunnan Conidiophores 410–580 × 13–17 µm, solitary or in groups of 2–4, erect, flexuous, unbranched, smooth, dark brown paler towards the apex, bulbous at base, 14–23 septate; Conidia 70–80 × 16–18 µm, single, obclavate, straight or curved, pale brown to brown, truncate and cicatrized at base, wider than apex, guttulate, 8–10-distoseptate Present [20] Absent [95,97] H. aquaticum On submerged decaying wood 19 H. arcautei On living leaves Scorpiurus subvillosa/Fabaceae Spain Conidiophores 35–50 × 7–8 µm, erect, simple, cylindrical, brownish-purple, 2–3 septate; Conidia 48–86 × 10.5–11 µm, cylindrical-fusoid, straight or slightly curved, light-brown chestnut, 3–8-distoseptate 20 H. asterinoides On living leaves of Eugenia sp./Myrtaceae Brazil Conidiophores 5–7 µm thick, fasciculate, rhizoid; Conidia 22–24 × 5–6 µm, fusoid, curved, colorless at each bottom, 3-distoseptate Absent [98] 21 H. asterinum On Liquidambar sp./ Altingiaceae U.S.A./Florida Conidiophores erect, simple, septate; Conidia 500–600 × 80 µm, clavate, 3–4-distoseptate Present [99] 22 H. astragali On leaves of Astragalus siversianus/Fabaceae Kyrgyzstan No information available Absent [100] India/Maharashtra Conidiophores 3–7 septate, unbranched, and of two types; shorter conidiophore uniformly wide, 62.4–72.8 × 7.8 µm, brown; longer ones narrow at the base and paler, gradually broadening and darkening towards the apex, 440–680 × 5.2–10 µm; Conidia yellow to brown, darkening at maturity, of two kinds; normal ones 23–93.6 × 26 µm, elliptical with hemispherical edges, widest at the middle, 0–10-distoseptate; a typical conidia abundant, forked or geniculate, septation forked, brown to dark brown, 5–8-distoseptate Absent [101] 18 23 H. atypicum On leaves of Triticum sp./Poaceae Life 2021, 11, 454 24 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 24 H. austriacum On dead corticated twigs of Fagus sylvatica/Fagaceae Austria/Döbling, Kahlenberg, Wien Conidiophores 275–700(–920) µm long, 11.5–19 µm wide at the base, tapering to 7–11 µm near the apex, solitarily or fasciculate, erect, simple, sub-cylindrical, straight or flexuous, thick-walled, smooth, brown to dark brown, paler near the apex, with well-defined small pores at the apex, 1–12 septate; Conidia (30–)35–48(–97) × (10.0–)13.7–16.5(–19.8) µm, tapering to 4.5–6.0 µm at the distal end, obpyriform to lageniform, straight or curved, smooth, pale brown, (4–)5–7(–10)-distoseptate, with a blackish-brown 3–6 µm wide scar at the base 25 H. avenae-pratensis On sheaths of Avena pratensis/Poaceae Germany Conidiophores 300 × 8–11 µm, solitary or fasciculate, dark-chestnut, septate; Conidia 70–107 × 16–21 µm, cylindrical or obclavate, light brown, on both sides paler, 5–11-distoseptate Absent [102] 26 H. bactridis On sheaths of Bactris sp./Arecaceae Brazil/Pará Conidiophores 200 × 3–4.5 µm, septate; Conidia 20–30 × 6–8 µm, fusoid, 6–7-distoseptate Absent [103] 27 H. bakeri On dead stems of Premnavestita sp./Lamiaceae Philippines Conidiophores 500–800 ×12 µm wide at base to below, 10 µm wide, erect, unbranched, dark; Conidia 80–150 × 17–22 µm, solitary, oblong, obclavate, 3–6-distoseptate Absent [104] Absent [105] Absent [106] 28 H. bambusicola On dead culm of Bambusa sp./Poaceae China/Sichuan Conidiophores 55–247 × 4–6 µm, fasciculate or solitary, simple, cylindrical, straight or flexuous, thick walled, smooth, brown, paler towards the apex, with well-defined small pores, 1–2 septate; Conidia 36–66 × 6–11 µm narrowing towards the apex to 2–4.5 µm wide, obclavate, straight or slightly flexuous, thin-walled 1–1.5 µm thick, smooth, pale brown, paler towards the apex, 5–8-distoseptate, scar not distinct at the base 29 H. bataticola On living leaves of Ipomoea batatas/Convolvulaceae Caucasus No information available Present [21] Life 2021, 11, 454 25 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References Sierra Leone Conidiophores 350–110 × 10–15 µm thick at the apex, 15–20 µm thick at the base, dense, fasciculate, simple, straight or flexuous, smooth-walled, dark brown, sometimes paler towards the apex, with well definded, small pores septate; Conidia 55-(86–)145 × 16–(17.2–)18 µm thick in broadest part, tapering to 3–4 µm the apex, obclavate, straight or flexuous, rostrate, smooth-walled, subhyaline to brown, 7–18-distoseptate, with a dark blackish brown to black scar ath the base Absent [85] Absent [107] H. bauhiniae On dead twigs of Bauhinia tomentosa/Fabaceae 31 H. belgaumense On litter, Calamus thwaitesii/Arecaceae India/Karnataka Conidiophores 140–250 × 6–9 µm, erect, straight to flexuous, unbranched, smooth, brown; Conidia 10–15 × 6–11 µm, solitary, dry, sub-spherical, dark brown, truncate at base, roundea at the apex, 1-distoseptate 32 H. bhawanii On leaves of Eragrostis japonica/Poaceae India/Bihar No information available Absent [108] 33 H. bigenum Palmae rotten petiole/Arecaceae Peru No information available Absent [109] 34 H. bondarzewii From grains of Triticum sp. and Secale sp./ Poaceae Russia, Ukraine No information available Present [60,110] 35 H. cacaliae Cacalia sonchifolia/Asteraceae Brazil No information available Absent [111] 36 H. cacaophilum From unfermented Cacao beans, Theobroma cacao/Malvaceae Dominican Republic/Santo Domingo No information available Absent [112] 37 H. cactacearum In young plants of Cereus species/Cactaceae Italy No information available Absent [113] H. caespitiferum Meliola spec. in leaf spots of living leafs of Omphalea pauciflora/ Euphorbiaceae Dominican Republic/Santo Domingo Conidiophores 150–300 × 6.5–8 µm, simple, dark-brown, septate; Conidia 18–42 × 8–11 µm, oblong to fusoid, dark-brown, constrict at septum, (3–)6–7-distoseptate Absent [92] 30 38 Life 2021, 11, 454 26 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 39 H. canephorae Coffea canephora/Rubiaceae Democratic Republic of the Congo/Zaire No information available Absent [114] 40 H. cantareirense On dead stems Brazil/São Paulo Conidiophores 7–12 µm thick, erect, fasciculate; Conidia 50–60 × 8–12 µm, clavate, brown, constrict at septum, 6–8-distoseptate Absent [115] 41 H. cantonense On decaying culms of Bambusa vulgaris/Poaceae China Conidiophores 80–95 × 6 µm; Conidia 50–62 × 8 µm, obclavate, 7–9-distoseptate Absent [116] 42 H. caperoniae On living leaves of Caperonia palustris/Euphorbiaceae Dominican Republic Conidiophores 100–300 × 3.5–5 µm, 2–5 fasciculate, simple, olive-brown; Conidia 22–55 × 4–6 µm, oblong-fusoid or subclavate, rarely cylindrical, yellow or gray-brown Absent [92] Dominican Republic/Santo Domingo Conidiophores 1–4 articulate, 200–350 µm long, very densely fasciculate, erect