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Research Article
An overview of Melanommataceae (Pleosporales, Dothideomycetes): Current insight into the host associations and geographical distribution with some interesting novel additions from plant litter
expand article infoDanushka S. Tennakoon, Kasun M. Thambugala§, Nimali I. de Silva|, Hai-Yan Song, Nakarin Suwannarach|, Fu-Sheng Chen, Dian-Ming Hu
‡ Jiangxi Agricultural University, Nanchang, China
§ University of Sri Jayewardenepura, Nugegoda, Sri Lanka
| Chiang Mai University, Chiang Mai, Thailand
Open Access

Abstract

Melanommataceous species exhibit high diversity with a cosmopolitan distribution worldwide and show a prominent saprobic lifestyle. In this study, we explored five saprobic species collected from plant litter substrates from terrestrial habitats in China and Thailand. A combination of morphological characteristics and multi-locus phylogenetic analyses was used to determine their taxonomic classifications. Maximum Likelihood and Bayesian Inference analyses of combined LSU, SSU, ITS and tef1-α sequence data were used to clarify the phylogenetic affinities of the species. Byssosphaeria poaceicola and Herpotrichia zingiberacearum are introduced as new species, while three new host records, Bertiella fici, By. siamensis and Melanomma populicola are also reported from litter of Cinnamomum verum, Citrus trifoliata and Fagus sylvatica, respectively. Yet, despite the rising interest in the melanommataceous species, there is a considerable gap in knowledge on their host associations and geographical distributions. Consequently, we compiled the host-species associations and geographical distributions of all the so far known melanommataceous species.

Key words

Biodiversity, multi-gene phylogeny, new host records, Pleosporales, saprobes, systematics

Introduction

Fungi occur in a wide range of ecosystems (Hawksworth 2001; De Silva et al. 2016; Hernández-Restrepo et al. 2017). With over 98,334 extant species, Ascomycota is the largest phylum of fungi and is widely distributed in terrestrial, freshwater and marine environments (Hill et al. 2021; Senanayake et al. 2022; Bánki et al. 2023). The class Dothideomycetes is estimated to have 32,365 species and is one of the most ecologically diverse group of ascomycetes (Hongsanan et al. 2020; Bánki et al. 2023). Pleosporales is considered the largest and most diverse order in the class Dothideomycetes (Zhang et al. 2012; Hongsanan et al. 2020). They exhibit a wide range of lifestyles (e.g. biotrophs, endophytes, epiphytes, hemibiotrophs, pathogens, saprobes) and can be found worldwide, including terrestrial, marine and freshwater environments (Jones et al. 2019; Hongsanan et al. 2020; Calabon et al. 2022; Gao et al. 2023). In addition, they are extremely adaptable to various ecological niches and can exist in anaerobic, aquatic, mutualistic, terrestrial and even in severe habitats, such as deserts (Zhang et al. 2012; Hyde et al. 2013; Hongsanan et al. 2020). Pleosporales species play significant functional roles from an agricultural, ecological and economic perspectives (Zhang et al. 2012; Raja et al. 2017; Wen et al. 2020; Pimenta et al. 2021). According to the recent outline of Wijayawardene et al. (2022), Pleosporales consists of 91 families.

Melanommataceae is one of the species-rich families in Pleosporales, Dothideomycetes. It was introduced by Winter (1885) to include Melanomma as the type genus and species have globose or depressed ascomata, fissitunicate asci, pigmented and phragmosporous ascospores (Tian et al. 2015; Hongsanan et al. 2020). The ordinal level placement of Melanommataceae was controversial for a long time and Barr (1983) introduced Melanommatales to accommodate all taxa which have trabeculate pseudoparaphyses. The nature of trabeculate pseudoparaphyses was broadly discussed by Liew et al. (2000) and illustrated that they are generally 1 μm or thinner, clearly anastomose between the asci and are embedded in a gelatinous matrix. Subsequently, Barr (1990) introduced five genera in Melanommataceae (Byssosphaeria, Keissleriella, Melanomma, Ostropella and Strickeria) based on erumpent to superficial ascomata and thick-walled peridium. Over time, Melanommataceae has been transferred from Melanommatales to Pleosporales with the revolution of DNA-based molecular phylogenetic studies (Eriksson 2006; Wang et al. 2007; Zhang et al. 2012; Hyde et al. 2013; Tian et al. 2015; Hongsanan et al. 2020).

The species of Melanommataceae have cosmopolitan distribution worldwide in temperate, subtropical and tropical regions (Hyde et al. 2013; Tian et al. 2015; Jaklitsch and Voglmayr 2017; Kularathnage et al. 2022). They can play a vital role as saprobes, endophytes or hyperparasites and occur on twigs or bark of various woody plants in terrestrial, marine or freshwater habitats (Tian et al. 2015; Hashimoto et al. 2017; Tennakoon et al. 2018; Hongsanan et al. 2020; Gao et al. 2023). In addition, some species (e.g. Seifertia alpina) have been recorded as plant pathogens and cause bud blight or bud blast disease of Rhododendron species (Glawe and Hummel 2006; Li et al. 2016a). Interestingly, several species have also been reported from soil (e.g. Herpotrichia gelasinosporoides, H. striatispora and Pleotrichocladium opacum), on lichen species (e.g. Aposphaeria ramalinae) and on mushroom species (e.g. Exosporiella fungorum on Thelephora sp.) (Karsten 1892; Pitard and Harmand 1911).

The cosmopolitan nature of Melanommataceae is further supported by the numerous novel genera and species that have been introduced in the past years. Based on the year of introduction, we compiled the data and revealed that nine genera were introduced between 1800 and 1899 and nine more genera between 1900 and 1999. In addition, 18 genera were introduced between 2000 and 2024 (Fig. 1). This rapid increase may be primarily due to the revolution in fungal taxonomical studies with DNA sequence-based molecular phylogenies in the last two decades. Interestingly, six genera were introduced in the year 2018, such as Marjia, Melanocucurbitaria, Melanodiplodia, Monoseptella, Pseudobyssosphaeria and Uzbekistanica (Fig. 2). Consequently, 36 genera are currently accepted in Melanommataceae, viz. Alpinaria, Aposphaeria, Asymmetricospora, Bertiella, Bicrouania, Byssosphaeria, Calyptronectria, Camposporium, Dematiomelanomma, Exosporiella, Fusiconidium, Herpotrichia, Mamillisphaeria, Marjia, Melanocamarosporioides, Melanocamarosporium, Melanocucurbitaria, Melanodiplodia, Melanomma, Monoseptella, Muriformistrickeria, Navicella, Neobyssosphaeria, Petrakia, Phragmocephala, Phragmotrichum, Pleotrichocladium, Praetumpfia, Pseudobyssosphaeria, Pseudodidymella, Pseudostrickeria, Sarimanas, Seifertia, Tumularia, Uzbekistanica and Xenostigmina (Hongsanan et al. 2020; Gao et al. 2023). Conversely, the species discoveries are much higher during the 1900 and 1999 period (226 species) and thirty-nine species have been introduced between 1800 and 1899. However, despite having introduced 18 genera during the 2000–2024 period, only 76 species have been introduced (Fig. 1). Nevertheless, it could be much more in future with the extensive taxon samplings, particularly in poorly studied countries/regions, substrates and hosts.

Figure 1. 

a The number of melanommataceous genera introduced in different time periods b the number of melanommataceous species introduced in different time periods (Source – MycoBank Database).

Figure 2. 

The melanommataceous genera introduced in different time periods (Source – MycoBank Database).

We are exploring the fungal diversity of plant litter substrates with the aim of clarifying their taxonomy, based on morphology coupled with multi-gene phylogeny (Thambugala et al. 2017; Tennakoon et al. 2018, 2021, 2023; Wanasinghe et al. 2018). Thus, we have collected five taxa from China and Thailand which belong to the family Melanommataceae. The objectives of this study are to identify the melanommataceous taxa associated with plant litter using both morphological and phylogenetic approaches and to provide an updated checklist of species in Melanommataceae. This study provides a database on melanommataceous species for future studies, increases knowledge of fungal diversity and helps to understand their global distribution and host associations.

Materials and methods

Sample collection and examination

Fresh fungal specimens were collected from plant litter (dead wood and leaves) from Chiang Mai, Thailand and Kunming, China. The collected specimens were taken back to the laboratory in zip lock bags and paper envelopes. All the samples were subjected for an incubation period (one day) in plastic boxes lined with wet tissue paper. The micro- and macro-morphological characteristics were observed as described by Tennakoon et al. (2023). Sections of ascomata were taken manually and were mounted in distilled water. A stereomicroscope (AXIOSKOP 2 PLUS Series, Göttingen, Germany) was used to examine the surface morphological characteristics of fungal fruiting bodies. Micro-morphological characteristics, such as asci, ascospores and pseudoparaphyses were examined using Axioskop 2 Plus (Göttingen, Germany) compound microscope. Images were taken using Canon Axiocam 506 color digital camera (Hanover, Germany) fitted to a Axioskop 2 Plus (Göttingen, Germany) compound microscope. The micro morphological characteristics, such as colour, shape, height and diameter of ascomata, asci, ascospores, peridium and pseudoparaphyses were recorded. Indian ink was used to inspect the existence of the mucilaginous sheath in ascospores. The prepared slides were permanently preserved using lactoglycerol and sealed by applying nail-polish around the margins of cover slips. All measurements were obtained using Tarosoft (R) Image Framework application. Adobe Photoshop CS3 Extended version 10.0 software (Adobe Systems, USA) was used to construct the photo plates. Specimens were deposited in the Herbarium of Department of Biology (CMUB) and Sustainable Development of Biological Resources Laboratory (SDBR), Faculty of Science, Chiang Mai University and Herbarium of Fungi, Jiangxi Agricultural University (HFJAU), Nanchang, China. The Faces of Fungi (FOF) and Index Fungorum (IF) numbers were obtained for new species (Byssosphaeria poaceicola and Herpotrichia zingiberacearum) as mentioned in Jayasiri et al. (2015) and Index Fungorum (2024).

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from fungal fruiting bodies on the natural substrate by using a DNA extraction kit (E.Z.N.A. ® Forensic DNA Kit, D3591-01, Omega BIO-TEK) following the manufacturer’s protocol. DNA products were intended for use as a template for PCR and stored at 4 °C and the duplicates were kept at -20 °C for long-term storage. Four genomic regions were amplified, the internal transcribed spacer (ITS) region (ITS1-5.8S-ITS2), the 28S large subunit rDNA (LSU), 18S small subunit rDNA (SSU) and the translation elongation factor 1-alpha gene (tef1-α). The primers ITS4 and ITS5 were used to amplify the ITS (White et al. 1990), LR0R and LR5 primers for LSU (Vilgalys and Hester 1990), NS1 and NS4 for SSU (White et al. 1990) and EF1-983F and EF1-2218R primers for tef1-α (Rehner 2001). The amplification reactions were performed in a total reaction volume of 25 µl, which contained 9.5 µl of sterilised distilled water, 12.5 µl of 2 × Power Taq PCR MasterMix (a premix and ready to use solution, including 0.1 Units/μl Taq DNA Polymerase, 500 μM dNTP Mixture each (dATP, dCTP, dGTP, dTTP) (Bioteke Co., China), 1 μl of each forward and reverse primers (stock concentration 10 pM) and 1 μl of DNA template. The polymerase chain reaction (PCR) thermal cycle programmes for ITS, LSU, SSU and tef1-α genes amplification were adjusted as described in Tennakoon et al. (2022). To check the quality of PCR products, agarose gel electrophoresis (1%) was conducted. The purified PCR products were subjected for sequencing at Sangon Biotech (Shanghai) Co., Ltd, China. Generated sequences were deposited in GenBank and accession numbers were listed (Table 1).

Table 1.

GenBank and culture collection accession numbers of species included in the phylogenetic study. The newly-generated sequences are shown in bold face.

