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Phytotaxa 609 (1): 017–044
https://www.mapress.com/pt/
Copyright © 2023 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Sajeewa Maharachchikumbura: 31 Jul. 2023; published: 17 Aug. 2023
https://doi.org/10.11646/phytotaxa.609.1.3
17
Isolation and characterization of novel Dothideomycetes species from forest soils
in Chiang Rai and Krabi (Thailand): additions to the diversity of Curvularia and
Verruconis
W. A. ERANDI YASANTHIKA1,2,3,9, DANUSHKA S. TENNAKOON4,5,10, ANTONIO R. GOMES DE FARIAS2,11,
K. W. THILINI CHETHANA1,2,12*, D. JAYARAMA BHAT6,7,13 & DHANUSHKA N. WANASINGHE8,14*
1 School of Science, Mae Fah Luang University, Chiang Rai, 57100, Thailand
2 Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, 57100, Thailand
3 Innovative Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Guang Dong Province, People’s Republic of
China
4 Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
5 Research Center of Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai 50200, Thailand
6 Department of Botany and Microbiology, College of Science, King Saud University, P.O. Box 2455, Riyadh 11451, Saudi Arabia
7 Formerly, Department of Botany, Goa University, Goa, India; House No. 128/1-J, Azad Co-Op Housing Society, Curca, P.O. Goa
Velha-403108, India
8 Center for Mountain Futures, Kunming Institute of Botany, Chinese Academy of Science, Honghe County 654400, Yunnan, People’s
Republic of China
9
�
eyasanthika@gmail.com; https://orcid.org/0000-0002-3757-3801
10
�
danushkasandaruwanatm@gmail.com; https://orcid.org/0000-0003-2306-1255
11
�
rfariasagro@gmail.com; https://orcid.org/0000-0003-4768-1547
12
�
kandawatte.thi@mfu.ac.th; https://orcid.org/0000-0002-5816-9269
13
�
bhatdj@gmail.com; https://orcid.org/0000-0002-3800-5910
14
�
dnadeeshan@gmail.com; https://orcid.org/0000-0003-1759-3933
*Corresponding authors:
�
dnadeeshan@gmail.com,
�
kandawatte.thi@mfu.ac.th
Abstract
Fungi are adapted to diverse environments, where the forest soils have complex fungal communities with a wide range of
lifestyles. This study aims to explore and isolate Dothideomycetes from the forest soils in Thailand. Sampling sites were
located in Chiang Rai and Krabi provinces. Cultures were obtained through soil dilution series, and the strains were subjected
to morphological observations and multigene phylogenetic analyses for identification. Maximum likelihood and Bayesian
inference analyses were conducted to clarify their phylogenetic affinities using partial nuclear ribosomal DNA (ITS, LSU,
and SSU) and the protein-coding genes (tef1-α and GADPH). Herein, two new species (Curvularia chiangraiensis sp.
nov. and Verruconis soli sp. nov.) and two new records (C. chiangmaiensis and V. thailandica) are described based on
the morphological and phylogenetic evidence. Curvularia chiangraiensis sp. nov. is distinguished by its relatively smaller
conidia and the presence of sympodial proliferation of conidiogenesis and Verruconis soli sp. nov. exhibits micronematous
conidiophores and obovoid conidia that transform into sub-cylindrical or ellipsoidal shapes, and becoming 1-septate when
mature. These unique characteristics set them apart from closely related taxa. The newly described taxa have been subjected
to a thorough comparison with closely related species, enabling a comprehensive analysis. The study offers detailed
descriptions and includes high-quality micrographs, with the aim of providing a comprehensive understanding of these
newly identified taxa
Keywords: 2 new records, 2 new species, Ascomycota, Pleosporales, Venturiales
Introduction
Fungi are diverse microorganisms that play important functional roles in soil ecosystems (Tedersoo et al. 2017, 2021,
Shi et al. 2022). In particular, they regulate nutrient cycling, disease suppression, and water dynamics, which promote
plant growth (Guo et al. 2020). The physical, chemical, and biological characteristics of soil are major drivers of fungal
distribution, resulting in various global taxonomic groups (Guo et al. 2020, Tedersoo et al. 2021, Coleine et al. 2022).
YASANTHIKA ET AL.
18 • Phytotaxa 609 (1) © 2023 Magnolia Press
As a result of the complexity of belowground ecosystems, the taxonomy of macro- and micro-organisms is challenging
(Tedersoo et al. 2014). There is a plethora of studies in the world on above-ground biodiversity compared to the
belowground (Guerra et al. 2020). For example, soils in Thailand are characterized by tropical climatic conditions, yet
studies on soil biota are limited (Osono et al. 2009, Amma et al. 2018). Despite the challenges, many researchers have
made considerable effort to resolve the taxonomy of soil-inhabiting fungi in Thailand at their higher and species levels
(Amma et al. 2018, Mongkolsamrit et al. 2020, Nundaeng et al. 2021).