to sub-erect, straight or slightly irregularly curved, almost straight ot curved, dark-brown to blackish, tip light-colored; Conidia 22–25 ×8–10 µm, 1–4 to each conidiopore, easily falling, ellipsoid to ovoid, with narrowed ends, or basal end narrowed-truncate, apical end rounded to acute, not caudate, central cells from dark-brown to brownish, and cells light brown to yellowish, 2–5-distoseptate Absent [117] 43 H. carpocrinum Parasite on perithecia of Meliola funebris on leaves of Omphalea sp./Euphorbiaceae (O. pauciflora) 44 H. carposaprum On Lycopersicon esculentum/Solanaceae British Guiana, Haiti, Mexico No information available Absent [118] 45 H. ceibae On leaves of Ceiba pentandra/Malvaceae Philippines No information available Absent [119] Present [85,120] Absent [121] 46 H. chlorophorae On dead twigs of Chlorophora regia/Moraceae Sierra Leone Conidiophores 120–270 × 7–10 µm thick at the base, often swollen towards the tip up to 12 µm, single or fasciculate, simple, straight or flexuous, smooth-walled, brown to dark brown, with 1–3 well-definded, small pores, septate; Conidia 52–(73–)102 × 8–(9.5–)11 µm, thick in the widest part narrowing gradually towards the apex to 3–5 µm, obclavate, straight or flexuous, smooth-walled, subhyaline to pale brown, 6–9-distoseptate, with a tather large dark blackish-brown to black scar at the base 47 H. chrysobalani On dry leaves of Chrysobalanus icaco/Chrysobalanaceae Dominican Republic/Bonao Conidiophores up to 6 µm, fasciculate, erect, 2–3 septate; Conidia 25–50 × 3–4 µm, fusoid, 2–4-distoseptate Life 2021, 11, 454 27 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References Ecuador/Tungurahua Conidiophores 200–350 × 4–6 µm, dense, erect, fasciculate, simple, straight or slightly curved, dark-brown or olive, septate; Conidia 32–50 × 9–11 µm, elongate-fusiform, blunt at both ends, curved, rarly straight, gray or olive-brown, 3–4-distoseptate Absent [122] 48 H. chusqueae On living and dying leaves of Chusquea serrulata/Poaceae 49 H. cibotii On leaves of Cibotium sp./Cibotiaceae U.S.A./Hawaii Islands No information available Absent [123] 50 H. ciliare - - No information available Absent [124] 51 H. citri On leaves of Citrus poonensis, Citrus tankart, Citrus ponki, and of Citrus sinensis var. brasiliensis/Rutaceae China/Taiwan No information available Absent [125] 52 H. claviphorum Rotten branch Peru No information available Absent [109] 53 H. cleosmatis On living leaves of Clematis sp./ Ranunculaceae (in foliisvivis Cleosmati soctandri) Dominican Republic Conidiophores 140–250(–300) µm long, 4–5 µm wide, solitary, erect, simple, dark-brown, often becoming paler; Conidia 28–52 × 6.5–9 µm, clavate or fusoid, yellow or pale olive-brownish, (3–)4–5-distoseptate Absent [92] 54 H. clusiae On leaves of Clusiarosa sp./Clusiaceae Dominican Republic Conidiophores 108–128 × 12–16.5 µm effuse, brown-black, irregular at based, or subbulbose, septate; Conidia 26–32 × 10–11.5 µm, fusoid, subfusoid or cylindrical, 4–8-distoseptate Absent [126] 55 H. coffeae On leaves of Coffea liberica/Rubiaceae Ghana Conidiophores 300–400 × 7–8 µm, effuse, nigro-olivaceas, aggregate, erect, cylindrical, rect or flexuous, olives-brown, septate; Conidia 45–55 × 8–10 µm, obovate, 3–5-distoseptate Absent [127] China/Guangxi Conidiophores 60–280 × 7.0–8.5 µm, fasciculate, simple, subcylindrical, straight or flexuous, thick-walled, smooth, dark brown, paler towards the apex, with 1–3 well-defined small pores at the apex, 1–2 septate; Conidia 100–147.5 µm long, 9.5–11 µm diam in the widest part, narrowing towards the apex to 3–4 µm diam, straight or slightly flexuous, smooth-walled, pale brown, sometimes verruculose at apex, 11–17-distoseptate, with a large dark blackish-brown scar at the base, 2–3 µm thick Absent [128] 56 H. conidiophorellum On dead branches of tree Life 2021, 11, 454 28 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 57 H. constrictum On dead branches of Trachycarpus fortunei/Arecaceae China/Guangdong Conidiophores single, simple, subcylindrical, straight or slightly flexuous, brown to dark brown, paler towards the apex, 1–3 septate; Conidia 57–120 × 9–12 µm, thick in the widest part, narrowing toward the apex to 2.5–5 µm, abruptly tapered to a truncate base, tretic, obclavate, straight or slightly flexuous, pale brown, paler toward to apex, 9–15-distoseptate, sometimes constricted at one or two septa 58 H. conviva On Hyphoderma caliciferum, the genus of crust fungi in the family Meruliaceae. Spain/Archipelago/ Balearic/Baleares Islands No information available Absent [130] 59 H. corchori On leaves of Corchorus capsularis/Malvaceae China/Taiwan No information available Absent [131] 60 H. crassiseptum Meliola abrupta Dominican Republic Conidiophores 30–50 × 2–3 µm, septate; Conidia 45–55(–65) × 12–14 µm, ovoid or elliptical, (2–)3-distoseptate Absent [86] 61 H. crotalariae On leavesof Crotalaria juncea/Fabaceae India/Assam No information available Absent [132] 62 H. crus-galli On living leaves of Echinochloa crus-galli (=Panicum crista-galli)/Poaceae Japan No information available Absent [133,134] 63 H. cubense On rachis of Roystonea regia/Arecaceae Cuba No information available Absent [135] 64 H. cucumerinum On living leaves of Cucumis sativus/Zingiberaceae Russia/Krym No information available Absent [136] 65 H. curvulum On decaying leaves of Zea mays/Poaceae Philippines Conidiophores 160–180 × 7–7.5 µm, fasciculate, filiform, septate; Conidia 25–35 × 8–9 µm, oblong-fusoid, narrow, 3(–4)-distoseptate Absent [137] 66 H. cuspidatum On decaying branches of Afzelia rhomboidea/Fabaceae Philippines Conidiophores 800–900 × 8–9 µm, fasciculate, filiform, multiseptate; Conidia 100–130 × 11–12 µm, obclavate, 8–12-distoseptate Absent [137] Absent [129] Life 2021, 11, 454 29 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References Conidiophores 100–130 × 4–5 µm, subfasciculate, filiform long, simple, fuliginous up paler, septate; Conidia 14–15 × 2.5 µm, cylindrical, apex rounded, base acuted, minute, pale fuliginous, 3-distoseptate Absent [138] 67 H. cylindricum On rotten wood Czech Republic/Bohemia 68 H. cymmartinii On leaves of Cymbopogon martinii/Poaceae India/Uttar Pradesh No information available Absent [108] 69 H. cyperi On Cyperus sp./ Cyperaceae Greece Conidiophores straight to subflexuous, greenish, paler at apex; Conidia 78 × 9 µm, fusoid, fuscidull, 5–8-distoseptate Absent [139] 70 H. dactylidis On leaves of Dactylis glomerata/Poaceae U.S.A./Pennsylvania No information available Absent [140] Present [85] 71 H. dalbergiae On dead branches of Dalbergia sissoo/Fabaceae