Fungal Species Strain/Voucher No. GenBank Accession Number
ITS LSU SSU tef–1α
Alpinaria rhododendri KT 2520 LC203335 LC203360 LC203314 LC203388
Aposphaeria corallinolutea MFLU 15-2752 KY554202 KY554197 KY554200 KY554205
A. corallinolutea GLMC 1355 MT153708 MT156159
Bertiella ellipsoidea MFLU 16-0583 KX765261 KX765262
B. ellipsoidea MFLUCC 17-2015 MG543922 MG543913
B. fici MFLUCC 20-0229 MW063223 MW079351 MW183786
B. fici NCYUCC 19-0260 MW063224 MW079352 MW183787
B. fici NCYUCC 19-0290 MW063225 MW079353 MW183788
B. fici CMUB 40045 PP460772 PP460764 PP475453
B. macrospora IL 5005 GU385150
B. macrospora SMH 3953 GU327744
Beverwykella pulmonaria CBS 283.53 KY189974 GU301804
Byssosphaeria diffusa AFTOL ID 1588 DQ678071 DQ678019 DQ677915
By. diffusa CBS 250.62 GU205239
By. jamaicana SMH 1403 GU385152 GU327746
By. jamaicana SMH 3085 GU385154
By. jamaicana SMH 3464 GU385153
By. macarangae MFLUCC 17-2655 MH389782 MH389778 MH389780 MH389784
By. musae MFLUCC 11-0146 KP744435 KP744477 KP753947 MH581149
By. phoenicis ZHKUCC 21-0122 ON180685 ON180683 ON180691 ON243583
By. phoenicis ZHKUCC 21-0123 ON180686 ON180684 ON180692 ON243584
By. rhodomphala SMH3086 GU385155
By. rhodomphala ANM 942 GU385160
By. rhodomphala SMH 3402 GU385170
By. rhodomphala GKM L153N GU385157 GU327747
By. salebrosa SMH 2387 GU385162 GU327748
By. schiedermayeriana SMH 1816 GU385159
By. schiedermayeriana SMH 1269 GU385158
By. schiedermayeriana SMH 3157 GU385163 GU327745
By. siamensis MFLUCC 10-0099 KT289895 KT289897
By. siamensis MFLUCC 17-1800 MG543923 MG543914 MG543917
By. siamensis HFJAU10336 PP460780 PP460773 PP460765 PP475454
By. taiwanense MFLUCC 17-2643 MH389783 MH389779 MH389781 MH389785
By. villosa GKM 204 N GU385151 GU327751
By. poaceicola HFJAU10337 PP460781 PP460774 PP460766 PP475455
By. poaceicola HFJAU10338 PP460782 PP460775 PP460767 PP475456
Fusiconidium aquaticum KUMCC 15-0300 KX641894 KX641895 KX641896
F. mackenziei MFLUCC 14-0434 KX611112 KX611114 KX611118
Herpotrichia herpotrichoides GKM 212N GU385169
H. herpotrichoides SMH 5167 GU385175
H. macrotricha GKM 196N GU385176 GU327755
H. macrotricha SMH 269 GU385177
H. macrotricha SMH 269 GU385177 GU327756
H. vaginatispora MFLUCC 13-0865 KT934252 KT934256 KT934260
H. xiaokongense KUMCC 21-0004 MZ408889 MZ408891 MZ394066
H. zingiberacearum HFJAU10332 PP460783 PP460776 PP460768 PP475457
H. zingiberacearum HFJAU10333 PP460784 PP460777 PP460769 PP475458
H. zingiberacearum HFJAU10334 PP460785 PP460778 PP460770 PP475459
Hysterium angustatum MFLU 16-1179 KX611363 KX611364 KX611365
Marjia tianschanica TASM 6120 MG828909 MG829019 MG829126 MG829206
M. tianschanica TASM 6121 MG828910 MG829020 MG829127 MG829206
Marjia uzbekistanica TASM 6122 MG828911 MG829021 MG829128 MG829208
Melanocucurbitaria uzbekistanica MFLUCC 17-0829 MG828912 MG829022 MG829129 MG829209
Melanodiplodia tianschanica TASM 6111 MG828914 MG829023 MG829130 MG829210
Me. tianschanica TASM 6112 MG828915 MG829024 MG829131 MG829211
Me. tianschanica MFLUCC 17-0805 MG828913 MG829025 MG829132 MG829212
Melanomma japonicum KT 3425 LC203320 LC203338 LC203292 LC203367
Mel. populicola CBS 543.70 NR_170056 NG_075164 NG_070237
Mel. populicola CPC 27203 MT223817 MT223910
Mel. populicola CBS 350 82 MT223815 JF740265
Mel. populicola CBS 130330 JF740328
Mel. populicola HFJAU10335 PP460786 PP460779 PP460771 PP475460
Mel. pulvis-pyrius KT 2110 LC203322 LC203340 LC203294 LC203368
Mel. pulvis-pyrius KT 2113 LC203323 LC203341 LC203295 LC203369
Mel. pulvis-pyrius AH 375 LC203324 LC203342 LC203296 LC203370
Mel. pulvis-pyrius KH 27 LC203325 LC203343 LC203297 LC203371
Mel. pulvis-pyrius CBS 124080 MH863349 GU456323 GU456302 GU456265
Monoseptella rosae MFLUCC 17-0815 MG828916 MG829026 MG829133 MG829213
Mo. rosae TASM 6114 MG828917 MG829027 MG829134 MG829214
Mo. tuberculata CBS 256.84 GU301851 GU349006
Muriformistrickeria rubi MFLUCC 15-0681 KT934253 KT934257 KT934261
Petrakia irregularis CBS 306.67 NR_164281 MH870670
Phragmocephala atra MFLUCC 15-0021 KP698721 KP698725 KP698729
P. garethjonesii MFLUCC 15-0018 KP698722 KP698726 KP698730
Pleotrichocladium opacum CBS 450.70 MH859791 KY853524
Pl. opacum CBS 709.92 KY853464 KY853526
Pseudostrickeria rosae MFLUCC 17-0643 MG828954 MG829065 MG829169 MG829234
Ps. mutabilis SMH 1541 GU385209
Sarimanas pseudofluviatile KT 760 LC001717 LC001714 LC001711
S. shirakamiense KT 3000 LC001715 LC001712
Seifertia azaleae DAOM 239136 EU030276
Se. shangrilaensis MFLUCC 16-0238 KU954100 KU954101 KU954102
Uzbekistanica rosae-hissaricae MFLUCC 17-0819 MG828976 MG829087 MG829187 MG829242
U. rosae-hissaricae MFLUCC 17-0820 GU269840 MG829088 MG829188 MG829243
Xenostigmina zilleri CBS 115685 GU269840 GU253857 LC203316 GU384553

Phylogenetic analyses

The obtained sequences were initially assembled (forward and reverse sequences) using SeqMan v. 7.0.0 (DNASTAR, Madison, WI). Assembled sequences were subjected to BLAST search in GenBank to obtain strains which have high similarities (https://blast.ncbi.nlm.nih.gov/). In addition, some other sequences for taxa in Melanommataceae were obtained using recent publications (Kularathnage et al. 2022; Gao et al. 2023). In total, 85 isolates were used for phylogenetic analyses including Hysterium angustatum (MFLU 16-1179) as the outgroup taxon. The combined dataset comprised four genes, ITS, LSU, SSU and tef1-α. Single gene sequences were aligned with MAFFT v.7.490 online application (Katoh and Standley 2013) (https://mafft.cbrc.jp/alignment/software/) and manually improved in necessary places. Aligned sequences were combined using BioEdit v.7.2.5 (Hall 1999).

The concatenated aligned dataset was analysed separately using Maximum Likelihood (ML) and Bayesian Inference (BI). Maximum Likelihood analysis was performed using the online portal CIPRES Science Gateway v. 3.3 (Miller et al. 2010), with RAxML-HPC v.8 on XSEDE (8.2.12) tool (Stamatakis et al. 2008; Stamatakis 2014) using the GTR+I+G model of nucleotide evolution. Evolutionary models for each barcode were determined using MrModelTest v. 2.3 (Nylander 2004) under the Akaike Information Criterion (AIC).

MrBayes 3.2.1 (Ronquist et al. 2012) was used to analyse Bayesian Inference phylogenies and was run with four chains of 3,000,000 generations and trees were sampled every 100th generation. The initial 20% of sampled data were discarded as burn-in. The phylograms were visualised using the FigTree v.1.4.0 tool (Rambaut 2012) and reorganised in Adobe Illustrator® CS3 (Version 15.0.0, Adobe®, San Jose, CA). The alignments and sequences were deposited in TreeBASE, submission ID: 31212 (http://www.treebase.org/) and GenBank (https://www.ncbi.nlm.nih.gov/), respectively.

Geographical distribution and host associations of melanommataceous species

We enumerated 340 species of Melanommataceae, grouped into 36 genera, along with their geographic distribution and host associations. The necessary data were obtained from published books, publications in reputable journals, Species Fungorum (https://www.speciesfungorum.org), MycoBank (https://www.mycobank.org/), the U.S. National Fungus Collections Fungus-Host Database (Farr and Rossman 2024) and online sources (Catalogue of Life Checklist). The gathered data were mentioned with appropriate references (Table 2). In total, 296 articles were studied for information on the 340 melanommataceous species that are currently legitimate (Table 2) and accessed through Google Scholar searches. MycoBank (https://www.mycobank.org/) was used to illustrate the nomenclature validity of the taxa. However, numerous species have not been verified using molecular data, so most species should be correctly identified, based on modern taxonomic concepts. Therefore, the distribution may vary slightly.

Table 2.

Host association and geographical distribution of reported melanommataceous species.