Dothideomycetes is a dominant group in Ascomycota which includes around 1,500 genera and is expected to hold
a high diversity in belowground ecosystems (Tedersoo et al. 2014, Jayasiri et al. 2019, Hongsanan et al. 2020a, b,
Das et al. 2021). Curvularia and Verruconis are common soil genera, yet only a few studies are available about them
in Thailand soils (Marin-Felix et al. 2017a, b, Hernández-Restrepo et al. 2020). Curvularia is a species-rich genus
in Pleosporaceae (Pleosporales, Dothideomycetes), and currently, 185 species are listed in Species Fungorum 2023
(Wijayawardene et al. 2022). Boedijn (1933) established this genus, typified by C. lunata. Molecular analyses are
essential in Curvularia species delimitation, as most of them share similar morphological characteristics (Manamgoda
et al. 2015). Curvularia species have a cosmopolitan distribution and are recorded from plants, terrestrial and aquatic
substrates, and air (Marin-Felix et al. 2020). They can be epiphytes, endophytes, pathogens (animal, human, and
plants) and/or saprobes (Tadych et al. 2012, Marin-Felix et al. 2017b, Ferdinandez et al. 2021) and usually occur in
their asexual state, but the sexual pseudocochliobolus state has been observed (Wijayawardene et al. 2021b) . Some
have been recorded as human pathogens (i.e., Curvularia spicifera and C. lunata) (Carter & Boudreaux 2004, de Hoog
et al. 2011, da Cunha et al. 2013, Madrid et al. 2014, Marin-Felix et al. 2020). In addition, Curvularia aeria, C. lunata,
and C. eragrostidis are plant pathogens that cause severe diseases to cash crops (Adikaram 2020, Ferdinandez et al.
2021). Curvularia borreriae, C. brachyspora, C. inaequalis, C. nicotiae, C. paraverruculosa, C. pseudointermedia,
C. pseudoprotuberata, C. shahidchamranensis, C. spicifera and C. subpapendorfii, have been reported from different
soil-based environments worldwide, such as cultivated, volcanic ash, and crude oil contaminated soils (Manamgoda
et al. 2015, Marin-Felix et al. 2017a, b, 2020, Hernandez-Restrepo et al. 2018). Curvularia alcornii, C. asianensis,
C. borreriae, C. hawaiiensis, C. hominis, C. lunata, and C. verruculosa strains have been isolated previously from
various hosts and substrates (i.e. cow and deer dung, Dactyloctenium aegyptium, Eleusine indica, and Zea mays) from
Thailand (Jeamjitt et al. 2006, Marin-Felix et al. 2017b, Ahebwa et al. 2020).
Zhang et al. (2011) established Sympoventuriaceae (Venturiales, Dothideomycetes) and typified by Sympoventuria.
Samerpitak et al. (2014) erected Verruconis to accommodate V. gallopava (≡ Ochroconis gallopava). Currently, ten
species are accepted in this genus (Species Fungorum 2023, Wijayawardene et al. 2022). Most Verruconis members
have been reported as saprobes or pathogens (on humans and animals) in terrestrial or aquatic environments. Verruconis
comprises several thermophilic species (i.e., V. gallopava) that can survive at 35–42 °C (Qiao et al. 2019, Shen et al.
2020). These species can be found in soil, hot springs, industrial effluents, plants, and humans (Zhang et al. 2018,
Huanraluek et al. 2019, Shen et al. 2020).
High-throughput sequencing methods are widely applied in soil-based fungal taxonomy studies (Yasanthika et al.
2022). Numerous soil-associated dothideomycetous taxa have been recovered from the tropics, though, the cultured
fraction contains only a few (Elnaiem et al. 2021, Tedersoo et al. 2021) due to limitations in culturing techniques.
However, cultures are important in species identification and biotechnology (Chethana et al. 2020, 2021, Wijayawardene
et al. 2021a, Yasanthika et al. 2022). This study aimed to investigate soil-inhabiting Dothideomycetes from tropical
forest soils in Thailand. Herein, we provide morpho-molecular illustrations to introduce two new Dothideomycetes
species (C. chiangraiensis and Verruconis soli) and two soil records (C. chiangmaiensis and V. thailandica) from
tropical forest soils in Thailand.
Materials & methods
Sampling, morphological studies, and isolation
Soil samples were collected from tropical forest areas of Chiang Rai and Krabi Provinces in Thailand, up to 10 cm
depths from the soil surface using a sterilized soil auger (Yasanthika et al. 2021). After collection, they were packed
in sterilized bags and stored in a box with ice during transportation. Upon reaching the laboratory, the samples were
stored at 4 °C until further analyses. Fungal isolations were done by the soil dilution method (up to 10-5), as described
in Yasanthika et al. (2020, 2022). Further, sub-culturing was done to obtain pure fungal colonies. Isolated strains were
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 19
incubated at 25 °C under day and night conditions for two to three months for culture sporulation. Macro- and micro-
morphological characteristics were examined using the sporulated cultures. Conidial attachments and mycelia were
picked using a sterilized needle and placed in a drop of distilled water on a slide for micro-morphological examinations.
Photomicrographs of fungal structures were captured and recorded using an OLYMPUS SZ61 compound microscope
and Canon EOS 600D digital camera mounted on a Nikon ECLIPSE 80i compound microscope. Measurements were
made with the Tarosoft (R) Image Frame Work program, and images were processed with Adobe Photoshop CS6
extended version 13.0.1 (Adobe Systems, USA). Dried cultures were prepared using agar with 2.5% glycerol as
herbarium specimens. Living cultures were deposited in the Mae Fah Luang University Culture Collection (MFLUCC)
and dried cultures at Mae Fah Luang University Herbarium (Herb. MFLU) Chiang Rai, Thailand. Faces of Fungi and
Index Fungorum numbers were obtained as described in Jayasiri et al. (2015) and Index Fungorum (2023), and the data
was submitted to the Greater Mekong Subregion database (Chaiwan et al. 2021).