Pakistan Conidiophores 300–1300 × 10–12(–15) µm, dense, fasciculate, simple, flexuous, smooth-walled, brown to dark brown, sometimes paler towards the apex, with well-definded small pores, septate; Conidia 58–(93–)125 × 12–(13.2–)14 µm thick in broadest part, tapering to gradually towards the apex to 3–5 µm, obclavate, straight or flexuous, smooth-walled, straw-coloured to pale brownwith, 5–17-distoseptate, large dark blackish-brown to black scar at the base 72 H. davillae On leaves of Davilla rugosa/Dilleniaceae U.S.A./San Francisco Conidiophores 4–6 µm, thick filiform, flexuous, unbranched, elongate, brown, septate; Conidia 40–70 × 4–6 µm, elongate-obclavate, narrower and paler, (1–)2–4-distoseptate Absent [141] H. decacuminatum In the dry twigs on Vitis vinifera/Vitaceae Italy Conidiophores 4 µm thick, extremely short-articulated, irregular, dark reddish-brown; Conidia 40–45 × 10 µm, long clavate, decacumina to tip, or cut down in pedicellum narrowed, pale brown-gray, 4–5-distoseptate Present [60,142] 74 H. delicatulum On stems of Umbelliferae or Apiaceae UK/Great Britain Conidiophores slender, subulate, multi-articultate, brown, paler at the tips; Conidia oblong, nearly colourless, with the apices very obtuse, consisting of about five swollen articulations, one or two of which have occasionally a vertical dissepiment Absent [143] 75 H. delphinii On stems of Delphinium brunonianum/Ranunculaceae Russia No information available Absent [144] 73 Life 2021, 11, 454 30 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 76 H. dendroideum On Acer sp./Sapindaceae U.S.A./South Carolina Conidiophores 1–2 short branchlets termintated, oblong, subfusiform, slightly curved, multiarticulate conidia; Conidia 60 µm long, each joint containing a globose nucleus Absent [145] 77 H. densum - - No information available Absent [146] 78 H. desmodii On Desmodium buergeri/ Fabaceae Japan No information available Absent [147] 79 H. diedickei No information available No information available No information available Absent [148] Absent [149] 80 H. dimorphosporum On decaying rotting stems of unknown liana Cuba Conidiophores 150–400 µm long, at the apex 9–12 µm, at the base 10–14 µm wide, single or fasciculate 2–10, simple, straight or flexuous, smooth, dark brown, paler towards the apex, septate; Conidia of two different types arising through pores a t the apex (1–4 pores) and late rally beneath the upper septa: (a) 19–24 × 8–10.5 µm, broadly ellipsoidal, ovoid or broadly fusiform, thick-walled, smooth, brown to dark brown, 1-distoseptate; (b) 24–65 µm long, 10–15 µm wide in the broadest part, tapering to 3.2–4.8 µm at the apex, obclavate, rostrate, straight or flexuous, pale brown, smooth, 6–9-distoseptate, with a dark brown scar at the base 81 H. dolichi On living leaves of Dolichos sp./Fabaceae Namibia Conidiophores 250–350 × 4–6 µm, erect, olive-brown; Conidia 27–38 × 5.5–8 µm, solitary, oblong-subfusoid, olive, 2–3-distoseptate Absent [87] 82 H. dongxingense Rhododendron sp. China No information available Absent [150] 83 H. elasticae - - No information available Absent [151] On leaves of Endiandra introrsa/Lauraceae Australia/New South Wales, Nightcap National Park Conidiophores 200–300 × 5–7 µm, solitary, erect, subcylindrical, straight to flexuous, unbranched, thick-walled, base bulbous, lacking rhizoids, brown, 8–16 septate; Conidia (35–)37–45(–57) × (7–)8(–9) µm, solitary or in short chains (2–3), obclavate, thick-walled, finely roughened, brown, 3(–4)-distoseptate Present [21,59] 84 H. endiandrae Life 2021, 11, 454 31 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 85 H. eragrostiellae On inflorescence and leaves of Eragrostis bifida/Poaceae India/Uttar Pradesh No information available Absent [108] 86 H. erythrinae On leaves of Erythrina suberosa/Leguminosae India/Karnataka Conidiophores 32–42 × 4–5 µm, simple, brownish-yellow; Conidia 39–62 µm at base, straight or vermiform, rounded at the apex and flat at the base, pale cinnamon-brown, 4–8-distoseptate Absent [91] Present [22] 87 H. erythrinicola On leaves of Erythrina humeana/Fabaceae South Africa/Eastern Cape Conidiophores 500–1200 × 6–10 mm, fasciculate, subcylindrical, unbranched, brown, becoming pale brown at apex, multiseptate; Conidia (70–)80–90(–110) × (9–)10–11(–12) mm, obclavate, straight to curved, apex subobtuse, smooth, medium brown, (6–)7–8(–12)-distoseptate 88 H. exasperatum On Dianthus barbatus/ Caryophyllaceae UK/Great Britain Conidiophores flexuous, knotted above, each knot bearing oblong conidia; Conidia 30–45 × 10–12 µm Absent [152] 89 H. feijoae On leaves of Acca sellowiana/Myrtaceae (syn: Feijoa sellowiana) North America/Hispaniola island No information available Absent [153] 90 H. ferrugineum On leaves of Hiraea sp. and Heteropterys sp./Malpighiaceae U.S.A./San Francisco Conidiophores 8–9 µm thick, filiform, yellow, septate; Conidia 50–62 × 11–14 µm, obclavate, subhyaline, last 2 septate hyaline-yellow to yellow Absent [141] 91 H. fici On leaves of Ficus retusa/Moraceae Philippines, Thailand Conidiophores fusciculate, long, nodulosis, septate; Conidia 18–20 × 5–6 µm, cylindrical, reddish-brown, 3-distoseptate Absent [137,154] 92 H. ficinum On leaves of Ficus ulmifolia/Moraceae Philippines Conidiophores 250 × 6 µm, filiform, septate; Conidia 50–60 × 6–8 µm, obclavate, 4–5-distoseptate Absent [137] 93 H. filicicola On leaves of Lygodium sp./Lygodiaceae and of Selaginella sp./Selaginellaceae Peru Conidiophores 400 × 3–5 µm thick, erect, simple, filiform, septate; Conidia 30–40 × 6–10 µm, cylindrical-fusoid or clavate, both side blunt, 3–5-distoseptate Absent [155] 94 H. flagellatum On mycelium of Meliola, in leaves of Ardisia disticha/Myrsinaceae Philippines Conidiophores 2.5–4 µm thick, erect, sub-hylaline Absent [156] 95 H. flumeanum On leaves of Bambusa sp./Bambuseae Philippines Conidiophores 90–100 × 6–7 µm, dense, fasticulate, filiform; Conidia 35–40 × 9–12 µm, obclavate, 3-distoseptate Absent [157] Life 2021, 11, 454 32 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 96 H. fumagineum On leaves of ficusulmifolia/ Moraceae Philippines Conidiophores 240–300 × 7 µm, filiform, septate; Conidia 35 × 9–10 µm, oblong-obclavate, 3-distoseptate Absent [137] 97 H. gibberosporum Musa cavendishii/Musaceae Somalia No information available Absent Present [158] 98 H. glabroides On Meliola glabroides, on Piper aduncum/ Piperaceae Puerto Rico Conidiophores 100–140 × 7 µm; Conidia 40–81 × 6–7 µm, 3–6-distoseptate Absent [159] 99 H. gleicheniae On leaves of Dicranopteris linearis (=Gleichenia dichotoma)/ Gleicheniaceae U.S.A./Hawaii Islands No information