Species Host Host family Locality References
Alpinaria rhododendri Rhododendron sp. Ericaceae Austria and Japan Hashimoto et al. (2017); Jaklitsch and Voglmayr (2017)
Aposphaeria allantella Solanum tuberosum Solanaceae Germany Farr and Rossman (2024)
Aposphaeria anomala Unidentified host Italy Farr and Rossman (2024)
Aposphaeria arachidis Arachis hypogaea Fabaceae India Kulkarni (1974); Mathur (1979); Farr and Rossman (2024)
Aposphaeria bambusae Bambusa sp. Poaceae Brazil Bresadola (1926); Farr and Rossman (2024)
Aposphaeria bombacis Bombax macrocarpum Malvaceae Germany Diedicke (1912)
Aposphaeria brunneotincta Castanea vesca Fagaceae The United States Thaxter (1922)
Aposphaeria buddlejae Buddleja davidii Buddlejaceae Ukraine Farr and Rossman (2024)
Aposphaeria calligoni Calligonum aphyllum Polygonaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria canavaliae Canavalia sp. Fabaceae Fiji Massee (1906); Dingley et al. (1981); Farr and Rossman (2024)
Aposphaeria caraganae Caragana arborescens Fabaceae Russia Farr and Rossman (2024)
Aposphaeria caricicola Carex rossii Cyperaceae The United States Farr and Rossman (2024)
Aposphaeria caulina Cerefolium sylvestre Apiaceae Finland Karsten (1905)
Aposphaeria charticola The United States Saccardo (1911)
Aposphaeria cladoniae Cladonia fimbriata Cladoniaceae Germany Allescher (1896)
Aposphaeria conica Quercus sp. Fagaceae Italy Saccardo and Traverso (1910)
Aposphaeria corallinolutea Fraxinus excelsior and Kerria japonica Oleaceae and Rosaceae The Netherlands De Gruyter et al. (2013)
Aposphaeria dendrophomoides Corylus avellana Betulaceae Italy Saccardo (1921)
Aposphaeria denudata Cydonia vulgaris Rosaceae Hungary Saccardo and Traverso (1910)
Aposphaeria desertorum Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria elymi Elymus arenarius Poaceae Germany Diedicke (1912)
Aposphaeria ephedrae Ephedra sp. Ephedraceae Kazakhstan and Ukraine Hayova and Minter (2009); Farr and Rossman (2024)
Aposphaeria epicorticalis Corylus avellana Betulaceae Italy Saccardo (1921)
Aposphaeria eragrostidis Eragrostis sp. Poaceae Eritrea, Ethiopia and Iraq Castellani and Ciferri (1950); Farr and Rossman (2024)
Aposphaeria eurotiae Eurotia eversmanniana Amaranthaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria ferrum-equinum Unidentified host Italy Tassi (1900)
Aposphaeria freticola Fagus sp. and Nothofagus antarctica Fagaceae, Nothofagaceae Argentina, Chile and Poland Farr (1973); Mulenko et al. (2008); Farr and Rossman (2024)
Aposphaeria gallicola Unidentified host Italy Farr and Rossman (2024)
Aposphaeria gregaria Salix sp. Salicaceae Germany Diedicke (1912)
Aposphaeria halimodendri Halimodendron halodendron Fabaceae Ukraine Farr and Rossman (2024)
Aposphaeria haloxyli Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria hapalophragmii Hapalophragmium acaciae Fabaceae Somalia Trotter (1931); Mujica and Vergara (1945); Farr and Rossman (2024)
Aposphaeria henryana Salix alba Salicaceae Italy Farr and Rossman (2024)
Aposphaeria heveae Hevea brasiliensis Euphorbiaceae Sri Lanka Petch (1917)
Aposphaeria hippuridis Hippuris vulgaris Plantaginaceae Germany Ade (1923)
Aposphaeria hospitae Kleinhovia sp. Malvaceae Sri Lanka Tassi (1900)
Aposphaeria humicola Unidentified host The Netherlands Oudemans (1902)
Aposphaeria ilicis Ilex aquifolium Aquifoliaceae Germany Diedicke (1912)
Aposphaeria iliensis Halimodendron halodendron Fabaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria jubaeae Jubaea spectabilis Arecaceae Chile Spegazzini (1921); Farr (1973); Mujica and Vergara (1945); Farr and Rossman (2024)
Aposphaeria kiefferiana Quercus sp. Fagaceae Italy Farr and Rossman (2024)
Aposphaeria kravtzevii Eurotia eversmanniana Amaranthaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria lentisci Pistacia sp. Anacardiaceae Greece and Spain Urries (1957); Pantidou (1973); Farr and Rossman (2024)
Aposphaeria lignicola Acacia arabica Fabaceae Pakistan Ahmad (1964)
Aposphaeria major Rubus parviflorus Rosaceae The United States Sydow and Sydow (1907)
Aposphaeria majuscula Vitis vinifera Vitaceae France Saccardo and Trotter (1913)
Aposphaeria martinii Sabal sp. Arecaceae United States Farr and Rossman (2024)
Aposphaeria mediella Pinus sp. Pinaceae Greece, Finland and Poland Karsten (1884); Pantidou (1973); Mulenko et al. (2008); Farr and Rossman (2024)
Aposphaeria melaleucae Melaleuca leucadendra Myrtaceae Australia Hennings (1903); Shivas and Alcorn (1996); Farr and Rossman (2024)
Aposphaeria mesembryanthemi Mesembryanthemum sp. Aizoaceae Portugal Costa and Camara (1952)
Aposphaeria mojunkumica Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Aposphaeria montbretiae Crocosmia sp. Iridaceae Azerbaijan and Georgia Siemaszko (1923)
Aposphaeria musarum Musa sapientum Musaceae Argentina Farr (1973); Farr and Rossman (2024)
Aposphaeria nigra Betula alba Betulaceae Germany Diedicke (1912)
Aposphaeria oxalidis Oxalis tuberosa Oxalidaceae Bolivia Farr and Stevenson (1963); Farr and Rossman (2024)
Aposphaeria pakistanica Unidentified host Pakistan Ahmad (1971)
Aposphaeria phellodendri Phellodendron amurense Rutaceae Ukraine Farr and Rossman (2024)
Aposphaeria pinea Pinus sylvestris Pinaceae France and Germany Saccardo (1884); Mulenko et al. (2008); Farr and Rossman (2024)
Aposphaeria pini-densiflorae Pinus densiflora Pinaceae Japan Sawada (1950); Farr and Rossman (2024)
Aposphaeria polonica Tilia platyphyllos Pinaceae Poland Mulenko et al. (2008); Farr and Rossman (2024)
Aposphaeria populea Populus sp. Salicaceae The United Kingdom Smith and Ramsbottom (1914); Dennis (1986)
Aposphaeria pulviscula Fagus sylvatica and Salix sp. Fagaceae and Salicaceae Austria, France, Iceland, Italy, the Netherlands and Ukraine Saccardo (1880); Spaulding (1961); Sutton (1980); Farr and Rossman (2024)
Aposphaeria punicina Punica granatum Lythraceae China and Malta Saccardo (1915); Tai (1979)
Aposphaeria purpurascens Acer pseudoplatanus Sapindaceae Italy Peyronel (1915)
Aposphaeria ramalinae Ramalina sp. (lichen) France Pitard and Harmand (1911)
Aposphaeria reaumuriae Reaumuria sp. Tamaricaceae Azerbaijan Farr and Rossman (2024)
Aposphaeria rhois Rhus oxyacantha Anacardiaceae Libya El-Buni and Rattan (1981); Trotter (1912)
Aposphaeria rostrata Unidentified host The Netherlands Oudemans (1902)
Aposphaeria rubefaciens Salix sp. Salicaceae Italy and the Netherlands Bubák and Kabát (1905)
Aposphaeria rudis Picea excelsa Pinaceae Finland Karsten (1905)
Aposphaeria salicis Salix sp. Salicaceae Germany and India Diedicke (1912); Mathur (1979)
Aposphaeria salicum Salix viminalis Salicaceae Germany Sydow and Sydow (1903)
Aposphaeria santolinae Santolina chamaecyparissus Asteraceae Ukraine Farr and Rossman (2024)
Aposphaeria sepulta Citrus aurantium Rutaceae Italy Saccardo (1884)
Aposphaeria sequoiae Sequoia sp. Cupressaceae Denmark Lind (1913)
Aposphaeria silenes Silene otites Caryophyllaceae Russia Farr and Rossman (2024)
Aposphaeria sphaerospora Betula alba Betulaceae Italy Peyronel (1918)
Aposphaeria striolata Populus deltoides Salicaceae The United States Saccardo (1916)
Aposphaeria taquarae Bambusoideae sp. Poaceae Brazil Viégas (1945)
Aposphaeria tiliana Tilia cordata Malvaceae Ukraine Gucevic (1977)
Aposphaeria tragopogonis Tragopogon dubius Asteraceae Romania Sandu-Ville and Mititiuc (1971)
Aposphaeria turmalis Diospyros virginiana Ebenaceae The United States Ellis and Everhart (1902); Cash (1952)
Aposphaeria ulmicola Ulmus sp. Ulmaceae The United Kingdom Saccardo (1884)
Aposphaeria zeae Zea mays Poaceae Azerbaijan and Georgia Farr and Rossman (2024)
Asymmetricospora calamicola Calamus caryotoides Arecaceae Australia Fröhlich and Hyde (1998); Zhang et al. (2012)
Bertiella botryosa Ulmus sp. Ulmaceae The United States Morgan (1904)
Bertiella ellipsoidea Unidentified host Thailand Hyde et al. (2016)
Bertiella fici Cinnamomum verum, Ficus septica Lauraceae, Moraceae China and Thailand Tennakoon et al. (2021); this study
Bertiella gelatinosa Unidentified host Brazil Almeida et al. (2017)
Bertiella rhodospila Cyrilla sp., Populus sp. and Quercus sp. Cyrillaceae, Fagaceae and Salicaea The United States Barr et al. (1986)
Bertiella striatispora Unidentified host India Niranjan and Sarma (2019)
Bicrouania maritima Atraphaxis spinosa and Halimione portulacoides Amaranthaceae, Polygonaceae France and Uzbekistan Kohlmeyer and Volkmann-Kohlmeyer (1990); Gafforov (2017)
Byssosphaeria alnea Alnus sp. Betulaceae The United States Barr (1984); Farr and Rossman (2024)
Byssosphaeria erumpens Litsea sp. Lauraceae China Chen and Hsieh (2004); Li and Zhuang (2008); Farr and Rossman (2024)
Byssosphaeria erythrinae Erythrina indica Fabaceae France Barr (1984)
Byssosphaeria guangdongense Phoenix roebelenii Arecaceae China Xiong et al. (2023)
Byssosphaeria hainanensis Unidentified host China Li and Zhuang (2008)
Byssosphaeria jamaicana Bambusa sp., Quercus sp. and Salix sp. Fagaceae and Poaceae, Salicaceae China, Czech Republic, Jamaica and Mexico Barr (1984); Sivanesan and Hsieh (1989); Wang et al. (2004); Medel (2007); Farr and Rossman (2024)
Byssosphaeria juniperi Juniperus sp. Cupressaceae The United States Wang et al. (2004); Farr and Rossman (2024)
Byssosphaeria macarangae Macaranga tanarius Euphorbiaceae China Tennakoon et al. (2018)
Byssosphaeria musae Musa sp. Musaceae Thailand Liu et al. (2015); Farr and Rossman (2024)
Byssosphaeria oviformis Saccharum arundinaceum Poaceae China and Jamaica Barr (1984); Lu et al. (2000); Wong and Hyde (2001); Farr and Rossman (2024)
Byssosphaeria phoenicis Phoenix roebelenii Arecaceae China Kularathnage et al. (2022)
Byssosphaeria poaceicola Arundo pliniana Poaceae China This study
Byssosphaeria rhodomphala Acer pseudoplatanus and Populus sp. Salicaceae and Sapindaceae Brazil, China, Poland and the United States Cooke (1887); Scheuer and Chlebicki (1997); Reblova (1997); Chen and Hsieh (2004); Wang et al. (2004)
Byssosphaeria salebrosa Vaccinium sp. Ericaceae The United States Barr (1984); Barr et al. (1986); Farr and Rossman (2024)
Byssosphaeria schiedermayriana Sambucus nigra Adoxaceae Austria Barr (1984)
Byssosphaeria semen Pyrus americana Rosaceae The United States Barr (1984)
Byssosphaeria siamensis Pandanus sp. and Citrus trifoliata Pandanaceae and Rutaceae China and Thailand Tian et al. (2015); Hyde et al. (2018); Farr and Rossman (2024); this study
Byssosphaeria taiwanense Macaranga tanarius Euphorbiaceae China Tennakoon et al. (2018)
Byssosphaeria xestothele Cornus florida and Robinia pseudoacacia Cornaceae and Fabaceae Sweden and the United States Barr (1984); Eriksson (2014); Farr and Rossman (2024)
Calyptronectria argentinensis Foeniculum piperitum and Manihot carthaginensis Apiaceae, Euphorbiaceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Calyptronectria indica Annona squamosa Annonaceae India Pande (2008)
Calyptronectria platensis Manihot carthagenensis Euphorbiaceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Camposporium antennatum Acacia aulacocarpa, Caesalpinia echinata, Cinnamomum japonicum, Cocos nucifera, Drymophloeus pachycladus, Eucalyptus globulus, Ficus erecta, Laurus nobilis, Machilus sp., Mucuna ferruginea, Neolitsea scrobiculata, Phoenix hanceana, Pinus massoniana, Quercus sp. and Trachycarpus fortunei Arecaceae, Fabaceae, Fagaceae, Lauraceae, Moraceae, Myrtaceae, and Pinaceae California, China, the United Kingdom and Venezuela Harkness (1884); Lu et al. (2000); Castaneda-Ruiz et al. (2003); Farr and Rossman (2024)
Camposporium appendiculatum Unidentified host China Hyde et al. (2020)
Camposporium atypicum Mesua ferrea Calophyllaceae India Koukol and Delgado (2021)
Camposporium cambrense Alnus sp., Carpinus betulus, Fagus sp., Freycinetia anksia, Laurus nobilis and Quercus sp. Betulaceae, Fagaceae, Lauraceae, and Pandanaceae China, New Zealand, Poland, Russia and the United Kingdom Hughes (1951); Whitton et al. (2002); Mulenko et al. (2008); Farr and Rossman (2024)
Camposporium chinense Unidentified host China Xu et al. (2021)
Camposporium dulciaquae Unidentified host Thailand Calabon et al. (2021)
Camposporium fusisporum Pandanus sp. Pandanaceae Brunei Whitton et al. (2002)