DNA extraction, PCR amplification, and sequencing
Pure fungal colonies grown in PDA (Potato Dextrose Agar) media for four weeks at 25 °C were scraped, and 50–100 mg
of the mycelia were subjected to the total genomic DNA extraction using the Biospin Fungus Genomic DNA Extraction
Kit (BioFlux®) (Hangzhou, P. R. China) according to the manufacturer’s instructions. Partial DNA sequences of
ribosomal coding genes; internal transcribed spacer region of ribosomal DNA (ITS), large subunit nuclear ribosomal
DNA region (LSU), and small subunit nuclear ribosomal DNA (SSU) genomic regions, and the protein-coding gene
regions, translation elongation factor 1-α (tef1-α) and glyceraldehyde-3-phosphate dehydrogenase (GAPDH) genes
were amplified with the primer pairs, ITS5/ITS4, LR0R/LR5, NS1/NS4, EF1-983F/EF1-2218R and gpd1/gpd2
respectively (Huanraluek et al. 2019, Qiao et al. 2019, Ferdinandez et al. 2021). PCR reactions were composed of
12.5 μL of 2 × Power Taq PCR MasterMix (a premixed and ready-to-use solution, including 0.1 Units/μL Taq DNA
Polymerase, 500 μM dNTP Mixture each (dATP, dCTP, dGTP, dTTP), 20 mM Tris-HCl pH 8.3, 100 Mm KCl, 3 mM
MgCl2, stabilizer and enhancer), 1 μL of each primer (10 μM), 1 μL genomic DNA extract and 8.5 μL double distilled
water. PCR thermal cycles of DNA amplification of LSU, ITS, GAPDH, and tef1-α regions were performed following
Shen et al. (2020) and Ferdinandez et al. (2021). PCR products were purified and sequenced by Qingke Company,
Kunming City, Yunnan Province, P. R. China.
Phylogenetic analyses
Newly generated sequences were searched in the BLAST tool against the GenBank database (https://blast.ncbi.nlm.
nih.gov/Blast.cgi) to determine the closely related taxa. Other sequences used in the analyses (TABLES 1 and 2) were
obtained from GenBank based on recently published data (Huanraluek et al. 2019, Qiao et al. 2019, Shen et al. 2020,
Ferdinandez et al. 2021), and the sequences retrieved from the GenBank. Sequence alignments were performed in
MAFFT v. 7.036 (Katoh et al. 2019) with default settings and refined where necessary using BioEdit v. 7.0.5.2 (Hall
1999). Evolutionary models were calculated using MrModeltest v. 2.3 (Nylander 2004) under the Akaike Information
Criterion (AIC). Bayesian analyses were conducted with MrBayes v. 3.1.2 (Huelsenbeck & Ronqvist 2001) to evaluate
Bayesian posterior probabilities (BYPP) (Rannala & Yang 1996, Zhaxybayeva & Gogarten 2002) by Markov Chain
Monte Carlo sampling (MCMC). For each dataset, six simultaneous MCMCs were run for 6,000,000 and 2,000,000
generations for Curvularia and Sympoventuriaceae, respectively, with trees sampled at every 1000th generation in both
scenarios. The first 25% of generated trees were discarded, and the remaining trees were used to calculate posterior
probabilities (BYPP) for the majority rule consensus tree (Cai et al. 2006). The BYPP values greater than 0.95 are
given above each node in FIGURE 1 and 2. Maximum likelihood (ML) trees were generated using RAxML-HPC2 on
XSEDE (8.2.8) (Stamatakis 2014) in the CIPRES Science Gateway platform (Miller et al. 2012), applying GTR+I+G
as the best fit model of evolution. ML bootstrap values (ML) equal to or greater than 60% are given above each node
in FIGURE. 1 and 2. Phylograms were visualized with FigTree v1.4.0 (Rambaut 2012) and reorganized in Microsoft
power-point (2007) and Adobe Illustrator® CS5 (Version 15.0.0, Adobe®, San Jose, CA). Newly generated nucleotide
sequences were deposited in the GenBank.
YASANTHIKA ET AL.
20 • Phytotaxa 609 (1) © 2023 Magnolia Press
TABLE 1. Species, strain codes, and GenBank accession numbers of the taxa used in the phylogenetic analyses of
Curvularia.