available Absent [123] 100 H. gossypii On living leaves and bracts of Gossypium sp./ Malvaceae North America Conidiophores 40–185 × 6.5–8.5 µm, singly or in groups of three to six, straight cylindrical to nodose or bent, brown, 5 septate; Conidia 35–118 × 11.7–18.4 µm, elliptical, curved, rarely straight, light to dark fuliginous, thick walled, rounded at the ends, 1–8-distoseptate Absent [160] 101 H. grewiae On leaves of Grewia sp./Malvaceae Democratic Republic of the Congo Conidiophores 80–120 × 5–8 µm, fasciculate, septate; Conidia 35–45 × 8–10 µm, fusoid, 2–4-distoseptate Absent [161] Absent [128] Absent [162] 102 103 H. guangxiense On dead branches of unidentified tree China/Guangxi, Shanglin Conidiophores 330–850 µm long, 15–20 µm wide just above the base and 8–13 µmwide toward the apex, fasciculate, simple, straight or flexuous, sub-cylindrical, thick-walled, smooth, brown, with 1–3 well-defined small pores at the apex, 1–4 septate; Conidia 76–110 µm long, 16–22 µm wide in the widest part, narrowing towards the apex to 3–6µm wide, straight or curved, obclavate, smooth, middle brown, paler towards the apex, 9–17-distoseptate, with a large dark blackish-brown scar at the base, 1.5–3.5 µm thick H. guianense Meliola guianensis parasitic on mycelium on living leaves of Theobroma cacao/ Malvaceae Guyana No information available Life 2021, 11, 454 33 of 53 Table 5. Cont. No. 104 105 106 Taxa H. heringerianum Host(Genus/Family) Tipuana speciosa/ Fabaceae H. hispanicum On dead corticated twigs of Juglans regia/Juglandaceae H. hispaniolae On living leaves of Manihot utilissima/ Euphorbiaceae Locality Morphology Sequence Data References Brazil No information available Absent [163] Asturias, Selviella, Spain Conidiophores 130–540 µm long, 13–22.5 µm wide at the base, tapering to 8–15 µm near the apex, solitarily or in small groups, erect, simple, straight or flexuous, thick-walled, subcylindrical, smooth, dark to blackish brown, paler near the apex, with well-defined small pores at the apex, 1–2 septate; Conidia 69–99(–130) × (17–)18–21(–24) µm, obclavate, straight or flexuous, thin-walled, smooth, pale brown, (4–)6–11(–14)-distoseptate, with a blackish-brown 4–6 µm wide scar at the base Present [21] Dominican Republic/Haiti Conidiophores sub-hyaline to light-grey, when old, with an almost hyaline tip; Conidia 14.8–(53.5–)81.4 × 7.4–(11–)14.8 µm, sub-hyaline to smoky, irregular, cylindric-elongate to ellipsoidal, straight or slightly curved, with the basal end applanate, 1–8-distoseptate Absent [112] Absent [67] 107 H. hunanense On dead branches of unidentified tree China/Zhangjiajie, Hunan Conidiophores 70–226 × 5–7 above, 8.5–14 µm base, solitary or fasciculate, simple, cylindrical, straight or flexuous, thick-walled, smooth, brown, well-defined small pores at the apex, 1–3 septate; Conidia 56–127 × 10–14 base, apex 2–4 µm, obclavate, straight or curved, smooth, middle brown, paler towards the apex, 4–12-distoseptate, blackish-brown scar at the base, 1.5 µm thick 108 H. hygrophilae On leaves of Hygrophila brasiliensis/Acanthaceae Dominican Republic No information available Absent [89] 109 H. insigne On leaves of Mallotus philippensis/ Euphorbiaceae Philippines Conidiophores 600–800 × 50 µm, fasciculate, filiform, blackish, septate; Conidia 45–55 × 7–8 µm, obclavate, often curved, 4–5-distoseptate Absent [137] Life 2021, 11, 454 34 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 110 H. insuetum On living leaves of Philodendron sodiroi (=Piplocarpha sodiroi)/Araceae Ecuador/Pichincha Conidiophores 2.5–5 µm thick, olive brown or dark brown; Conidia 17–38 × 7–12 µm, oblong, ellipsoid or oblong-ellipsoid fusiform and often subclavate, rarely cylindrical, often straigtly, rarely curved, olive brown or dark-brown, (3–)5–7(–9)-distoseptate, scared or a little more often in the middle constricted 111 H. ipomoeae On leaves of Ipomoea reptans/ Convolvulaceae China/Taiwan No information available Absent [130] H. iranicum On living leaves of Indigofera sp./Fabaceae Iran/Bandar Abbas Conidiophores 40–75 × 6-9 µm, dense, curved, rarely straight, dark-brown, septate; Conidia 36(–42) × 7–11 µm, oblong, narrowly ellipsoid or curved, obtuse at both ends, straight or curved, sometimes irregular, olive, 1–3-distoseptate Absent [164] Italy Conidiophores (190–)330–600 × (12–)16–18(−20) µm, aggregated, erect, straight or slightly flexuous, unbranched, cylindrical, dark brown, 13–25 septate; Conidia 58–78 × 15–19(−23) µm, obclavate, straight or curved, pale brown to brown, slightly truncate and black at base, rounded, narrowed, 6–11-distoseptate Absent [61] Present [21] Absent [165] 112 113 H. italicum On dead branch of Alnus glutinosa/Betulaceae 114 H. juglandinum On dead corticated twigs of Juglans regia/Juglandaceae Austria/ Niederösterreich/Gießhübl, Italy Conidiophores (175–)215–325(–455) µm long, 11–23 µm wide at the base, 8.5–14 µm wide near the slightly inflated apex, fasciculate, erect, simple, straight or flexuous, thick-walled, sub-cylindrical, smooth, brown to dark brown, darker to black at the apex, the latter with a well-defined apical pore; Conidia (69–)89–145(–205) × (15.0–)16.5–20.0(–25.0) µm, rostrate, straight or flexuous, thin-walled, smooth, pale brown, (5–)9–17(–20)-distoseptate, blackish-brown scar at the base 115 H. juglandis Juglans sp./Juglandaceae China, Yunnan No information available Absent [122] Life 2021, 11, 454 35 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References Kenya Conidiophores 250–550 × 8–12 µm, solitary, unbranched; Conidia 30–90 × 8–10 µm, in the broadest part, uniformly tapering to 2–4 µm wide at at the apex, solitary, simple straight or somewhat curved, obclavate, rostrate, subhyaline, smooth, 4–15-distoseptate Absent [166] 116 H. kakamegense On dead attached twig of Uvariopsis congensis/ Annonaceae 117 H. kalakadense On dead unidentified twig India/Tamil Nadu Conidia 13–15 µm Absent [21] 118 H. kalopanacis On dead wood of Kalopanax septemlobus/ Araliaceae Russia/Primorye No information available Absent [167] 119 H. kok-saghyz In seeds of Taraxacum kok-saghyz/Asteraceae Russia No information available Absent [168] 120 H. kyllingae Kyllinga sp./ Cyperaceae Uganda No information available Absent [169] 121 H. lablab On leaves of Dolichos lablab/Fabaceae China/Taiwan No information available Absent [130] On MEA and PDA Conidiophores 100–300 × 4–6(–7) µm, erect, subcylindrical, thick-walled, medium brown, multiseptate; Conidia (35–)70–110(–170) × (6–)7–8(–11) µm, obclavate to subcylindrical, straight to slightly curved, thick-walled, medium brown, (3–)4–6(–10)-distoseptate