Camposporium himalayanum Phoenix sp. Arecaceae India Adamcik et al. (2015)
Camposporium hyalinum Elaeagnus macrophylla, and Fagus sp. Elaeagnaceae and Fagaceae The United Kingdom Abdullah (1980); Kirk and Spooner (1984)
Camposporium hyderabadense Borassus flabellifer, Machilus thunbergii and Mucuna ferruginea Arecaceae, Fabaceae, and Lauraceae China, India and Japan Rao and Rao (1964); Ichinoe (1971); Farr and Rossman (2024)
Camposporium indicum Borassus flabellifer Arecaceae India Rao and Rao (1964)
Camposporium japonicum Acacia confusa, Betula pendula, Cinnamomum japonicum, Freycinetia arborea, Litchi chinensis, Machilus thunbergii, Mucuna ferruginea, Paulownia kawakamii and Quercus sp. Betulaceae, Lauraceae, Pandanaceae, Paulowniaceae, and Sapindaceae China and Japan Ichinoe (1971); Matsushima (1980); Farr and Rossman (2024)
Camposporium laundonii Aralia elata, Pasania edulis and Rosa sp. Araliaceae, Fagaceae, and Rosaceae Japan and New Zealand Ellis (1976); Watanabe (1993); Farr and Rossman (2024)
Camposporium lycopodiellae Lycopodiella inundata Lycopodiaceae Germany Hyde et al. (2020)
Camposporium marylandicum Unidentified host The United States Shearer (1974)
Camposporium microsporum Borassus flabellifer Arecaceae India Rao and Rao (1964)
Camposporium multiseptatum Unidentified host China Hyde et al. (2020)
Camposporium ontariense Acer saccharum and Freycinetia arborea Sapindaceae and Pandanaceae Canada and the United States Matsushima (1983); Whitton et al. (2002)
Camposporium pellucidum Betula pendula, Caesalpinia echinata, Carpodetus serratus, Elaeagnus sp., Fagus sp., Laurus sp., Pasania glabra, Picea abies, Rhopalostylis sp. and Sorbus aucuparia Areceae, Betulaceae, Elaeagnaceae, Fabaceae, Lauraceae, Pinaceae, Rosaceae, and Rousseaceae Brazil, Japan, Poland, New Zealand, Russia and the United Kingdom Hughes (1951); Matsushima (1983); Mulenko et al. (2008); Farr and Rossman (2024)
Camposporium quercicola Quercus germana Fagaceae Mexico Mercado Sierra et al. (1995)
Camposporium ramosum Freycinetia sp. Australia and the United States Whitton et al. (2002)
Camposporium scolecosporium Unidentified host Papua New Guinea Kobayasi (1971)
Camposporium septatum Unidentified host Thailand Hyde et al. (2020)
Camposporium valdivianum Sophora microphylla Fabaceae The United States Koukol and Delgado (2021)
Camposporium verruculosum Clematis vitalba Ranunculaceae Italy Koukol and Delgado (2021)
Dematiomelanomma yunnanense Hypericum monogynum and Rubus parvifolius Hypericaceae and Rosaceae China Gao et al. (2023)
Exosporiella fungorum Thelephora sp. (leathery earthfans/mushroom sp.) Sweden Karsten (1892)
Fusiconidium mackenziei Clematis vitalba Ranunculaceae Italy Li et al. (2017)
Herpotrichia alligata Opuntia sp. Cactaceae Sweden and the United States Barr (1992); Farr and Rossman (2024)
Herpotrichia alpincola Aconitum sp. Ranunculaceae Hungary and Slovakia Rehm (1906)
Herpotrichia arizonica Carnegiea gigantea Cactaceae The United States Barr (1992); Farr and Rossman (2024)
Herpotrichia australis Sclerocarya caffra Anacardiaceae South Africa Bose (1961)
Herpotrichia bakeri Sambucus javanica Viburnaceae Philippines Teodoro (1937); Farr and Rossman (2024)
Herpotrichia bambusana Bambusa vulgaris Poaceae Brazil Hennings (1908); Eriksson and Yue (1998)
Herpotrichia boldoae Boldea fragrans and Peumus boldus Monimiaceae Chile Spegazzini (1910); Mujica and Vergara (1945); Spaulding (1961); Farr (1973); Farr and Rossman (2024)
Herpotrichia brasiliensis Unidentified host Brazil Rick (1933)
Herpotrichia brenckleana Urtica gracilis Urticaceae Sweden Eriksson (2014); Farr and Rossman (2024
Herpotrichia caesalpiniae Caesalpinia sepiaria Fabaceae South Africa Doidge (1948); Sivanesan (1971)
Herpotrichia calamicola Calamus caryotoides Arecaceae Australia Fröhlich and Hyde (2000); Farr and Rossman (2024)
Herpotrichia callimorpha Chamaenerion angustifolium, Salix sp. and Xanthophyllum flavescens Onagraceae, Polygalaceae, and Salicaceae Denmark and India Munk (1957); Rabenhorst (1869); Pande (2008); Farr and Rossman (2024)
Herpotrichia caulogena Silene nutans Caryophyllaceae Luxembourg Feltgen (1903)
Herpotrichia chilensis Proustia pungens Asteraceae Chile Feltgen (1903); Farr (1973); Farr and Rossman (2024)
Herpotrichia cirrhostoma Unidentified host Sri Lanka Berkeley and Broome (1873); Petch (1912)
Herpotrichia dalisayi Unidentified host The Philippines Hyde and Aptroot (1998)
Herpotrichia decidua Unidentified host The United States Barr (1992); Farr and Rossman (2024)
Herpotrichia detzneriae Detzneria tubata Plantaginaceae Papua New Guinea Kobayasi (1971); Shaw (1984); Farr and Rossman (2024)
Herpotrichia diffusa Juglans cinerea and Populus sp. Juglandaceae Palestine and the United States Ellis and Everhart (1892); Ellis (1895)
Herpotrichia ellisii Abies sp. Pinaceae Canada Barr (1992); Farr and Rossman (2024)
Herpotrichia ephedrae Ephedra distachya Ephedraceae France Kuhnholtz-Lordat and Barry (1949)
Herpotrichia fusispora Unidentified host China Chen and Hsieh (2004); Farr and Rossman (2024)
Herpotrichia gelasinosporoides Soil India Von Arx (1981)
Herpotrichia henkeliana Phragmites communis Poaceae Germany Sydow and Sydow (1921)
Herpotrichia herbarum Achillea millefolium Asteraceae Wehmeyer (1952); Barr (1992)
Herpotrichia herpotrichoides Carya sp., Epilobium sp., Rhododendron hirsutum, Ribes sp. and Rubus sp. Ericaceae, Juglandaceae, Grossulariaceae, Onagraceae, and Rosaceae Austria, Denmark, Germany, Poland, the United States and the United Kingdom Cannon (1982); Barr (1984); Cannon et al. (1985); Mulenko et al. (2008); Zhang et al. (2012); Tian et al. (2015); Farr and Rossman (2024)
Herpotrichia hippocrateae Hippocratea grahamii Celastraceae India Tilak and Talde (1975); Pande (2008)
Herpotrichia indica Duranta plumieri Verbenaceae India Anahosur (1970); Pande (2008); Farr and Rossman (2024)
Herpotrichia laricina Larix decidua Pinaceae Luxembourg Feltgen (1901)
Herpotrichia leptospora Unidentified host Kirschstein (1911)
Herpotrichia lignicola Unidentified host Belgium Bose (1961)
Herpotrichia macrotricha Acer spicatum, Agropyron repens, Agrostis alba, Arundinaria sp., Carex sp., Cocos nucifera, Eupatorium formosanum, Fagus sylvatica, Fraxinus sp., Rubus sp. and Solidago sp. Arecaceae, Asteraceae, Cupressaceae, Cyperaceae, Fagaceae, Oleaceae, Poaceae, Rosaceae, and Sapindaceae China, India, the United Kingdom and the United States Dennis (1978); Saccardo (1883); Barr (1968, 1984); Pande (2008); Chen and Hsieh (2004); Farr and Rossman (2024)
Herpotrichia mangrovei Unidentified host China Jones and Vrijmoed (2003)
Herpotrichia melanotricha Hevea brasiliensis Euphorbiaceae Congo Saccas (1954)
Herpotrichia millettiae Millettia sp. Fabaceae Malaysia Sivanesan (1971)
Herpotrichia monospermatis Butea monosperma Fabaceae India Pande (2008); Farr and Rossman (2024)
Herpotrichia mulleri Artemisia nilagirica and Butea monosperma Asteraceae, Fabaceae India Pande (2008); Farr and Rossman (2024)
Herpotrichia myriangii Carica papaya Caricaceae Java Raciborski (1909)
Herpotrichia nectrioides Melastomataceae sp. Melastomataceae Brazil Rehm (1901)
Herpotrichia nigra Abies sp., Calocedrus decurrens, Cedrus libani, Chamaecyparis nootkatensis, Juniperus sp., Phyllodoce sp., Picea sp., Pinus sp., Pseudotsuga menziesii and Rhododendron sp. Cupressaceae, Ericaceae, and Pinaceae Austria, Canada, France, Germany, Italy, Norway, Poland, Switzerland, Turkey, Ukraine and the United States Shaw (1973); Ginns (1986); French (1989); Farr and Rossman (2024)
Herpotrichia nigrotuberculata Elaeis guineensis and Phyllostachys reticulata Arecaceae and Poaceae Japan and Tanzania Pirozynski (1972); Farr and Rossman (2024)
Herpotrichia nypicola Nypa fruticans Arecaceae Malaysia Hyde et al. (1999); Farr and Rossman (2024)
Herpotrichia occulta Eucalyptus sp. Myrtaceae Brazil Rick (1933)
Herpotrichia ochrostoma Fraxinus excelsior Oleaceae Luxembourg Feltgen (1903)
Herpotrichia palmicola Calamus caryotoides, Daemonorops sp. and Licuala ramsayi Arecaceae Australia and China Hyde et al. (1999); Fröhlich and Hyde 2000; Zhuang (2001); Farr and Rossman (2024)
Herpotrichia pandei Saccharum spontaneum Poaceae India Bose (1961)
Herpotrichia petrakiana Fagus sylvatica Fagaceae Bose (1961)
Herpotrichia philippinensis Alstonia scholaris Apocynaceae The Philippines Rehm (1914); Teodoro (1937); Farr and Rossman (2024)
Herpotrichia pinetorum Herpotrichia pinetorum Pinaceae Austria and India Winter (1885); Mueller (1958); Petrak (1962); Farr and Rossman (2024)
Herpotrichia quinqueseptata Abies lasiocarpa, Larix europaea, Picea sp. and Populus tremula Pinaceae and Salicaceae Canada, Czech Republic, Germany, Sweden and the United States Shaw (1973); Minter (1981); Ginns (1986); Eriksson (2014); Farr and Rossman (2024)
Herpotrichia rara Tanacetum vulgare Asteraceae Germany Kirschstein (1935)
Herpotrichia rhenana Epilobium angustifolium, Rhododendron hirsutum and Ribes sp. Ericaceae, Grossulariaceae, and Onagraceae Austria and the United States Fuckel (1870); Remler (1979); Farr and Rossman (2024)
Herpotrichia rhodospiloides Populus deltoides Salicaceae The United States Peck (1909)
Herpotrichia rhodosticta Carex paniculata and Populus sp. Cyperaceae and Salicaceae Germany and the United States Saccardo (1882); Samuels (1976); Farr and Rossman (2024)
Herpotrichia setosa Betula glandulosa and Myrica gale Betulaceae and Myricaceae Canada and Ireland Barr (1992); Farr and Rossman (2024)
Herpotrichia striatispora Soil South Africa Papendorf and Arx (1966)
Herpotrichia symphoricarpi Symphoricarpos sp. Caprifoliaceae The United States Barr (1984); Farr and Rossman (2024)
Herpotrichia tenuispora Urtica dioica Urticaceae Germany Kirschstein (1906)
Herpotrichia vaginatispora Trifolium sp. Fabaceae Italy Tian et al. (2015)
Herpotrichia villosa Unidentified host Brazil Samuels and Müller (1978)
Herpotrichia xiaokongensis Prunus sp. Rosaceae China Hyde et al. (2021)
Herpotrichia zingiberacearum Hedychium coronarium Zingiberaceae China This study
Mamillisphaeria dimorphospora Unidentified host Australia Hyde et al. (1996)
Marjia tianshanica Cerasus tianshanica Rosaceae Uzbekistan Wanasinghe et al. (2018)
Marjia uzbekistanica Rosa sp. Rosaceae Uzbekistan Wanasinghe et al. (2018)
Melanocamarosporioides ugamica Lonicera altmannii Caprifoliaceae Uzbekistan Pem et al. (2019)
Melanocamarosporium galiicola Galium sp. Rubiaceae Italy Wijayawardene et al. (2016)
Melanocucurbitaria uzbekistanica Acer pubescens Sapindaceae Uzbekistan Wanasinghe et al. (2018)
Melanodiplodia tianschanica Rosa sp. Rosaceae Uzbekistan Wanasinghe et al. (2018)
Melanomma acanthophilum Cereus quisco Cactaceae Chile Spegazzini (1923); Farr (1973); Farr and Rossman (2024)
Melanomma afflatum Unidentified host The United States Shear (1941)
Melanomma anceps Unidentified host Jawa Höhnel (1909)
Melanomma andinum Bulnesia retamo Zygophyllaceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Melanomma artemisiae-maritimae Artemisia maritima Asteraceae Russia Lobik (1927); Farr and Rossman (2024)
Melanomma aspegrenii Carpinus betulus, Cornus sp. and Fagus sylvatica Betulaceae, Cornaceae, and Fagaceae Poland Fuckel (1870); Mulenko et al. (2008); Farr and Rossman (2024)
Melanomma aurantiicola Citrus sp. Rutaceae Paraguay Spegazzini (1920); Farr (1973); Farr and Rossman (2024)
Melanomma aurantiiphila Citrus sp. Rutaceae Paraguay Spegazzini (1920); Farr (1973); Farr and Rossman (2024)
Melanomma australiense Unidentified host Australia Hyde and Goh (1999)
Melanomma brachythele Sambucus sp. Adoxaceae The United Kingdom Saccardo (1878)
Melanomma bubakii Campanula stricta Campanulaceae Turkey Bubák (1914)
Melanomma cacheutense Baccharis glutinosa Asteraceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Melanomma caesalpiniae Caesalpinia cearense Fabaceae Brazil Hennings (1908)
Melanomma caryophagum Carya sp. and Juglans sp. Juglandaceae The United States Fairman (1921)
Melanomma castillejae Castilleja pallida Orobanchaceae Siberia Farr and Rossman (2024)
Melanomma ceratoniae Ceratonia siliqua Fabaceae Spain Farr and Rossman (2024)