Taxon Culture accession number ITS GAPDH tef1-α
Bipolaris maydis CBS 136.29TKJ909769 KM034845 KM093793
B. panici-miliacei CBS 199.29TKJ909773 KM042896 KM093788
Curvularia aeria CBS 294.61THF934910 HG779148 NA
C. affinis CBS 154.34TKJ909780 KM230401 KM196566
C. ahvazensis CBS 144673TKX139029 MG428693 MG428686
C. akaii CBS 318.86TLT631340 LT715797 NA
C. akaiiensis BRIP 16080TKJ415539 KJ415407 KJ415453
C. alcornii MFLUCC 10-0703TJX256420 JX276433 JX266589
C. americana UTHSC08-3414THE861833 HF565488 NA
C. andropogonis CBS 186.49TLT631354 LT715835 NA
C. annelliconidiophori CGMCC3.19352T MN215641 MN264077 MN263935
C. arcana CBS 127224TMN688801 MN688828 MN688855
C. asiatica MFLUCC 10-0711TJX256424 JX276436 JX266593
C. australiensis BRIP 12044TKJ415540 KJ415406 KJ415452
C. australis BRIP 12521TKJ415541 KJ415405 KJ415451
C. austriaca CBS 102694TMN688802 MN688829 MN688856
C. bannonii BRIP 16732TKJ415542 KJ415404 KJ415450
C. beasleyi BRIP 10972TMH414892 MH433638 MH433654
C. beerburrumensis BRIP 12942TMH414895 MH433634 MH433657
C. boeremae IMI 164633TMH414911 MH433641 NA
C. borreriae CBS 859.73TLT631355 LT715838 NA
C. bothriochloae BRIP 12522TKJ415543 KJ415403 KJ415449
C. brachyspora CBS 186.50THG778983 KM061784 KM230405
C. buchloes CBS 246.49TKJ909765 KM061789 KM196588
C. cactivora CBS 580.74 MN688803 MN688830 MN688857
C. canadensis CBS 109239TMN688804 MN688831 MN688858
C. caricae-papayae CBS 135941TLT631350 LT715816 –
C. chiangmaiensis CPC 28829TMF490814 MF490836 MF490857
C. chiangmaiensis LC12044 MN215651 MN264087 MN263945
C. chiangmaiensis LC12030 MN215650 MN264086 MN263944
C. chiangmaiensis MFLUCC 22-0084 OP564987 OP859014 OP859018
C. chiangraiensis MFLUCC 22-0091TOP581428 OP859013 OP859017
C. chlamydospora UTHSC 07-2764THG779021 HG779151 NA
C. chonburiensis MFLUCC 16-0375 T MH275055 MH412747 NA
C. clavata BRIP:61680 KU552205 KU552167 KU552159
C. coatesiae BRIP 24261TMH414897 MH433636 MH433659
C. coicicola HSAUP 990901 AB453880 NA NA
C. coicis CBS 192.29THF934917 HG779130 JN601006
C. coimbatorensis SZMC 22225TMN628310 MN628306 MN628302
C. colbranii BRIP 13066TMH414898 MH433642 MH433660
......continued on the next page
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 21
TABLE 1. (Continued)
Taxon Culture accession number ITS GAPDH tef1-α
C. comoriensis CBS 110673 LT631357 LT715841 NA
C. crassiseptata CBS 503.90TLT631310 LT715882 MN688859
C. crustacea BRIP 13524 TKJ415544 KJ415402 KJ415448
C. curculiginis YZU 181230 MK507796 MK507794 MK507795
C. cymbopogonis CBS 419.78THG778985 HG779129 NA
C. dactyloctenicola CPC 28810TMF490815 MF490837 MF490858
C. dactyloctenii BRIP 12846TKJ415545 KJ415401 KJ415447
C. determinata CGMCC 3.19340TMN215653 MN264088 MN263947
C. deightonii CBS 537.70TLT631356 LT715839 NA
C. eleusinicola USJCC-0005TMT262877 MT393583 MT432925
C. elliptiformis CGMCC 3.19351TMN215656 MN264091 MN263950
C. ellisii CBS 193.62TJN192375 JN600963 JN601007
C. eragrosticola BRIP 12538TMH414899 MH433643 MH433661
C. eragrostidis CBS 189.48THG778986 HG779154 NA
C. falsilunata CGMCC 3.19329TMN215660 MN264093 MN263954
C. flexuosa CGMCC3.19447TMN215663 MN264096 MN263957
C. geniculata CBS 187.50TKJ909781 KM083609 KM230410
C. gladioli CBS 210.79 LT631345 LT715802 NA
C. graminicola BRIP 23186aTJN192376 JN600964 JN601008
C. guangxiensis CGMCC3.19330TMN215667 MN264100 MN263961
C. gudauskasii DAOM 165085 AF071338 AF081393 NA
C. harveyi BRIP 57412TKJ415546 KJ415400 KJ415446
C. hawaiiensis BRIP 11987TKJ415547 KJ415399 KJ415445
C. heteropogonicola BRIP 14579TKJ415548 KJ415398 KJ415444
C. heteropogonis CBS 284.91TKJ415549 JN600969 JN601013
C. hominis UTHSC 09-464THG779011 HG779106 NA
C. homomorpha CBS 156.60TJN192380 JN600970 JN601014
C. inaequalis CBS 102.42TKJ922375 KM061787 KM196574
C. intermedia CBS 334.64 HG778991 HG779155 NA
C. iranica IRAN 3487CTMT551122 MN266487 MN266490
C. ischaemi CBS 630.82THG778992 HG779131 NA
C. kenpeggii BRIP 14530TMH414900 MH433644 MH433662
C. khuzestanica CBS 144736TMH688044 MH688043 NA
C. kusanoi CBS 137.29TJN192381 LT715862 KM196592
C. lamingtonensis BRIP 12259TMH414901 MH433645 MH433663
C. lonarensis CBS 140569TKT315408 KY007019 NA
C. lunata CBS 730.96TJX256429 JX276441 JX266596
C. lycopersici Strain 11 KY883347 KY883345 NA
C. manamgodae CGMCC3.19446TMN215677 MN264110 MN263971
C. malina CBS 131274TJF812154 KP153179 KR493095
......continued on the next page
YASANTHIKA ET AL.