Present [21] 122 H. leucadendri On leaves of Leucadendron sp./ Proteaceae South Africa/Western Cape Province, Helderberg Nature Reserve 123 H. leucosykes On Meliola, on leaves of Leucosyke capitellata/ Urticaceae Philippines Conidiophores 300 × 7–8 µm, erect, brown, septate; Conidia 30 × 8 µm, 3-distoseptate Absent [156] China/Guangxi, Nanning Conidiophores 127–700 µm long, 9.5–18 µm diam just above the base and 8.5–10 µm diam towards the apex, solitary, simple, straight or flexuous, smooth or verruculose, thickwalled, dark brown, with 1–3 well-defined small pores at the apex, 1–4 septate; Conidia 24–38.5 × 9.5–13 µm, obclavate, straight or slightly curved, rostrate or pseudorostrate, smoothwalled, pale brown, subhyaline towards the apex, 4–6-distoseptate, with a large dark blackish-brown scar at the base, 1–2 µm thick Absent [128] 124 H. ligustri On dead branches of Ligustrum quihoui/ Oleaceae Life 2021, 11, 454 36 of 53 Table 5. Cont. No. 125 Taxa H. litseae Host(Genus/Family) Locality Morphology Sequence Data References Litsea polyantha/ Lauraceae India/Assam No information available Absent [170] Conidiophores 500 × 4–6 µm, erect, flexuous, cylindrical, smooth to rough-walled, medium brown, multiseptate; Conidia (25–)40–55(–65) × (7–)8–9 µm, subcylindrical, straight, smooth, medium brown, apex obutuse, base somewhat obconic, (3–)4–6(–7)-distoseptate Present [171] 126 H. livistonae On leaves of Livistona australis/Arecaceae Australia/New South Wales, Murramarang National Park 127 H. longisinuatum Palmae rotten trunk Peru Conidiophores 20–75 × 3.5–5 µm; Conidia 65–220(–1000) × 8–10.5 µm, solitary, long, narrowly obclavate, 9–22-distoseptate Absent [109] 128 H. lonicerae On Lonicera sp./ Caprifoliaceae Brazil No information available Absent [111] 129 H. lophirae On leaves of Lophiraalata sp./Ochnaceae Sierra Leone Conidiophores 110–200 × 3–4 µm thick, simple, bluntly rounded ends; Conidia solitary 15–29 × 3.5–4.5 µm, oblong or oblong-cylindrical, hook or curved, smooth, olive- brown, 1–2 guttulate, 1–3-distoseptate Absent [172] 130 H. lunzinense No information available No information available No information available Absent [173] 131 H. lusitanicum On Alnus glutinosa/ Betulaceae Portugal No information available Absent [174] 132 H. lycopersici On Solanum lycopersicum/Solanaceae Guinea No information available Absent [175] 133 H. machaerii On Machaerium sp./ Fabaceae Brazil No information available Absent [111] UK/Great Britain Conidiophores erect, simple, fusiform, 7–10 septate; Conidia 0.5-0.65 × 0.1 mm Absent [176] Japan Conidiophores 150–270 µm long, 9.5–13 µm thick at the apex, 8.5–13.5 µm thick at the base, single or fasciculate, straight or flexuous, smooth walled, brown to dark brown, sometimes paler toward the apex, septate; Conidia 100–203 × 12.5–22.5 µm tapering gradually to 2.5–5 µm thick near the apex, solitary, obclavate or rostrate, straight or flexuous, pale olive-brown to pale brown, paler toward the apex, 7–18-distoseptate, with a blackish-brown to black scar, 4–7 µm thick Present [177,178] 134 135 H. macilentum H. magnisporum On rotten wood On dead fallen branches of an unknown woody plant Life 2021, 11, 454 37 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 136 H. makilingense On dead branches of Paramignya monophylla/Rutaceae Philippines Conidiophores 400–600 × 7–9 µm, dense, erect, curved, brown, septate; Conidia 100–300 × 10–12 µm, obclavate, 12–18-distoseptate Absent [179] 137 H. manihotis on living leaves of Manihot sp./ Euphorbiaceae Brazil Conidiophores 50–95 × 4–6 µm, 4–6 septate; Conidia 40–50 × 6–8 µm, vermiform, clavate to subfusoid, olives, 4–7-distoseptate Absent [180] 138 H. marantae On leaves of Maranta arundinacea/Marantaceae China/Taiwan No information available Absent [130] 139 H. massarinum Berchemia racemose/ Rhamnaceae Japan Conidiophores 380–810 × 7–9 wide at the apex, 13.5–21 wide at the base µm, 15–25 septate; Conidia 17–56.5 × 5–9 µm, tretic, solitary or in short chains (5–6), obclavate, rostrate, pale brown, smooth, with or without guttules, 1–8-distoseptate Present [19] 140 H. mattiroloi On branches of Sideroxylon oxyacantha/Sapotaceae Etiopia No information available Absent [181] 141 H. mayaguezense On culms and leaves of Paspalum conjugatum/ Poaceae Puerto Rico Conidiophores 300–500 × 18–22 µm; Conidia 135–155 × 35–45 µm, fusoid to clavate, 3–4-distoseptate Absent [182] 142 H. melastomacearum On Meliolamelastomacearum, on Miconiaracemose/ Melastomataceae Puerto Rico Conidiophores 280 × 3 µm; Conidia 14–21 × 3.5–6 µm, ellipsoid, 3-distoseptate Absent [159] 143 H. meliae On leaves of Melia azedarach/Meliaceae Dominican Republic Conidiophores 250–350 × 15–22 µm, simple, aggregated, branched, olive-brown to black, septate; Conidia 70–100 × 12–15 µm, elongate, fusoid, or clavate Absent [183] 144 H. melioloides On leaves of Uvaria sp./Annonaceae Philippines Conidiophores 250–300 × 6–8 µm; Conidia 35–45 × 9–10 µm, obclavate, 3-distoseptate Absent [137] England, Italy Conidiophores 100–550 × 8–14 µm, fasciculate, simple, flexuous, cylindrical, smooth-walled, dark brown, with a pore at the apex and often 1–2, septate; Conidia 60–(114–)160 × 12–(17–)22 µm thick in broadest part, tapering to 4–10 µm near the apex, obclavate, smooth-walled, pale to mid golden-brown, 9–17-distoseptate, with 5–7 µm wide at the scar Present [184] 145 H. microsorum On twigs of Quercus ilex/ Fagaceae Life 2021, 11, 454 38 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 146 H. microsporum From soil India/Maharashtra Conidiophores 234–468 × 10.8 µm, pale brown, 10–16 septate; Conidia 26–41 × 22 µm, fusoid, widest at the middle, brown, 2–7-distoseptate Absent [18] 147 H. minimum On dead decorticatd branches UK/Great Britain/England Conidiophores erect, simple, septate; Conidia 12–14 × 3–4 µm, fusiform, obtuse at the ends, triseptate, scarcely constricted, hyaline Absent [185] Absent [129] 148 H. multiseptatum On dead branches China/Guangdong Conidiophores 390–650 × 10–14 µm wide at the base, 7–9 µm at apex, simple, subcylindrical, straight or slightly flexuous, smooth-walled, brown to dark brown, paler towards the apex, with 1–3 well definded, small pores, 1–3 septate; Conidia 78–190 µm long, 11–16 µm thick in the widest part, narrowing toward the apex to 3–6 µm, tretic, straight or slightly flexuous, obclavate or whip-like, smooth-walled, pale brown paler toward the apex, 13–25-distoseptate, with a dark blackish-brown scar at the base 149 H. nadsonii On fibers of Gossypium sp./Malvaceae Russia No information available Absent [186] Present [187] 150 H. nanjingense On dead branches of