Melanomma chilense Proustia pungens Asteraceae Chile Spegazzini (1910); Farr (1973); Farr and Rossman (2024)
Melanomma citricola Citrus medica Rutaceae Bangladesh and India Sydow et al. (1911); Rao (1969); Farr and Rossman (2024)
Melanomma conjunctum Thuja plicata Cupressaceae The United States Farr and Rossman (2024)
Melanomma cryptostegiae Cryptostegia grandiflora Apocynaceae India Pande (2008); Farr and Rossman (2024)
Melanomma cucurbitarioideum Pentaphylloides fruticosa Rosaceae China Yuan and Barr (1994); Farr and Rossman (2024)
Melanomma dactylosporum Unidentified host Brazil Rehm (1901)
Melanomma dinghuense Unidentified host China Inderbitzin and Huang (2001)
Melanomma distinctum Pentaphylloides fruticosa Rosaceae Russia Vasilyeva (1987)
Melanomma drimydis Drymis sp. Winteraceae Brazil Rehm (1901)
Melanomma dryinum Quercus sp. Fagaceae Belgium Mouton (1900)
Melanomma dzungaricum Eurotia eversmanniana Amaranthaceae Kazakhstan Vasilyeva (1987)
Melanomma ebeni Ebenus stellata Fabaceae Iran González Fragoso (1918)
Melanomma epiphytica Bambusa sp. Poaceae Indonesia Raciborski (1909); Eriksson and Yue (1998); Farr and Rossman (2024)
Melanomma gigantica Unidentified host India Pande (1979)
Melanomma glumarum Oryza sativa Poaceae China, India, Japan and the Philippines Watson (1971); Reinking (1919); Farr and Rossman (2024)
Melanomma gregarium Populus sp. Salicaceae The United States Cash (1953); Farr and Rossman (2024)
Melanomma halimodendri Halimodendron halodendron Fabaceae Kazakhstan Farr and Rossman (2024)
Melanomma haloxyli Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Melanomma helianthemi Helianthemum rupifragum Cistaceae Ukraine Gucevic (1969)
Melanomma heraclei Heracleum pubescens Apiaceae Ukraine Dudka et al. (2004); Farr and Rossman (2024)
Melanomma herpotrichum Populus sp. Salicaceae Luxembourg Feltgen (1903)
Melanomma japonicum Fagus crenata Fagaceae Japan Hashimoto et al. (2017); Farr and Rossman (2024)
Melanomma jenynsii Unidentified host The United Kingdom Saccardo (1883)
Melanomma juniperi Juniperus virginiana Cupressaceae The United States Fairman (1905); Cash (1953); Farr and Rossman (2024)
Melanomma langloisii Salix nigra Salicaceae The United States Farr and Rossman (2024)
Melanomma lithophilae Sobolewskia lithophila Apocynaceae Ukraine Dudka et al. (2004); Farr and Rossman (2024)
Melanomma longicolle Acer sp. and Citrus limon Aceraceae and Rutaceae Italy and the United Kingdom Saccardo (1875); Farr and Rossman (2024)
Melanomma marathawadense On paper India Tilak and Kale (1970)
Melanomma margaretae Dryas octopetala Rosaceae Poland Mulenko et al. (2008); Farr and Rossman (2024)
Melanomma martinianum Unidentified host New Zealand Saccardo (1883)
Melanomma mate Ilex paraguensis Aquifoliaceae Argentina Spegazzini (1908); Farr (1973); Farr and Rossman (2024)
Melanomma medium Acer negundo, Calligonum sp. and Tamarix sp. Sapindaceae and Tamaricaceae Canada, Italy and the United Kingdom Saccardo (1878); Farr (1973); Farr and Rossman (2024)
Melanomma mindorense Arenga mindorensis Arecaceae The Philippines Rehm (1913)
Melanomma mojunkumica Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Melanomma moravicum Unidentified host Slovakia Farr and Rossman (2024)
Melanomma mutabile Solanum dulcamara Solanaceae Luxembourg Feltgen (1901)
Melanomma myricae Myrica gale Myricaceae Sweden Eriksson (2014); Farr and Rossman (2024)
Melanomma nigriseda Fagus sp. Fagaceae The United States Fairman (1922)
Melanomma obliterans Unidentified host The United Kingdom Saccardo (1883)
Melanomma obtusissimum Unidentified host Cuba Farr and Rossman (2024)
Melanomma oryzae Oryza sativa Poaceae Japan Farr and Rossman (2024)
Melanomma oxysporum Quercus sp. Fagaceae The United States Hawksworth (1985)
Melanomma panici-miliacei Panicum miliaceum Poaceae Siberia Farr and Rossman (2024)
Melanomma philippinense Unidentified host The Philippines Sydow and Sydow (1914)
Melanomma populicola Cornus sp., Fagus sylvatica, Picea abies, Populus sp., Quercus sp. and Sorbus aucuparia Cornaceae, Fagaceae, Pinaceae, Rosaceae, and Salicaceae China, Croatia, Germany and the Netherlands Crous et al. (2020); Farr and Rossman (2024); this study
Melanomma praeandinum Salvia gilliesii Lamiaceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Melanomma pulveracea Unidentified host China Teng (1936)
Melanomma pulvis-pyrius Acer sp., Albizia julibrissin, Alhagi sp., Alnus sp., Berberis sp., Betula sp., Bupleurum fruticosum, Campsis radicans, Carpinus betulus, Celtis australis, Corylus sp., Larix decidua and Pinus sylvestris Apiaceae, Betulaceae, Berberidaceae, Bignoniaceae, Cannabaceae, Fabaceae, Pinaceae, and Sapindaceae Canada, Czech Republic, Japan, Poland, Russia, Scotland, Sweden and Ukraine Mulenko et al. (2008); Hashimoto et al. (2017); Crous et al. (2020); Farr and Rossman (2024)
Melanomma pyriostictum Unidentified host The United Kingdom Cooke (1887)
Melanomma rhododendri Rhododendron sp. Ericaceae Austria, Belgium, Germany, Italy, Luxembourg, the Netherlands, Switzerland, the United Kingdom and the United States Rehm (1881, 1906); Bommer and Rousseau (1890); Farr and Rossman (2024)
Melanomma ribis Ribes sp. Grossulariaceae The United States Barr (1990)
Melanomma rubicundum Lythrum sp. Lythraceae Sweden Bommer and Rousseau (1890); Farr and Rossman (2024)
Melanomma sanguinarium Unidentified host Saccardo (1878)
Melanomma saviczii Thymus pseudohumillimus Lamiaceae Ukraine Dudka et al. (2004); Farr and Rossman (2024)
Melanomma scrophulariae Scrophularia rupestris Scrophulariaceae Ukraine Gucevic (1967)
Melanomma sordidissimum Eriobotrya japonica Rosaceae Argentina Spegazzini (1909); Farr (1973); Farr and Rossman (2024)
Melanomma sparsum Abies sp. Pinaceae Switzerland Fuckel (1873)
Melanomma spiniferum Morus alba Moraceae The United States Cash (1953); Farr and Rossman (2024)
Melanomma subandinum Atriplex pamparum Amaranthaceae Argentina Cash (1953); Farr and Rossman (2024)
Melanomma subdispersum Betula sp., and Fagus sp. Betulaceae and Fagaceae Canada, Ireland, Poland, and the United Kingdom Conners (1967); Mulenko et al. (2008); Farr and Rossman (2024)
Melanomma submojunkumica Haloxylon aphyllum Amaranthaceae Kazakhstan Farr and Rossman (2024)
Melanomma thespesiae Thespesia sp. Malvaceae India Farr and Rossman (2024)
Melanomma trevoae Trevoa trinervia Rhamnaceae Chile Farr (1973); Farr and Rossman (2024)
Melanomma vile Quercus sp. Fagaceae Sweden Fuckel (1870)
Melanomma xylariae Unidentified host Brazil Höhnel (1907)
Monoseptella rosae Rosa sp. Rosaceae Uzbekistan Wanasinghe et al. (2018)
Muriformistrickeria rosae Rosa sp. Rosaceae Italy Wanasinghe et al. (2018)
Muriformistrickeria rubi Rubus sp. Rosaceae Italy Tian et al. (2015)
Navicella costaricensis Unidentified host The United States El-Shafie et al. (2005)
Navicella diabola Castanopsis sp. Fagaceae China Aptroot (2003)
Navicella pallida Elaeocarpus sp. Elaeocarpaceae Papua New Guinea Aptroot and Iperen (1998)
Navicella pileata Fraxinus sp., Salix fragilis, Tilia sp. and Quercus sp. Fagaceae, Malvaceae, Oleaceae, and Salicaceae Finland, Lithuania and Norway Fabre (1879); Holm and Holm (1988); Farr and Rossman (2024)
Navicella xinjiangensis Lonicera hispida, Haloxylon ammodendron Amaranthaceae and Caprifoliaceae China Yuan and Zhao (1994); Farr and Rossman (2024)
Neobyssosphaeria clematidis Clematis vitalba Ranunculaceae The United Kingdom Phukhamsakda et al. (2020)
Petrakia aesculi Aesculus turbinata Sapindaceae Japan Jaklitsch and Voglmayr (2017)
Petrakia deviata Acer campestre Sapindaceae Georgia and Switzerland Watzl (1937); Gross et al. (2017)
Petrakia echinata Acer sp. Sapindaceae Italy, Slovakia, Switzerland and the United States Sydow and Sydow (1913); Li et al. (2016a); Gross et al. (2017); Farr and Rossman (2024)
Petrakia fagi Fagus crenata Fagaceae Japan Beenken et al. (2020)
Petrakia greenei Acer saccharinum Sapindaceae The United States Beenken et al. (2020)
Petrakia irregularis Acer pseudoplatanus Sapindaceae The Netherlands and Poland Mulenko et al. (2008); Farr and Rossman (2024)
Petrakia juniperi Juniperus sp. Cupressaceae Germany Bedlan (2017)
Petrakia liobae Fagus sylvatica Fagaceae Switzerland Beenken et al. (2020)
Petrakia minima Fagus japonica Fagaceae Japan Hashimoto et al. (2017); Beenken et al. (2020)
Petrakia paracochinensis Miscanthus floridulus Poaceae China Wong et al. (2002)
Phragmocephala atra Rhopalostylis sapida and Urtica sp. Arecaceae, Urticaceae New Zealand and the United Kingdom Mason and Hughes (1951); McKenzie et al. (1992)
Phragmocephala elegans Gymnanthes lucida Euphorbiaceae Brazil and Cuba Castañeda (1985)
Phragmocephala elliptica Elaeagnus sp., Filipendula denudata, Laurus sp. Quercus robur and Sambucus sp. Adoxaceae, Elaeagnaceae, Fagaceae, Lauraceae, and Rosaceae Canada, Russia, Ukraine and the United Kingdom Hughes (1979); Dennis (1986); Holubtsova and Andrianova (2008); Farr and Rossman (2024)
Phragmocephala garethjonesii Unidentified host China Su et al. (2015)
Phragmocephala glanduliformis Corticium coronatum, Picea obovate and Quercus sp. Corticiaceae, Fagaceae, and Pinaceae Austria Hughes (1955); Farr and Rossman (2024)
Phragmocephala hughesii Unidentified host China Wu and Zhuang (2005)
Phragmocephala minima Abies balsamea and Fagus sylvatica Fagaceae and Pinaceae Canada and the United Kingdom Mason and Hughes (1951); Conners (1967)
Phragmocephala prolifera Populus tremuloides and Urtica dioica Salicaceae and Urticaceae Belgium and Canada Hughes (1979); Farr and Rossman (2024)
Phragmocephala stemphylioides Carya sp., Cistus sp. and Pistacia lentiscus Anacardiaceae, Cistaceae, and Juglandaceae Brazil, Canada, China and Italy Hughes (1958); Zhuang (2005); Lunghini et al. (2013); Farr and Rossman (2024)
Phragmotrichum andamanense Strobilanthes sp. Acanthaceae India Bhat and Kendrick (1993)
Phragmotrichum chailletii Abies sp. and Picea sp. Pinaceae Canada, Switzerland, Romania and the United States Kunze and Schmidt (1823); Ginns (1986); Farr and Rossman (2024)
Phragmotrichum karstenii Acer platanoides Sapindaceae Finland Sutton and Pirozynski (1965)
Phragmotrichum rivoclarinum Acer sp., Alnus sp. and Salix sp. Betulaceae, Salicaceae, and Sapindaceae Canada, Italy and the United Kingdom Sutton and Pirozynski (1966); Ginns (1986); Farr and Rossman (2024)
Phragmotrichum vassiljevae Alnus kamtschatica Betulaceae Russia Melnik (1984)
Pleotrichocladium opacum Soil Spain Hernández-Restrepo et al. (2017)
Praetumpfia obducens Fraxinus excelsior Oleaceae Austria and Sweden Jaklitsch and Voglmayr (2017)
Pseudobyssosphaeria bambusae Bamboo sp. Poaceae Thailand Hyde et al. (2018)
Pseudostrickeria muriformis Origanum vulgare Lamiaceae Italy Tian et al. (2015)
Pseudostrickeria ononidis Ononis spinosa Fabaceae Italy Tian et al. (2015)
Pseudostrickeria rosae Rosa sp. Rosaceae Italy Wanasinghe et al. (2018)
Sarimanas pseudofluviatile Unidentified host Japan Liu et al. (2015)
Sarimanas shirakamiense Swida controversa Cornaceae Japan Liu et al. (2015)
Seifertia alpina Rhododendron ponticum Ericaceae Austria Beenken et al. (2020)
Seifertia azaleae Ledum groenlandicum, Leucopogon costatus and Rhododendron sp. Ericaceae Australia, Canada, China, Germany, Italy, Japan, the Netherlands, New Zealand, Panama, Switzerland, the United Kingdom and the United States White and Hamilton (1935); Partridge and Morgan-Jones (2002); Farr and Rossman (2024)
Seifertia shangrilaensis Rhododendron decorum Ericaceae China Li et al. (2016b)
Tumularia aquatica Alnus glutinosa, Phragmites sp. and Quercus sp. Betulaceae, Fagaceae, and Poaceae South Africa and the United Kingdom Sivanesan (1984); Marvanová and Descals (1987); Farr and Rossman (2024)
Tumularia tuberculata Fagus sylvatica, Quercus sp. Fagaceae Hungary Gönczöl (1976)
Uzbekistanica pruni Prunus armeniaca Rosaceae Russia Hyde et al. (2020)
Uzbekistanica rosae-hissaricae Rosa sp. Rosaceae Uzbekistan Wanasinghe et al. (2018)
Uzbekistanica vitis-viniferae Vitis vinifera Vitaceae Ukraine Crous et al. (2020)
Uzbekistanica yakutkhanika Rosa sp. Rosaceae Uzbekistan Wanasinghe et al. (2018)
Xenostigmina aceris Acer macrophyllum Sapindaceae The United States Hashimoto et al. (2017)