22 • Phytotaxa 609 (1) © 2023 Magnolia Press
TABLE 1. (Continued)
Taxon Culture accession number ITS GAPDH tef1-α
C. mebaldsii BRIP 12900TMH414902 MH433647 MH433664
C. micropus CBS 127235THE792934 LT715859 NA
C. microspora GUCC 6272TMF139088 MF139106 MF139115
C. miyakei CBS 197.29TKJ909770 KM083611 KM196568
C. mosaddeghii IRAN 3131CTMG846737 MH392155 MH392152
C. muehlenbeckiae CBS 144.63THG779002 HG779108 NA
C. nanningensis HGUP11005TMH885321 MH980005 MH980011
C. neergaardii BRIP 12919TKJ415550 KJ415397 KJ415443
C. neoindica IMI 129790TMH414910 MH433649 MH433667
C. nicotiae BRIP 11983TKJ415551 KJ415396 KJ415442
C. nodosa CPC 28800TMF490816 MF490838 MF490859
C. nodulosa CBS 160.58 JN601033 JN600975 JN601019
C. oryzae CBS 169.53TKP400650 HG779156 KM196590
C. oryzae-sativae CBS 127725TMN688808 MN688835 MN688863
C. ovariicola BRIP 15882 JN192384 JN600976 JN601020
C. pallescens CBS 156.35TKJ922380 KM083606 KM196570
C. palmicola MFLUCC 14-0404TMF621582 NA NA
C. pandanicola MFLUCC 15-0746TMH275056 MH412748 MH412763
C. panici OKI-1 AB164703 NA NA
C. panici-maximi USJCC-0006TMN044757 MN053040 MN053009
C. paraverruculosa FMR 17656 TLR736641 LR736646 LR736649
C. patereae CBS 198.87TMN688810 MN688837 MN688864
C. penniseti CBS 528.70 MH859833 LT715840 NA
C. perotidis CBS 350.90THG778995 HG779138 KM230407
C. petersonii BRIP 14642TMH414905 MH433650 MH433668
C. phaeospara CGMCC 3.19448TMN215686 MN264118 MN263980
C. pisi CBS 190.48TKY905678 KY905690 KY905697
C. plantarum CGMCC3.19342 MN215688 MN264120 MN263982
C. platzii BRIP 27703bTMH414906 MH433651 MH433669
C. polytrata CGMCC3.19338TMN215691 MN264123 MN263984
C. portulacae BRIP 14541TKJ415553 KJ415393 KJ415440
C. prasadii CBS 143.64TKJ922373 KM061785 KM230408
C. protuberans CGMCC 3.19360TMN215693 MN264125 MN263986
C. protuberata CBS 376.65TKJ922376 KM083605 KM196576
C. pseudobrachyspora CPC 28808TMF490819 MF490841 MF490862
C. pseudoclavata CBS 539.70TMN688817 MN688844 MN688869
C. pseudoellisii CBS 298.80TMN688818 MN688845 MN688870
C. pseudointermedia CBS 553.89TMN688819 MN688846 MN688871
C. pseudointermedia USJCC-0003 MT262876 MT432927 MT432924
C. pseudolunata UTHSC09-2092THE861842 HF565459 NA
......continued on the next page
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 23
TABLE 1. (Continued)
Taxon Culture accession number ITS GAPDH tef1-α
C. pseudoprotuberata CBS 385.69TMN688821 MN688848 MN688873
C. pseudorobusta UTHSC 08-3458 HE861838 HF565476 NA
C. radici-foliigena CGMCC 3.19328TMN215695 MN264127 MN263988
C. radicicola CGMCC 3.19327TMN215699 MN264131 MN263992
C. ravenelii BRIP 13165TJN192386 JN600978 JN601024
C. reesii BRIP 4358TMH414907 MH433637 MH433670
C. richardiae BRIP 4371TKJ415555 KJ415391 KJ415438
C. robusta CBS 624.68TKJ909783 KM083613 KM196577
C. rouhanii CBS 144674TKX139030 MG428694 MG428687
C. ryleyi BRIP 12554TKJ415556 KJ415390 KJ415437
C. saccharicola CGMCC MN215701 MN264133 MN263994
C. sacchari-officinarum CGMCC3.19331TMN215705 MN264137 MN263998
C. senegalensis CGMCC 3.9578TEF175940 NA NA
C. shahidchamranensis IRAN 3133C MH550084 MH550083 NA
C. sichuanensis BN9 MH483998 NA NA
C. siddiquii CBS 196.62TMN688823 MN688850 NA
C. simmonsii USJCC-0002TMN044753 MN053011 MN053005
C. soli CBS 222.96TKY905679 KY905691 KY905698
C. sorghina BRIP 15900TKJ415558 KJ415388 KJ415435
C. spicifera CBS 274.52 JN192387 JN600979 JN601023
C. sporobolicola BRIP 23040bTMH414908 MH433652 MH433671
C. subpapendorfii CBS 656.74TKJ909777 KM061791 KM196585
C. suttoniae FMR 10992 THE861828 HF565479 LR736651
C. tamilnaduensis SZMC 22226TMN628311 MN628307 MN628303
C. thailandicum MFLUCC 15-0747 MH275057 MH412749 MH412764
C. tribuli CBS 126975TMN688825 MN688852 MN688875
C. trifolii ICMP 6149 KM230395 KM083607 JX266600
C. tripogonis BRIP 12375TJN192388 JN600980 JN601025
C. tropicalis BRIP 14834TKJ415559 KJ415387 KJ415434
C. tsudae ATCC 44764TKC424596 KC747745 KC503940
C. tuberculata CBS 146.63TJX256433 JX276445 JX266599
C. umbiliciformis CGMCC 3.19346TMN215711 MN264142 MN264004
C. uncinata CBS 221.52THG779024 HG779134 NA
C. variabilis CPC 28815TMF490822 MF490844 MF490865
C. verruciformis CBS 537.75 HG779026 HG779133 NA
C. verrucosa CBS 422.93 MN688826 MN688853 MN688876
C. verruculosa CBS 150.63 KP400652 KP645346 KP735695
C. vietnamensis FMR 17659 TLR736642 LR736644 LR736647
C. warraberensis BRIP 14817TMH414909 MH433653 MH433672
C. xishuangbannaensis KUMCC17-0185TMH275058 MH412750 MH412765
The newly generated sequences are indicated in bold, and type strains are indicated with T (NA =Not available)
YASANTHIKA ET AL.