an unidentified tree China/Jiangsu, Nanjing Conidiophores 250–470 × 6.9–7.7 µm, solitary or fasciculate, simple, straight or flexuous, thick-walled, sub-cylindrical, smooth, brown to dark brown, with well-defined small pores at the apex,1–4 septate; Conidia 64.5–170.5 µm long, 7.3–10.3 µm wide in the widest part, narrowing towards the apex to 5.0–6.8 µm wide, subulate or nearly whip-like, straight or curved, thin-walled, smooth, pale brown, 6–17-distoseptate, with a blackish-brown scar at the base, 1.4–2.7 µm thick 151 H. naviculare On leaves of Euphorbia sp./ Euphorbiaceae Brazil/Tubarão Conidiophores 6–8 µm thick, branched, often curved, yellow, septate; Conidia 50–84 × 11–16 µmnaviculiform, hyaline at length, very pale with brown Absent [188] 152 H. naviculatum On dead herbaceous stems of Solidago sp./Asteraceae U.S.A./New York No information available Absent [189] 153 H. newbouldiae On leaves of Newbouldialaevis/ Bignoniaceae Guinea No information available Absent [190] Life 2021, 11, 454 39 of 53 Table 5. Cont. No. 154 Taxa H. novae-zelandiae Host(Genus/Family) On dead wood and bark of Vitex lucens/ Lamiaceae Locality Morphology Sequence Data References New Zealand Conidiophores 165 µm long, 4.8–7(–9) µm, erect, single or in groups, simple, or once-branched at the base, straight or flexuous, subcylindrical, brown to dark brown below, very pale brown to subhyaline above, 15 septate; Conidia 13.5–16.2 × 7.2–9.0 µm, solitary, obovoid, sometimes slightly, smooth, the 2 lower cells being brown and the distal cell paler with a dark band of wall overlying each septum, 2-distoseptate Absent [191] Absent [128] 155 H. obpyriforme On dead branches of unidentified tree China/Guangxi Conidiophores 225–460 µm long, 9.5–13 µm diam just above the base and 6–8.5 µm diam towards the apex, arising singly from the upper cells of the stromata, simple, subcylindrical, straight or flexuous, dark brown, paler towards the apex, with well-defined small pores at the apex,1–3 septate; Conidia 47–74 µm long, 14–19 µm diam in the widest part, narrowing in diameter towards the apex to 2.5–5 µm, straight or slightly curved, obpyriform, smooth-walled, middle brown, paler towards the apex, 5–9-distoseptate, with a large dark blackish-brown scar at conidium base, 1–2 µm thick 156 H. ocoteae On Meliola ocoteae, on Guareatrichilioides Puerto Rico Conidiphores 135–200 × 4 µm, septate; Conidia 20–28 × 4–6 µm, 3-distoseptate Absent [159] 157 H. oligosporum Holotype of Sporidesmium olivaceum: on rotten branches of Tilia sp. Lectotype of Coryneumoligosporum, here designated: on rotten branches of Corylus sp. Epitype of Sporidesmiumolivaceum and of Coryneumoligosporum: on dead corticated twigs of Tiliacordata sp. Austria, Czech Republic, Germany From Epitype specimen [21] Conidiophores (17–)22–35(–46) × (8.0–)8.5–10.5(–11.5) µm, densely crowded, erect, simple, straight, cylindrical to slightly swollen at the apex, brown to dark brown, darker at the apex, smooth, 0–2 septate; Conidia (37–)59–80(–124) × (14.8–)15.8–18.0(–20.0) µm, tapering to 4–10.5 µm at the distal end, with 4–8 µm wide, dark brown to black scar at the base, obclavate, sometimes rostrate, straight or curved, smooth but occasionally wrinkled with age, pale brown to brown, paler toward the apex, 6–12(–16)-distoseptate Present [21,124] 158 H. olisipponense Culture from the perithecia stage of Pyrenophora polytricha - No information available Absent [192] Life 2021, 11, 454 40 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 159 H. oplismeni On leaves of Oplismenus cotnpositus/Poaceae China/Taiwan No information available Absent [130] 160 H. orchidacearum On leaves of Neottia ovata (=Listera ovata)/Orchidaceae France No information available Absent [193] 161 H. orthospermum On rotten wood U.S.A./New York Conidiophores 50–60 × 5 µm, erect, simple, fasciculate, straight, dark, 3–4 septate; Conidia 60–80(–110) × 10–12 µm, cylindrical, straight, apex rounded, tuncated at base, 12–14-distoseptate Absent [194] 162 H. oryzae-microsporae On Oryza sativa/Poaceae Japan No information available Absent [195] Absent [128] 163 H. ovoideum On dead branches of tree China/Jilin Conidiophores 380–510 × 15–25 µm diam just above the base, 7.5–10 µm diam towards the apex, arising singly from the upper cells of the stromata, simple, subcylindrical, straight or flexuous, thick-walled, smooth, brown to dark brown, paler towards the apex, with 1–3 well-defined small pores at the apex, 1–6 septate; Conidia 27–61 × 13–21 µm diam in the widest part, narrowing towards the apex to 4.5–8.5 µm, straight, ovoid, to ellipsoidal, smooth-walled, moderately brown, paler towards the apex, 3–8-distoseptate, with a large dark blackish-brown scar at the base, 1.5–2.5 µm thick 164 H. pachystelae On living leaves of Synsepalumm solo(=Pachystelam solo)/ Sapotaceae Tanzania Conidiophores 300–350 × 6–8 µm, erect, simple, septate; Conidia 35–50 × 10–13 µm, fusoid or oblong clavate or lanceolate, 3-5-distoseptate Absent [196] H. palaestinum On stems and flowers of Dianthus sp./ Caryophyllaceae Israel Conidiophores 30–160 × 6–8 µm, fasciculate, 8–16 aggregate, simple, bent, thick-walled, coffin terminal obtuse, thin, yellow or colorless, 5–7 septate; Conidia 60–120 × 9–12 µm, solitary, obclavate, rectiusculis or curved, pale-olive, minute-granule, thick-walled, towards colorless above, 5–7-distoseptate Absent [197] 165 Life 2021, 11, 454 41 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References Brasil/Pará, Papua New Guinea From reference specimen [190]; Conidiophores 132.5–195 × 5–6 µm, solitary, erect, simple, cylindrical, straight or flexuous, smooth, brown, light brown at the apex, 7–10 septate; Conidia 38–53 × 8–11 base, 3–4 µm apex, solitary, in small chains, obclavate or cylindrical, straight or slightly curved, simple, smooth, brown with light brown at apical cell, 6–10-distoseptate Absent [198,199] 166 H. palmigenum On rotten fruit of Cocos nucifera/Arecaceae; On petriole and rachis from reference specimen 167 H. panici On leaves of Panicum maximum/Poaceae Indonesia/Java Conidiophores 115–180 × 8–10 µm; Conidia (35–)50–75 × (7–)10–13 µm, ellipsoidal-truncate, ellipsoidal-elongate, dull-brown, (1–)3(–4)-distoseptate Absent [159] 168 H. papulosum On bark of Malus sylvestris or Pyrus communis/Rosaceae West Virginia No information available Absent [200] 169 H. parathesicola On Meliola parathesicola, on Parathesis serrulate/Primulaceae Puerto Rico Conidiophores 120 × 4 µm, solitary; Conidia 17–20 × 4–6 µm, base truncate, apex beaked, beak often 7 µm long, 1–3-distoseptate Absent [159] 170 H. paulense On leaves of