The distribution map was prepared using MapChart programme (https://www.mapchart.net/index.html), a platform from which a personalised map of the world using different colours can be generated. The Sankey diagram was created to show the species distribution through plant host families using the free online tool SankeyMATIC by Steve Bogart (www.sankeymatic.com).

Results

Phylogenetic analyses

Phylogenetic analyses of combined LSU, SSU, ITS and tef1-α sequences comprised 3480 characters including gaps. Hysterium angustatum (MFLU 16-1179) was used as the outgroup taxon. The RAxML analysis of the combined dataset yielded a best scoring tree (Fig. 3). The final ML optimisation likelihood value was -16620.192034. There were 35.32% undetermined characters or gaps and 1071 distinct alignment patterns. Estimated base frequencies were A = 0.247534, C = 0.235205, G = 0.272353, T = 0.244907; substitution rates AC = 1.775315, AG = 3.161447, AT = 1.766474, CG = 1.002405, CT = 11.511066, GT = 1.000; proportion of invariable sites I = 0.535605; gamma distribution shape parameter α = 0.513218. The Bayesian analysis has resulted in 30,000 trees after 3,000,000 generations. Bootstrap support values for ML higher than 70% and BYPP greater than 0.90 are given above each branch respectively (Fig. 3). All analyses (ML and BYPP) showed similar topologies and concurred with previous studies (Pem et al. 2019; Kularathnage et al. 2022; Gao et al. 2023).

Figure 3. 

Phylogram generated from Maximum Likelihood analysis is based on combined LSU, SSU, ITS and tef1-α sequence data. The tree is rooted with Hysterium angustatum (MFLU 16-1179). The new isolates are in red and ex-type strains are indicated in bold face. Bootstrap support values ≥ 70% from the Maximum Likelihood (ML) and Bayesian Posterior Probabilities (BYPP) values ≥ 0.90 are given above the nodes, respectively.

According to the phylogeny, our collection HFJAU10336, HFJAU10337 and HFJAU10338 cluster within Byssosphaeria species. HFJAU10337 and HFJAU10338 cluster together and show an independent lineage, sister to B. phoenicis isolates (ZHKUCC 21-0122 and ZHKUCC 21-0123) with 83% ML and 0.99 BYPP statistical support. The isolate HFJAU10336 clusters with B. siamensis isolates (MFLUCC 10-0099, MFLUCC 17-1800 and KUMCC 21-0339) in a monophyletic clade. In addition, CBUB 40045 groups with Bertiella fici in a 100% ML and 1.00 BYPP supported clade. Three isolates (HFJAU10332, HFJAU10333 and HFJAU10334) cluster with Herpotrichia species, and show a close phylogenetic relationship with H. melanotricha (GKM 196N, SMH 269 and JCM 14419). As well as HFJAU10335 groups with Melanomma populicola (CBS 130330, CBS 350.82, CBS 543.70 and CPC 272203) in a 99% ML and 0.99 BYPP supported clade.

Taxonomy

Bertiella (Sacc.) Sacc

Notes

Bertiella was established by Saccardo and Sydow (1899) to include B. macrospora as the type species. The species have superficial ascomata, cylindrical-clavate asci and hyaline, 1-septate (when immature) and pale brown, 3-septate (when mature) ascospores (Tian et al. 2015; Hongsanan et al. 2020). To the present time, there are six Bertiella species in Species Fungorum (2024). Of them, molecular data are available only for three species. Bertiella species have been reported from six different plant families, Cyrillaceae, Fagaceae, Lauraceae, Moraceae, Salicaceae and Ulmaceae (Fig. 10).

Bertiella fici Tennakoon, C.H. Kuo & K.D. Hyde, Fungal Diversity 108: 29 (2021)

Fig. 4

Description

Saprobic on dead leaves of Cinnamomum verum J. Presl (Lauraceae). Sexual morph: Ascomata 160–220 × 230–280 µm ( = 180 × 240 μm, n = 15), solitary or scattered, semi-immersed to superficial, appeared as black dots on host surface, globose to subglobose, glabrous, unilocular, ostiolate. Peridium 12–20 μm wide, thick-walled, carbonaceous, composed of several layers of brown to dark brown pseudoparenchymatous cells, cells towards the inside hyaline, arranged in a textura angularis, fusing at the outside indistinguishable from the host tissues. Hamathecium comprising numerous, 1–2 µm wide, hyaline, septate, cellular pseudoparaphyses. Asci 50–60 × 7.5–8.5 μm ( = 52 × 7.8 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindrical-clavate, short pedicellate, apically rounded, with a distinct ocular chamber. Ascospores 14–18 × 4–5 μm ( = 15 × 4.2 μm, n = 40), overlapping, 1–2-seriate, fusiform, initially hyaline, becoming yellowish-brown at maturity, 1-septate, slightly curved, slightly constricted at the septum, guttulate, smooth-walled. Asexual morph: Undetermined.

Figure 4. 

Bertiella fici (CMUB 40045, new host record) a appearance of ascomata on the host b, c close-up of ascomata d vertical section of an ascoma e peridium f pseudoparaphyses g, h asci i–n ascospores. Scale bars: 75 µm (d); 10 µm (e); 20 µm (f–h); 7 µm (i–n).

Material examined

Thailand, Chiang Rai, Doi Mae Salong Mountain, on a dead leaf of Cinnamomum verum (Lauraceae), 15 June 2020, D. S. Tennakoon, DMS002 (CMUB 40045).

Known hosts

Cinnamomum verum and Ficus septica (Tennakoon et al. 2021; this study).

Known distribution

China and Thailand (Tennakoon et al. 2021; this study)

Notes

Bertiella fici was introduced by Tennakoon et al. (2021) from dead leaves of Ficus septica in China. The morphological characteristics of our collection (CMUB 40045) tally well with the B. fici in having solitary or scattered, semi-immersed to superficial ascomata, cylindrical to cylindrical-clavate asci and yellowish-brown, 1-septate ascospores with overlapping size ranges (Tennakoon et al. 2021). Multi-gene phylogeny (LSU, SSU, ITS and tef1-α) also indicates that our collection nested with B. fici isolates in a 100% ML and 1.00 BYPP supported clade. This was further confirmed by having only two nucleotide differences in the LSU and SSU genes between our collection and the type of Bertiella fici. Thus, we conclude our collection as a new host record of Bertiella fici from Cinnamomum verum. In addition, this is the first Bertiella fici record from Thailand.

Byssosphaeria Cooke

Notes

Cooke and Plowright (1879) established Byssosphaeria to accommodate B. keithii as the type species. Byssosphaeria species have superficial ascomata with bright yellow, orange or red flat apices around the ostiole, with dependent hyphal appendages that merge with the subiculum below and hyaline ascospores (Tian et al. 2015; Tennakoon et al. 2018). Species have cosmopolitan distribution as saprobes in various plant substrates (e.g. dead leaves, wood). As well, Byssosphaeria species have been reported from 15 plant families (Fig. 10). Currently, there are 18 accepted species in Species Fungorum (2024).

Byssosphaeria poaceicola Tennakoon & D.M. Hu, sp. nov.

Fig. 5

Etymology

Named after the host family (Poaceae) where this fungus was collected.