24 • Phytotaxa 609 (1) © 2023 Magnolia Press
TABLE 2. Species, strain codes, and GenBank accession numbers of the taxa used in the phylogenetic analyses of
Sympoventuriaceae.
Taxon Culture accession number ITS LSU SSU
Acroconidiellina arecae NFCCI 3696 KX306747 KX306776 NA
Clavatispora thailandica MFLUCC 17-2237 MH065721 MH062960 MH062967
Echinocatena arthrinioides CBS:144202 MH107890 MH107937 NA
Fusicladium ramoconidii CBS 462.82 MH861516 MH873263 NA
Matsushimaea fasciculata GUCC 18239 MZ503725 MZ503758 MZ503651
M. monilioides CBS 143867TLT883468 LT883469 NA
Mycosisymbrium cirrhosum MTCC 12435 KR259883 KR259884 KR259885
Ochroconis anellii CBS 284.64TFR832477 KF156138 NG_062993
O. anomala CBS 131816TNA KF156137 NG_062986
O. cordanae CBS 475.80TKF156022 KF156122 NG_062985
O. crassihumicola CBS 120700 KJ867429 KJ867430 KJ867431
O. humicola CBS 116655THQ667521 KF156124 KF156068
Scolecobasidiella avellanea CBS 772.73 NA EF204505 EF204520
Scolecobasidium podocarpicola CBS 146057TMN562138 MN567645 NA
S. terreum P043 NA EU107306 EU107356
Sympoventuria capensis CBS 120136TKF156039 KF156104 NG_061163
Venturia inaequalis CBS 594.70 KF156040 MH871648 KF156093
Ve. inaequalis CBS 815.69 NA MH877728 GU296204
Veronaeopsis simplex CBS 588.66TKF156041 EU041877 NG_061164
Ver. simplex CBS 129155 NA MH878033 NA
Verruconis calidifluminalis CBS 125817 NA KF156107 KF156045
V. calidifluminalis CBS 125818TMH875239 KF156108 KF156046
V. gallopava CBS 547.81 NA KF156109 NA
V. gallopava CBS 119640 NA KF156111 KF156049
V. gallopava CBS 100437 NA KF156113 KF156050
V. gallopava CBS 863.95 NA KF156114 KF156052
V. gallopava CBS 116660 NA KF156115 KF156048
V. gallopava CBS 118.91 HQ667551 KF156110 KF156047
V. gallopava CBS 437.64THQ667553 KF282656 NG_062995
V. gallopava CBS 867.95 HQ667561 KF282657 KF156051
V. hainanensis YMF1.04165T MK244397 MK248269 MF536879
V. heveae MFLUCC 17-0092TMH602349 MH602348 NA
V. mangrovei NFCCI-4389 MN782362 MN241144 MN241147
V. mangrovei NFCCI 4390TMN782361 MN241145 NG_070311
V. panacis SYPF 8337TMF536882 MF536880 MK248267
V. pseudotricladiata YMF1.04915TMK244396 MK248270 MK248268
V. soli MFLUCC 22-0082TOP581411 OP581410 OP581426
V. soli MFLUCC 22-0090 OP581412 OP564981 OP581427
V. terricola CBS:131795 MK810925 MK810811 NA
V. thailandica CBS 145768TNR_166359 NG_068350 NA
V. thailandica GUCC 18267 MZ503753 MZ503786 MZ503679
V. thailandica MFLUCC 22-0083 OP564986 OP564978 OP565004
V. verruculosa CBS 119775 KF156014 KF282668 KF156055
Yunnanomyces pandanicola MFLUCC 17-2260TMH388369 MH376743 NG_070295
Y. phoenicis MFLUCC 19-0253 NA MK976737 MK976739
The newly generated sequences are indicated in bold, and type strains are indicated with T (NA = Not available).