Myrtaceae Brazil/São Paulo Conidiophores 3–4.5 µm thick, brown, septate; Conidia 15–24 × 4 µm, fusoid, brown, 3-distoseptate Absent [115] 171 H. penniseti On leaves of Pennisetum glaucum (=Pennisetum typhoides)/Poaceae India/Uttar Pradesh No information available Absent [108] 172 H. philippinum On decaying leaves of Arenga mindorensis/ Arecaceae Philippines Conidiophores 300–400 × 6–7 µm, fasciculate, filiform, curved, septate; Conidia 33–35 × 8–9 µm, obclavate, 4-distoseptate Absent [137] 173 H. philodendri On Meliola philodendri, on Philodendron krebsii/Araceae Puerto Rico Conidiphores 400 × 3–4 µm; Conidia 24–35 × 5–9 µm, clavate, 3-distoseptate Absent [159] 174 H. phomatae On bark of Acer pennsylvanicum/ Sapindaceae U.S.A./New York No information available Absent [189] 175 H. phyllantheum On dead branches hanging down of Phyllanthus sp./Phyllanthaceae Philippines Conidiophores 180–200 × 4.7–6 µm, fillifrom, blackedned, septate; Conidia 80–90 × 9–10 µm, obclavate, long, 9–11-distoseptate Absent [137] Life 2021, 11, 454 42 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 176 H. piperis On leaves of Piper betle/Piperaceae China/Taiwan No information available Absent [130] 177 H. portoricense On dead branches hanging down of Phyllanthus sp./Phyllanthaceae Philippines Conidiophores 25–250 × 2–5 µm; Conidia 30–60 × 6–10 µm, elongate-fusoid, olive-brown or brown, (2–)4-distoseptate Absent [86,201] 178 H. proliferatum On grain of Triticum sp./Poaceae India/Maharashtra Colony on PDA; Conidiophores 292–510 × 7–13.8 µm, unbranched, pale, olivaceous, 5–20 septate; Conidia 23–126 × 11.5–13.8 µm; cylindrical, olivaceous, 3–13-distoseptate Absent [101] Absent [128] 179 H. pseudomicrosorum On dead branches of unidentified tree China/Changbaishan, Jilin Conidiophores 155–288 × 11–15 µm, fasciculate, simple, cylindrical, straight or flexuous, smooth, dark brown, paler towards the apex, with 1–3 well-defined small, 1–4 septate; Conidia 82–142 × 17–27 µm in the widest part, narrowing towards the apex to 3–6 µm diam, tretic, straight or slightly flexuous, obclavate, smooth-walled, brown, paler towards the apex, 7–16-distoseptate, with a large dark blackish-brown scar at the base, 2–4 µm thick 180 H. pseudotsugae On bark and resin exudations of Pseudotsuga taxifolia var. glauca/Pinaceae U.S.A. Conidiophores scattered on aerial hyphae with usually one at each cell; Conidia 65–105 × 14–15 µm, opaque, black or greenish black, smooth walls, with 8–14-distoseptate Absent [202] 181 H. purpurascens On leaves of Panicum purpurascens/Poaceae U.S.A./Florida No information available Absent [203] 182 H. pyracanthae Pyracantha sp./Rosaceae Portugal No information available Absent [204] U.S.A. Conidiophores (115–)133–226(–300) µm long, 14–20 µm wide at the base, tapering to 10–15 µm near the apex, solitarily or fasciculate, simple, straight or flexuous, cylindrical, thick-walled, smooth, brown to dark brown, with well-defined small pores at the apex; Conidia 60–100 × 15–22 µm, straight or flexuous, obclavate, smooth-walled, brown, 8–10-distoseptate, with blackish-brown to black scar at the base Absent [21] 183 H. quercicola On dead corticated branches of Quercus cf. reticulata/Fagaceae Life 2021, 11, 454 43 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) Locality Morphology Sequence Data References 184 H. quercinum On dead corticated twigs of Quercus petraea/Fagaceae Conidiophores (40–)74–199(–332) µm long, 11–18 µm wide at the base, tapering to 8.5–13.5 µm near the apex, solitarily or fasciculate, simple, straight or flexuous, cylindrical, smooth, Austria/Niederösterreich, brown to dark brown, with well-defined small pores at the apex, Spitzerberg 1–5 septa; Conidia (47–)78–130(–201) × (13.2–)15.3–18.0(–20.5) µm, straight or flexuous, rostrate, smooth-walled, brown, 8–13(–20)-distoseptate, with blackish-brown to black scar at the base 185 H. repens On bark of dead Acer grandidentatum/ Sapindaceae U.S.A./Utah, Red Butte Canyon Conidia 40–45(–60) × 8–9 µm, sub-oblong, 5–12-distosepate Absent [205] 186 H. reyesii On dead branch of Guioa sp./ Sapindaceae Philippines Conidiophores 130 × 8–10 µm, erect, brown, septate; Conidia 34–112 × 8–13 µm, tereti-fusoid, brown, ends hyaline, 5–14-distosepate Absent [137] 187 H. rhodomyrti On leaves of Rhodomyrtus tomentosa/Myrtaceae China/Guangdong Conidia 42–60 × 17–20 µm, fusoid, brown, 5–7-distosepate Absent [206] 188 H. rhopaloides On decraying stem of Brassica oleracea/Brassicaceae Britain, France, Germany, Italy, Portugal Conidiophores short, dark brown-black, 12–14 septate; Conidia 0.04–0.1 mm long, 0.08 mm wide, straight or slightly curved Absent [207,208] 189 H. schelkownikowii On branches Armenia, Azerbaijan, Georgia, Russia No information available Absent [209] 190 H. scolecoides On dry woood Germany/Reichenberg Conidiophores simple, branched; Conidia 80 × 7.5 µm, torulus, fusciculate, septate, yellow Absent [96] 191 H. sechiicola On Sechium edule/ Cucurbitaceae Puerto Rico No information available Absent Present [210] 192 H. sichuanense On dead branches of tree China/Sichuan No information available Absent Present [211] Present [21] Life 2021, 11, 454 44 of 53 Table 5. Cont. No. 193 Taxa H. solani Host(Genus/Family) On stem of Solanum nigrum/Solanaceae (type); Citrus linella; Leucaena glauac; Solanum dulcamara; S. nigrum; Solanum tuberosum Locality Morphology Sequence Data References England, Guernsey, New Zealand, New Guinea, Sierra Leone, Wales Conidiophores 120–600 × 9–15 µm thick near base, 6–9 µm thick near the apex, erect, simple, straight or flexuous, smooth or occasionally, brown to very dark brown, paler near apex, septate, with small pores at apex, 1–8 septate; Conidia (24–)39–85 × (7–)9–11 µm, straight or curved, obclavated, smooth-walled, subhyaline to brown, 2–8-distosepate, with a welll-defind dark brown to black scar at base Present [212] Absent [213] 194 H. solitarium On leaves of Iris sp./Iridaceae U.S.A./Minnesota Conidiophores 60–150 × 6 µm, solitary, slightly fasciculate, erect, swollen at the base, lighter colored at the apex, dark brown, septate; Conidia 24–30 × 8–9 µm, oblong-elliptical, sometimes slightly curved, dark brown, at first 2–4 guttulate, 3–5-distosepate 195 H. spirotrichum On withered leaves of Cyrtophyllum fragrans/Gentianaceae Singapore Conidiophores 190–220 × 6 µm, fasciculate, filiform, brown, septate; Conidia 23–25 × 9 µm, oblong-obclavate, gently curving, brown, 3-distosepate Absent [214] 196 H. spurirostrum On dead branches of tree China/Sichuan No information available Absent [211] 197 H. subapiculatum On dead wood of Sambucus callicarpa/Adoxaceae