Holotype

HFJAU10338.

Description

Saprobic on dead stem of Arundo pliniana Turra (Poaceae). Sexual morph: Ascomata 550–650 × 600–800 µm ( = 610 × 715 μm, n = 10), solitary to gregarious, superficial, dark brown to black, setose, coriaceous, unilocular, globose to subglobose, non-papillate, apex rounded with an orange to yellow ostiole, ostiole central, with pore-like opening, periphysate. Peridium 30–45 μm wide, thick-walled, composed of 6–7 layers of dark brown cells, orange to yellow near ostiole, arranged in textura angularis. Hamathecium 1–2.5 μm wide, comprising dense, filiform, anastomosing, septate, trabeculate pseudoparaphyses, embedded in a gelatinous matrix. Asci 165–180 × 12–15 μm ( = 171 × 13 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical clavate, apically rounded, long pedicellate (30–40 μm), with an indistinct ocular chamber. Ascospores 32–40 × 7–8 μm ( = 36 × 7.5 μm, n = 30), overlapping, 1–2-seriate, ellipsoid to fusiform, initially hyaline, pale brown when mature, 1-septate, constricted at the septum, slightly curved, guttulate, smooth-walled. Asexual morph: Undetermined.

Figure 5. 

Byssosphaeria poaceicola (HFJAU10338, holotype) a, b appearance of ascomata on the host c vertical section of an ascoma d ostiole e peridium f pseudoparaphyses g–i asci j–o ascospores. Scale bars: 200 µm (c); 20 µm (d, e); 50 µm (f–i); 10 µm (j–o).

Material examined

China, Yunnan Province, Kunming, on a dead stem of Arundo pliniana (Poaceae), 22 July 2016, D. S. Tennakoon, KDS30 (HFJAU10338, holotype); ibid. 28 August 2016, KDS29 (HFJAU10337, paratype).

Notes

In the combined LSU, SSU, ITS and tef1-α phylogenetic analysis, two strains of Byssosphaeria poaceicola (HFJAU10337 and HFJAU10338) formed a monophyletic clade sister to By. phoenicis strains (ZHKUCC 21-0122 and ZHKUCC 21-012) with 83% ML and 0.99 BYPP statistical support. Morphologically, they share similarities in having superficial, dark brown to black, coriaceous, non-papillate ascomata, cylindrical clavate asci and ellipsoid to fusiform, 1-septate, pale brown ascospores (Kularathnage et al. 2022). Although Byssosphaeria poaceicola can be distinguished from By. phoenicis in their size differences of asci (165–180 × 12–15 μm vs. 100–160 × 10–15 μm) and ascospores (32–40 × 7–8 μm vs. 25–30 × 5–7 μm) (Kularathnage et al. 2022). In addition, a comparison of the 497 nucleotides across the ITS (+5.8S) gene region of By. poaceicola and By. phoenicis shows 16 base pair differences (3.21%). It is interesting to notice that the Byssosphaeria species have not been collected much from Poaceae hosts, except for bamboo species (Jiang et al. 2022). Based on this finding, it appears that Byssosphaeria species can adapt to a variety of habitats, although there are limited studies to investigate their diversity on various hosts and regions.

Byssosphaeria siamensis Boonmee, Q. Tian & K.D. Hyde, Fungal Diversity 74: 283 (2015)

Fig. 6

Description

Saprobic on dead stem of Citrus trifoliata L. (Rutaceae). Sexual morph: Ascomata 250–400 × 300–500 µm ( = 320 × 410 μm, n = 10), solitary to gregarious, superficial, dark brown to black, setose, coriaceous, unilocular, globose to subglobose, non-papillate, apex rounded with an orange to yellow ostiole, ostiole central, with pore-like opening. Peridium 20–35 μm wide, thick-walled, composed of several layers of dark brown cells, orange to yellow near ostiole, arranged in textura angularis to textura prismatica. Hamathecium 1–2.5 μm wide, comprising dense, filiform, anastomosing, septate, pseudoparaphyses, embedded in a gelatinous matrix. Asci 110–130 × 11–13 μm ( = 120 × 12 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical clavate, apically rounded, long pedicellate (20–35 μm), with an ocular chamber. Ascospores 30–40 × 6.5–8 μm ( = 31 × 7 μm, n = 30), overlapping, 1–2-seriate, ellipsoid to fusiform, initially hyaline, pale brown when mature, 1-septate, constricted at the septum, slightly curved, smooth-walled or verrucose. Asexual morph: Undetermined.

Figure 6. 

Byssosphaeria siamensis (HFJAU10336, new host record) a appearance of ascomata on the host b close-up of ascomata c vertical section of an ascoma d ostiole e peridium f pseudoparaphyses g, h asci i–m ascospores. Scale bars: 200 µm (c); 20 µm (d, e); 50 µm (f–h); 10 µm (i–m).

Material examined

China, Yunnan Province, Kunming, on a dead stem of Citrus trifoliata (Rutaceae), 27 July 2016, D. S. Tennakoon, KDS27 (HFJAU10336).

Known hosts

Citrus trifoliata (this study).

Known distribution

China, Thailand (Tian et al. 2015; Hyde et al. 2018; this study).

Notes

The morphological characteristics of our collection (HFJAU10336) similar to the type of Byssosphaeria siamensis in their superficial, dark brown to black, setose, non-papillate ascomata, cylindrical clavate asci and ellipsoid to fusiform, pale brown, 1-septate ascospores (Tian et al. 2015). In addition, both share overlapping size ranges of asci (110–130 × 11–13 μm vs. 112–148 × 10–16 μm) and ascospores (30–40 × 6.5–8 μm vs. 40.5–50 × 7–11 μm) (Tian et al. 2015). However, our collection is lacking a mucilaginous sheath which is present in the type species (Tian et al. 2015). According to the multi-gene phylogeny, our collection nested with By. siamensis isolates in 86% ML and 0.98 BYPP supported clade and close to the isolate MFLUCC 17-1800 with 88% ML, 0.99 BYPP support. Therefore, based on both morphology and phylogeny evidence, we introduce our collection as a new host record of B. siamensis from Citrus trifoliata in China.

Herpotrichia Fuckel

Notes

The diverse genus Herpotrichia was established by Fuckel (1868) to include two species, H. rhenana and H. rubi, but without designating a type species. Thus, Bose (1961) assigned H. rhenana as the lectotype. Subsequently, Holm (1979) assigned H. herpotrichoides (synonymous with H. rubi) as the generic type (Cannon 1982). The species have erumpent to superficial ascomata, clavate to cylindrical asci with hyaline to pale brown, 1-septate ascospores (Sivanesan 1984; Tian et al. 2015). The asexual morph is pyrenochaeta-like with or without setae on the surface of the pycnidia (Sivanesan 1984; Hongsanan et al. 2020). Herpotrichia species have been reported as saprobes, mostly in dead wood substrates in 34 plant families (Fig. 10).

Herpotrichia zingiberacearum Tennakoon & D.M. Hu, sp. nov.

Fig. 7

Etymology

Named after the host family (Zingiberaceae) where this fungus was collected.

Holotype

HFJAU10332.

Description

Saprobic on dead stem of Hedychium coronarium J. Koenig (Zingiberaceae). Sexual morph: Ascomata 250–350 × 240–320 µm ( = 298 × 262 μm, n = 10), solitary to clustered, superficial, dark brown to black, setose, coriaceous, unilocular, globose to subglobose, rounded apex broadly cap-like, ostiolate. Peridium 15–25 μm wide, thick-walled, composed of 4–5 layers of dark brown to black cells, arranged in textura angularis. Hamathecium 1–3 μm wide, comprising dense, filiform, anastomosing, septate, branched pseudoparaphyses, embedded in a gelatinous matrix. Asci 85–98 × 10–14 μm ( = 94 × 12 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical clavate, apically rounded, short pedicellate, with an ocular chamber. Ascospores 25–30 × 5–6 μm ( = 28 × 5.2 μm, n = 30), overlapping, 1–2-seriate, fusoid with narrowly rounded ends, hyaline, 1-septate, straight to slightly curved, constricted at the septum, with large guttules, surrounded by an expanded gelatinous sheath pointed at both ends, 2.5–4 μm wide, smooth-walled. Asexual morph: Undetermined.

Figure 7. 

Herpotrichia zingiberacearum (HFJAU10332, holotype) a, b appearance of ascomata on the host c vertical section of an ascoma d peridium e pseudoparaphyses f–h asci i–l ascospores m ascospore stained in Indian ink showing a mucilaginous sheath. Scale bars: 100 µm (c); 20 µm (d); 50 µm (e–h); 15 µm (i–m).

Material examined

China, Yunnan Province, Kunming, on a dead stem of Hedychium coronarium (Zingiberaceae), 20 July 2016, D. S. Tennakoon, KDS12 (HFJAU10332, holotype); ibid. 21 August 2016, KDS13 (HFJAU10333, paratype); ibid. 25 August 2016, KDS18 (HFJAU10334, paratype).

Notes

Herpotrichia zingiberacearum is isolated from the dead stem of Hedychium coronarium (Zingiberaceae). The newly-generated sequences H. zingiberacearum (LSU, SSU, ITS and tef1-α) formed a monophyletic clade closely related to H. macrotricha with 90% ML and 0.99 BYPP statistical support. Morphologically, they share similarities in having dark brown to black, setose, coriaceous ascomata, cylindrical clavate asci and hyaline, 1-septate ascospores (Mugambi and Huhndorf 2009). However, H. zingiberacearum can be distinguished from H. macrotricha in their smaller asci (85–98 × 10–14 μm vs. 115–145 × 11–13 μm) and ascospores (25–30 × 5–6 μm vs. 30–35 × 4–6 μm) (Tanaka and Hosoya 2006). On the other hand, H. zingiberacearum (HFJAU10332) differs from H. macrotricha (GKM 196N) by a comparison of the 694 nucleotides across the tef1-α gene region which shows 20 base pair differences (3.02%). This finding addresses future studies to explore the diversity of Herpotrichia species in different geographic regions and host plants, as these species have not previously been described from Zingiberaceae hosts (Table 2).

Melanomma Nitschke ex Fuckel

Notes

Melanomma was validly established by Fuckel (1870) with M. pulvis-pyrius as the type species. These species have cosmopolitan distribution worldwide and characterised in having carbonaceous ascomata and hyaline or brown, 2–3-septate ascospores (Tian et al. 2015; Crous et al. 2020). The asexual morph can be either coelomycetes or hyphomycetes (Hyde et al. 2011). Currently, 94 species are accepted in this genus (Species Fungorum 2024) and have been reported from 39 plant families (Fig. 10).

Melanomma populicola Crous & R.K. Schumach, Fungal Systematics and Evolution 6: 201 (2020)

Fig. 8

Basionym

Aposphaeria populina Died., Krypt.-Fl. Brandenburg (Leipzig) 9: 206 (1912).

Synonym

Melanomma populinum (Died.) Phukhams. & K.D. Hyde [as ‘populina’], Fungal Diversity 83: 49. 2017.

Description

Saprobic on dead stem of Fagus sylvatica L. (Fagaceae). Sexual morph: Ascomata 130–200 × 200–300 µm ( = 150 × 255 μm, n = 15), solitary or scattered, immersed, erumpent through host surface, black, multi-loculate, globose to subglobose, ostiolate. Peridium 10–15 μm wide, thick-walled, carbonaceous, composed of several layers of light brown to dark brown pseudoparenchymatous cells, cells towards the inside hyaline, arranged in a textura angularis, fusing at the outside indistinguishable from the host tissues. Hamathecium comprising numerous, 1–2 µm wide, hyaline, septate, filiform pseudoparaphyses. Asci 80–110 × 6–8 μm ( = 96 × 7 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, apically rounded, short pedicellate with furcate end, with an indistinct ocular chamber. Ascospores 13–17 × 4–5.2 μm ( = 15 × 4.8 μm, n = 35), overlapping, 1–2-seriate, ellipsoid, initially hyaline, becoming light brown at maturity, 3-septate, straight to slightly curved, slightly constricted at the septa, guttulate, smooth-walled. Asexual morph: See Crous et al. (2020).

Figure 8. 