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 25
Results
Phylogenetic analyses of Curvularia
Both ML and Bayesian (BYPP) analyses resulted in trees with similar topologies which did not differ significantly
(data not shown). Similarly, their topologies agree with previous studies based on multigene analyses (Marin-Felix et
al. 2020, Ferdinandez et al. 2021). A best-scoring ML tree obtained from the combined ITS-GAPDH-tef1-α alignment
comprises 158 strains of Curvularia and two outgroup taxa, Bipolaris maydis (CBS 136.29) and B. panici-miliacei
(CBS 199. 29). TABLE 3 displays all the statistical data related to the combined gene tree. Compared to the combined
gene tree, single gene trees (ITS, GAPDH, and tef1-α) showed similar topologies with low bootstrap supports for
closely related taxa. The approximate length of the combined alignment is 2127 bp, and single gene alignments are ITS:
600 bp, GAPDH: 589 bp, and tef1-α: 938 bp. Many Curvularia species are difficult to identify by the morphological
differences (Marin-Felix et al. 2017, 2020, Ferdinandez et al. 2021). Combined phylogeny is important for resolving
Curvularia, and GAPDH is considered the phylogenetically most informative marker (Marin-Felix et al. 2017, 2020,
Ferdinandez et al. 2021).
Our isolate, MFLUCC 22-0091, grouped sister to the type of C. brachyspora (CBS 186.50) and C. simmonsii
(USJCC-0002) with 77% ML and 0.98 BYPP bootstrap supports (FIGURE 1), and we introduce it as a new species,
C. chiangraiensis. Above three species clustered with C. aeria and C. homomorpha. In this cluster (clade IV), C.
brachyspora (CBS 186.50) was introduced from soils in Java, C. aeria and C. homomorpha were originally isolated
from air in Brazil and USA, respectively, and C. simmonsii (USJCC-0002) was introduced from Panicum maximum
in Sri Lanka. From these, C. brachyspora and C. aeria have been reported on Eleusine indica and Lactuca sativa in
Thailand, respectively (Marin-Felix et al. 2017, 2020, Pornsuriya et al. 2018, Ferdinandez et al. 2021).
Our other collection (MFLUCC 22-0084) formed a well-supported clade with C. chiangmaiensis strains
(CPC28829, LC12044, LC12030), with 94% ML and 1.00 BYPP support; thus, we report MFLUCC 22-0084 as C.
chiangmaiensis and this species is sister to C. lunata (CBS 730.96) shown in clade III (FIGURE 1).
Most of the Curvularia strains in our phylogenetic tree (FIGURE 1) are from type strains. Considering their host/
substrates or habitats indicated in FIGURE 1, many are recorded from plant-based substrates as saprobes, pathogens,
or endophytes. Of the remaining, 11 are from human or animal substrates, eight are from the air, one is from aquatic
habitats, and 13 are isolated from soils (FIGURE 1). In an attempt to understand the relationship between ecology and
phylogeny of selected soil inhabiting Curvularia species shown in FIGURE 1, we separated them into clades (I, II, III,
IV, and V).
Clade IV contains five type species, representing soil, air, and human substrates (soil: C. brachyspora CBS
186.50 and C. chiangraiensis F001; air: C. homomorpha CBS 156.60 and C. aeria CBS 294.61). However, these
are recorded from other substrates as well (not included in FIGURE 1), such as C. brachyspora and C. aeria isolated
from humans and plants (Torda et al. 1997, da Cunha et al. 2013, Pornsuriya et al. 2018, Ayvar-Serna et al. 2022), and
C. homomorpha isolated from plants (Manamgoda et al. 2011). However, C. simmonsii USJCC-0002 has only been
recorded from plant substrates (Ferdinandez et al. 2021).
Clade II includes four species distributed in plants, air, and soil (C. borreriae CBS 859.73, C. pallescens CBS
156.35, C. trifolii ICMP 6149 and C. coatesiae BRIP 24261). Among them, C. borreriae (CBS 859.73) and C.
pallescens (CBS 156.35) were isolated from the soil and the air, respectively. Curvularia trifolii (ICMP 6149) have
been isolated from plants and air (Almaguer et al. 2013, Marin-Felix et al. 2020), while C. coatesiae (BRIP 24261)
from plants (Laundon et al. 1971).
Clade III contains C. chiangmaiensis (Type: CPC 28829, LC12044, LC12030, and MFLUCC 22-0084) and
C. lunata (Type: CBS 730.96). Both species have strains recorded from the soil. The type of C. chiangmaiensis is
recorded from plants (Raza et al. 2019). The type strain of C. lunata is found in humans and has also been recorded
from plants (Basha et al. 2002, Marin-Felix et al. 2020). In the phylogenetic tree (FIGURE 1), there are isolates from
the soil in clade I (C. inaequalis and C. pseudoprotuberata) and V (C. shahidchamranensis and C. nicotiae), clustering
with other species that are isolated from plants and air. Based on the above, soil-associated Curvularia species have a
phylogenetic affinity to cluster and show similar ecological adaptations required to switch the trophic nature and shift
among different substrates.
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26 • Phytotaxa 609 (1) © 2023 Magnolia Press
FIGURE 1. Maximum likelihood phylogenetic tree generated of the combined ITS-GAPDH-tef1-α sequence data for Curvularia. Bootstrap
support values of maximum likelihood greater than 60% and Bayesian posterior probabilities (BYPP) greater than 0.95 are indicated above
the nodes. Newly added strains are in blue and ex-type strains are in bold. The tree is rooted to Bipolaris maydis (CBS13629P) and B.
panici-miliacei (CBS 19929). Isolated substrates/ habitat is indicated in triangles. Black: Human/ animal, green: plants, brown: soil, pink:
air, blue: aquatic habitat. unknown:empty
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 27
FIGURE 1. (Continued) Maximum likelihood phylogenetic tree generated of the combined ITS-GAPDH-tef1-α sequence data for
Curvularia.