U.S.A./Washington Conidiophores 8–10 µm thick; Conidia 35–80 × 12–16 µm, oblong or subfusiform, 6–7-distosepate Absent [215] China/Guangdong Conidiophores 120–200 × 10–12 (basal), above 6–8.5 µm thick, simple or fasciculate, erect, subcylindrical, brown, pores 1–3 µm, septate; Conidia 72–125 × 9–11.5 µm, obclavate, straight or flexuous, subhyaline, apex 2.5–5 µm, black at tip, 6–9-distosepate, dark blackish-brown scar Absent [129] China/Yunnan Conidiophores 239–423 × 8.5–15.5 µm, solitary or in group of 2–4, unbranched, straight or curved, smooth, dark brown, paler towards to the apex, bulbous at base 9–14 septate; Conidia 41–55 × 14.5–18.5 µm, straight or curved, wider below than apex, truncate and dark at base, apically rostrate and pale, smooth, pale brown to mid-brown, guttulate, 6–10-distosepate Present [24] 198 199 H. subhyalinum H. submersum On living leaves of Phoenix hanceana/Arecaceae On submerged decaying wood Life 2021, 11, 454 45 of 53 Table 5. Cont. No. 200 Taxa H. subsimile Host(Genus/Family) On withered and dead leaves of Bruguiera sexangula (=Bruguiera eriopetala)/Rhizophoraceae Locality Morphology Sequence Data References Singapore Conidiophores 200–250 × 8–9 µm, filiform-fasciculate, brown, septate; Conidia (38–)45–50 × 11–12(–14) µm, brown, 3-distoseptate Absent [216] Present [22] 201 H. syzygii On bark canker of Syzygium sp./Myrtaceae South Africa/Eastern Cape Province Conidiophores 150–400 × 10–15 mm, fasciculate, unbranched, clavate at apex, dark brown, multiseptate; Conidia (70–)80–100(–150) × (19–)22–23(–25) mm, obclavate, curved, apex subobtuse, warty, inner surface striate, medium brown, (7–)9–12-distoseptate 202 H. theobromae On leaves of Theobroma cacao/Malvaceae Italy Conidiophores 500–1000 µm, erect, 6–10 septate; Conidia 60–160 × 12–20 µm, obclavate to tereti-obclavate Absent [217] 203 H. theobromicola On rotten leaves of Theobroma cacao/Malvaceae Dominican Republic/Moca Conidiophores 20–33 × 3.5–5 µm olives-brown, septate; Conidia 46–58 × 10–13.5 µm, elliptic-oblong or subfusoid, irregular, 3–5-distoseptate Absent [218] 204 H. tritici On seedhead of Triticum vulgare/Poaceae Tanzania Conidiophores 3.5–5 µm, thick fasciculate, erect, sepate; Conidia 12–25 × 4–7 µm, subcylindrical-oblong, clavate or fusoid, 2–4-distoseptate, constrict at septum Absent [219] 205 H. tritikernelis On kernels of Triticum aestivum/Poaceae India/Bihar No information available Absent [108] 206 H. turbinatum On unidentified wood Great Britain Conidiophores simple, slender; Conidia elongated, turbinatis, tuncatus, apiculate, brown, 4–7-distoseptate Absent [220] 207 H. ubangiense On leaves of Coffea sp./Rubiaceae Democratic Republic of the Congo/Ubangi River Conidiophores (2–)3–6 µm, fasciculate, erect, branched, septate; Conidia 30–60 × 5–8 µm, fusoid, 3–4-distoseptate Absent [221] 208 H. ustilaginoideum On flowers of Panicum spicatum/Poaceae Democratic Republic of the Congo Conidiophores 3–3.5 µm thick, fasciculate; Conidia 10–50 × 3.5–4.5 µm, cylindrical or subfusoid, blunted, 1–5-distoseptate Absent [121] Brazil/Pernambuco Conidiophores 150–200 × 10–14 µm, erect, unbranched, straight or flexuous, cylindrical, slightly inflated at the apex, smooth, brown, 5–7 septate; Conidia 29–58 × 4–7 µm, cylindrical-obclavate, subcylindrical, oblong or navicular, dry, trick-walled, with wall verrucose or verruculose, gray-brown, lumina pale yellow, (0–)1–4-distoseptate Absent [222] 209 H. varium On decaying leaves of unidentified plants Life 2021, 11, 454 46 of 53 Table 5. Cont. No. Taxa Host(Genus/Family) H. varroniae On leaves of Varronia sp./Boraginaceae 211 Locality Morphology Sequence Data References Puerto Rico Conidiophores 160–200 × 4 µm; Conidia 27–44 × 6–7 µm, 3-distoseptate Absent [223] H. velutinum Fagus sylvatica dead corticated twigs/saprobic on decaying wood submerged in stream Austria, Wien, Döbling, Kahlenberg/China From reference specimen [20]; Conidiophores 530–655 × 16–18 µm, erect or flexuousk, unbranched, dark brown, 17–23 septate; Conidia 67–79 × 15–19 µm, single, obclavate, straight or curved, smooth, pale brown to brown, 7–9-distoseptate, rounded at apex, guttulate when young, non-guttulate at maturity Present [16,20] 212 H. viticis On leaves of Vitis sp./Vitaceae Brazil/Pará Conidiophores 80 × 2–3 µm, fasciculate, septate; Conidia 12–20 × 2.5–3.5 µm cylindrical, 1–3-distoseptate Absent [223] 213 H. wagateae On leaves of Moullava spicata (=Wagatea spicata)/Fabaceae India/Karnataka Conidiophores 81–125 × 1.5 –2.5 µm, yellowish-brown, multiseptate; Conidia 15.5–28 × 3–4 µm, clavate-cylindric, cinnamon-yellow, rounded at both ends, 2–4-distoseptate Absent [91] 214 H. warpuriae On stem of Warpuria clandestina/Acanthaceae Great Britain/England Conidiophores 300–500 × 6–8 µm; Conidia 115–190 × 12–14 µm, obclavate, 8–11-distoseptate Absent [224] 215 H. xanthosomatis On leaves of Xanthosoma violaceum/Araceae Dominican Republic/Moca Conidiophores 35–90 µm long, septate; Conidia 185 × 24 µm, fusoid, subfusoid to subclavate, 3–7(-10)-distoseptate Absent [225] 216 H. xylopiifolii On Asterina, on Xylopia sericea/Annonaceae Brazil/Pernambuco Conidiophores 85–305 × 5.5–8 µm, erect, 3–5 septate; Conidia 38–62 × 8–13.5 µm, cylindrical or clavate, 3–6-distoseptate Absent [226] 210 Life 2021, 11, 454 47 of 53 Author Contributions: Conceptualization, S.K., K.D.H., S.C.K., C.S. and S.T.; Data curation, S.K.; Methodology, S.K., A.M. and S.T.; Resources, K.D.H., S.C.K., J.X. and S.L.; Supervision, K.D.H. and S.T.; Writing—original draft, S.K., K.D.H., S.C.K., A.M., C.S., L.A.P.D., C.M.N., J.X. and S.T.; Writing—review & editing, S.K., K.D.H., S.C.K., S.T. and S.L. All authors have read and agreed to the published version of the manuscript. Funding: Saowaluck Tibpromma would like to thank the International Postdoctoral Exchange Fellowship Program (number Y9180822S1), CAS President’s International Fellowship Initiative (PIFI) (number 2020PC0009), China Postdoctoral Science Foundation and the Yunnan Human Resources, and Social Security Department Foundation for funding her postdoctoral research. Kevin D. Hyde thanks the Thailand Research Funds for the grant “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Subregion (RDG6130001)”. Samantha Karunarathna thanks CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (number 2018PC0006) and the National Science Foundation of China (NSFC) for funding this research work under project code 31750110478. This work was partly supported by Chiang Mai University. Institutional Review Board Statement: Not applicable. 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