Melanomma populicola (HFJAU10335, new host record) a appearance of ascomata on the host b close-up of ascoma c vertical section of an ascoma d peridium e pseudoparaphyses f–h asci i–n ascospores. Scale bars: 75 µm (c); 10 µm (d); 40 µm (e–h); 10 µm (i–n).

Material examined

China, Kunming, on a dead stem of Fagus sylvatica (Fagaceae), 25 July 2016, D. S. Tennakoon, KDS25 (HFJAU10335).

Known hosts

Populus canadensis, Picea abies, Quercus sp., Sorbus aucuparia (Crous et al. 2020; this study).

Known distribution

China, Germany, The Netherlands (Crous et al. 2020; this study).

Notes

Melanomma populicola was introduced by Diedicke (1912) as Aposphaeria populina, which was a phoma-like species collected from Populus canadensis. An epitype for A. populina was established by De Gruyter et al. (2013). Tibpromma et al. (2017) synonymised A. populina under Melanomma and erected as M. populina. Subsequently, this was validated by Crous et al. (2020) and erected as M. populicola. Morphological characteristics of our collection (HFJAU10335) fit well with the M. populicola in having black, globose to subglobose ascomata, cylindrical, apically rounded asci and ellipsoid, 3-septate, light brown ascospores. In addition, there are overlapping size ranges of asci (80–110 × 6–8 μm vs. 91–106 × 6.5–7.5 μm) and ascospores (13–17 × 4–5.2 μm vs. 15.1 × 5 μm) (Crous et al. 2020). Multi-gene phylogeny also shows that our collection groups with M. populicola isolates in a 99% ML and 0.99 BYPP supported clade. Therefore, we introduce our collections as a new host record of M. populicola from Fagus sylvatica.

Geographical distribution and host associations of melanommataceous species

Based on the data collected, it appears that the members of Melanommataceae are widely distributed around the world, comprising subtropical, tropical and temperate regions, such as Austria, Australia, Brazil, Canada, Finland, Germany, Japan, India, Thailand, Papua New Guinea, South Africa, Ukraine and the United States). The highest number of species have been reported from the United States (48 species). This is followed by China (35 species), Italy (30 species), India (25 species), Germany (23 species) and the United Kingdom (22 species) (indicated by red areas, Fig. 9). Moreover, Japan (17 species), Brazil (16 species), Ukraine (16 species), Canada (15 species), Kazakhstan (13 species), Poland (13 species) and Austria (11 species) are indicated by green areas (Fig. 9). Twenty-four countries have the species number range between 3 and 10, for instance Argentina, Russia and Sweden (10 species), Uzbekistan (9 species), Australia, France and Switzerland (8 species), Chile (7 species), Luxembourg, New Zealand, the Philippines and Thailand (6 species), Finland (5 species), Belgium and South Africa (4 species) and three species in nine countries (Azerbaijan, Czechia, Denmark, Georgia, Hungary, Papua New Guinea, Slovakia, Spain and Sri Lanka) (indicated by blue areas, Fig. 9). In addition, 36 countries reported two or a single species (indicated by yellow areas, Fig. 9). This may be because those countries have very limited taxonomic investigations on melanommataceous species. Consequently, it would be important to conduct more comprehensive collections, carry out more taxonomic studies and identify the species in those poorly-explored countries (e.g. African continent). In Asia, the highest number of melanommataceous species have reported from China, India and Thailand (Fig. 9), while Austria, Germany, Italy, Poland and the United Kingdom in Europe also have high numbers.

Figure 9. 

Distribution of so far reported species in Melanommataceae worldwide. Colour gradient shows the number of recorded species from lowest (yellow) to highest (red) and no records (grey).

Some species exist over multiple continents, while others appear to have limited distribution and are currently reported from one or a few countries (Fig. 9). Nine species of Melanommataceae are intercontinental and occur on more than five countries. Some highly diverse species include Aposphaeria pulviscula, Camposporium cambrense, C. pellucidum, Herpotrichia herpotrichoides, H. nigra, H. quinqueseptata, Melanomma pulvis-pyrius, M. rhododendri and Seifertia azaleae (Table 2). As well, nine species have been reported from four countries (Byssosphaeria jamaicana, By. rhodomphala, Camposporium antennatum, Herpotrichia macrotricha, Melanomma glumarum, M. subdispersum, Petrakia echinata, Phragmocephala elliptica and Phragmotrichum chailletii) and nine species from three countries (Aposphaeria eragrostidis, A. freticola, A. mediella, Camposporium hyderabadense, Melanomma medium, M. populicola, Navicella pileata, Phragmocephala stemphylioides and Phragmotrichum rivoclarinum). In contrast, the distribution of 313 species has been limited to two or to one country, based on the current information. We believe that the distribution of those species may be much higher with future collections and taxonomical investigations.

When focusing on the host association of melanommataceous species, most of have been discovered in decaying wood or submerged woody substrates (e.g. Aposphaeria corallinolutea, A. rudis, Asymmetricospora calamicola, Bertiella ellipsoidea, Byssosphaeria juniperi, Calyptronectria argentinensis, Camposporium chinense, C. marylandicum, Herpotrichia alpincola, Marjia uzbekistanica, Melanomma dinghuense, Phragmocephala garethjonesii and Sarimanas pseudofluviatile) (Table 2). Some species have been recorded from decaying leaves (e.g. Bertiella fici, Byssosphaeria musae, Phragmocephala elegans). As well, it is noteworthy to mention that some species have been recorded from soil (e.g. Herpotrichia gelasinosporoides, H. striatispora and Pleotrichocladium opacum). In addition, Aposphaeria ramalinae has been collected from a lichen species (Ramalina implectens) in France (Pitard and Harmand 1911) and Exosporiella fungorum from a mushroom species (Thelephora sp. fibre vase or earth fan mushroom) in Sweden (Karsten 1892).

According to the host associations, some species of Melanommataceae are highly diverse and have been collected from more than 10 host species (e.g. Camposporium antennatum, C. japonicum, C. pellucidum, Herpotrichia macrotricha, H. nigra and Melanomma pulvis-pyrius), while some are from more than five host species (e.g. Camposporium cambrense, Herpotrichia herpotrichoides, Melanomma populicola and Phragmocephala elliptica). The highest number of melanommataceous species have reported from the plant family Fagaceae (32 species). The most common Fagaceae hosts are Fagus spp. (e.g. F. crenata, F. sylvatica) and Quercus species (e.g. Q. germana). This is followed by Fabaceae (24 species), Rosaceae (23 species), Salicaceae (22 species), Poaceae (19 species), Pinaceae and Arecaceae (18 species), Sapindaceae (16 species), Betulaceae (15 species), Amaranthaceae (12 species), Asteraceae (10 species), Ericaceae (9 species), Cupressaceae, Euphorbiaceae and Lauraceae (7 species), Rutaceae (6 species) and Malvaceae, Oleaceae and Pandanaceae (5 species) (Fig. 10). In addition, a range of 2–5 species have recorded from 30 plant families and single melanommataceous species have reported from 35 plant families. Thus, up to date, the species of the family have been reported from 82 plant families. However, some species have been collected from unidentified hosts and, thus, their host association remained unresolved (e.g. Aposphaeria anomala, A. pakistanica, Bertiella ellipsoidea, B. gelatinosa, Camposporium appendiculatum and C. chinense). The host specificity for most of the species has not yet been clarified, as they have been recorded from various plant families. Though, it was noted by Jaklitsch and Voglmayr (2017) that Praetumpfia obducens may be host specific to Fraxinus species.

Figure 10. 

The melanommataceous species distribution through plant host families.

Discussion

The members of Melanommataceae have been well-studied in last two decades, leading to many exciting discoveries (17 new genera and 76 new species), which may be mostly linked to the progress made in the DNA sequence data. The initial classification of these species was primarily based on morphological characteristics (e.g. globose or depressed ascomata, trabeculate pseudoparaphyses, fissitunicate asci, pigmented ascospores) associated with hand drawings (Saccardo 1883; Winter 1885; Morgan 1904; Spegazzini 1909; Shear 1941). However, morphology-based identification is challenging since most species can share similar characteristics, which may lead to misinterpretations. Thus, a combination of DNA-based molecular phylogeny and morphological traits has become a widely accepted tool for current fungal classifications (Tian et al. 2015; Hashimoto et al. 2017; Jaklitsch and Voglmayr 2017; Purahong et al. 2017; Kularathnage et al. 2022). Mostly, the large subunit (28S, LSU), small subunit (18S, SSU), internal transcribed spacers (ITS1–5.8S–ITS2), translation elongation factor 1 gene (tef1-α) and RNA polymerase second largest subunit (rpb2) molecular markers have been used in current phylogenetic analyses of Melanommataceae (Tian et al. 2015; Tennakoon et al. 2018; Pem et al. 2019; Kularathnage et al. 2022).

Melanommataceae genera are highly varied, both morphologically and phylogenetically. Of them, some are highly diverse with numerous species (e.g. Aposphaeria: 83 species, Melanomma: 81 species, Herpotrichia: 61 species, Camposporium: 25 species, Byssosphaeria: 18 species and Petrakia: 10 species), while some have fewer species (Phragmocephala: 9 species, Bertiella: 6 species, Navicella: 5 species, Phragmotrichum: 5 species, Uzbekistanica: 4 species, Calyptronectria: 3 species, Pseudostrickeria: 3 species, Seifertia: 3 species, Marjia: 2 species, Muriformistrickeria: 2 species, Sarimanas: 2 species, and Tumularia: 2 species). In addition, some genera are monotypic and need more collections for their expansion (e.g. Alpinaria, Asymmetricospora, Bicrouania, Dematiomelanomma, Exosporiella, Fusiconidium, Mamillisphaeria, Melanocamarosporioides, Melanocamarosporium, Melanocucurbitaria, Melanodiplodia, Monoseptella, Neobyssosphaeria, Pleotrichocladium, Praetumpfia, Pseudobyssosphaeria and Xenostigmina). However, it is noteworthy to mention that some genera currently lack molecular data and, thus, their phylogenetic position is uncertain (e.g. Asymmetricospora, Bicrouania, Calyptronectria, Exosporiella, Mamillisphaeria and Navicella) (Hongsanan et al. 2020). As well, only two species of Aposphaeria have molecular data out of the 84 species listed in the Species Fungorum (2024). Consequently, to resolve their phylogenetic placements, further collections and investigations are essentially required.

In this study, we introduced two new species and three new host records collected from China and Thailand. The new species, Byssosphaeria poaceicola and Herpotrichia zingiberacearum can be distinguished from related species in their morphology and DNA molecular data. The morphological characteristics of the new host records strongly tally with their type species and phylogeny analyses also provide evidence for their placements. These new host records also demonstrate their adaptability to a broad range of habitats and there could be many more. Thus, many fungal species and host associations are waiting for us and we should undertake further explorations.

Acknowledgements

Dr. Yang Gao is thanked for his support in depositing the herbarium specimens. The authors would like to thank Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for his guidance on the fungal nomenclature.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

No funding was reported.

Author contributions

Conceptualisation, D.S.T. N.I.d.S. and D.M.H; methodology, D.S.T. and N.I.d.S; software, D.S.T. and N.I.S.; validation, K.M.T., N.I.d.S., H.Y.S., N.S., F.S.C. and D.M.H; formal analysis, D.S.T. and N.I.d.S; investigation, K.M.T., N.I.d.S., H.Y.S., N.S., F.S.C. and D.M.H; resources, D.S.T; data curation, D.S.T. and N.I.d.S; writing – original draft preparation, D.S.T., K.M.T., N.I.d.S., N.S. and D.M.H; writing – review and editing, K.M.T., N.I.d.S., H.Y.S., N.S., F.S.C. and D.M.H; supervision, D.M.H; project administration, D.M.H; funding acquisition, D.M.H. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Danushka S. Tennakoon https://orcid.org/0000-0003-2306-1255

Kasun M. Thambugala https://orcid.org/0000-0002-6210-0504

Nimali I. de Silva https://orcid.org/0000-0002-1577-280X

Nakarin Suwannarach https://orcid.org/0000-0002-2653-1913

Dian-Ming Hu https://orcid.org/0000-0002-4750-2871

Data availability

All of the data that support the findings of this study are available in the main text.

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