Phylogenetic analyses of Sympoventuriaceae
The concatenated ITS, LSU, and SSU alignment contained 51 strains of Sympoventuriaceae with Venturia inaequalis
(CBS 594.70 and CBS 815.69) as the outgroup taxon. The analyses resulted in ML and Bayes trees with similar
topologies. TABLE 3 displays all the statistical data related to the combined gene tree.
Our collections (MFLUCC 22-0082 and MFLUCC 22-0090) formed a distinct lineage within clade A closer to
sub-clades B and C, with low bootstrap supports (< 60% ML and 0.95 BYPP) (FIGURE 2). Sub-clade B comprises
Verruconis hainanensis (YMF1-04165) and V. pseudotricladiata (YMF1-04915), and sub-clade C includes V. mangrovei
(NFCCI 4390, NFCCI-4389). In addition to these three sub-clades, several other Verruconis species clustered in Clade
YASANTHIKA ET AL.
28 • Phytotaxa 609 (1) © 2023 Magnolia Press
A. This topology agrees with that of Zhang et al. (2018), Qiao et al. (2019), and Hernández-Restrepo et al. (2020). Since
these studies used different gene combinations and did not include all the available species, several inconsistencies and
topological dissimilarities are observed compared to our study. Therefore, we introduce our collections (MFLUCC 22-
0082 and MFLUCC 22-0090) as new species in this genus. Another one of our isolates (MFLUCC 22-0083) grouped
with V. thailandica (CBS 145768, GUCC 18267) with 97% ML bootstrap and 1.00 BYPP, separated from V. terricola
(CBS131795), and thus identified as V. thailandica.
As shown in FIGURE 2, we tried to figure out the relationship between the ecology and phylogeny of Verruconis
species. Most of the genera in Sympoventuriaceae have been recorded from plants. Exceptionally in Verruconis, two
species were recorded from terrestrial plants (V. heveae MFLUCC 17-0092, V. paniacs SYPF 8337), one from mangrove
habitats (V. mangrovei; NFCCI 4390, NFCCI-4389), two from submerged plants (V. hainanensis YMF1.04165 and V.
pseudotricladiata YMF1.04915) and one from hot springs (V. calidifluminalis CBS 125817, CBS 125818).
The remaining members are recorded from the soil (FIGURE 2). Except for our species V. soli (MFLUCC 22-
0082 and MFLUCC 22-0090), the other three are grouped in the same clade. Besides, three strains of V. thailandica
(CBS 145768, GUCC 18267, MFLUCC 22-0083) have been isolated from China and Thailand, all of which are from
soil (Hernández-Restrepo et al. 2020, Wei et al. 2022). Verruconis found from plant parts are recorded as saprobes.
Verruconis gallopava is mostly recorded as a human/animal pathogen but has also been found in soil (not in FIGURE
2) (Geltner et al. 2015). However, Verruconis has not been recorded from the air yet. Thus, we propose that Verruconis
is adapted to shift among lifestyles, preferably to the saprobic mode and soil habitats.
TABLE 3. Maximum-likelihood (ML) and Bayesian (BI) analyses results for each sequenced dataset.
Analyses Curvularia Sympoventuriaceae
Number of taxa 158 51
Gene regions ITS, GAPDH and tef1-α ITS, LSU and SSU
Number of character positions (including gaps) 2128 2726
ML optimization likelihood value -16896.440230 -19902.347775
Distinct alignment patterns in the matrix 749 1148
Number of undetermined characters or gaps (%) 16.74% 29.11%
Estimated base frequencies A 0.233125 0.251049
C 0.301275 0.219774
G 0.240690 0.295852
T 0.224910 0.23332
Substitution rates AC 0.982364 10 .905323
AG 3.653798 1.955792
AT 1.292861 1.245423
CG 1.143987 1.084109
CT 6.835565 4.832464
GT 1.000000 1.000000
Proportion of invariable sites (I) 0.612646 0.461707
Gamma distribution shape parameter (α) 0.721694 0.409261
Number of generated trees in BI 12002 4002
Number of trees sampled in BI after 25% were discarded as burn-in 9002 3002
Final split frequency 0.007976 0.007914
The total of unique site patterns 957 1149
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FIGURE 2. Maximum likelihood phylogenetic tree based on combined SSU-ITS-LSU sequence data for Sympoventuriaceae. Bootstrap
support values of maximum likelihood greater than 60% and Bayesian posterior probabilities (BYPP) greater than 0.95 are indicated above
the nodes. Newly added strains are in blue and ex-type strains are in bold. The tree is rooted to Venturia inaequalis (CBS 594.70 and CBS
815.69). Isolated substrates are indicated in triangles. Black: Human/ animal, green: plants, brown: soil, purple: rock/ sediments, blue:
aquatic habitat. unknown: empty
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30 • Phytotaxa 609 (1) © 2023 Magnolia Press
Taxonomy
Curvularia chiangraiensis Yasanthika & A.R. Gomes de Farias, sp. nov.
Index Fungorum number: IF559984, Faces of Fungi number: FoF 12886, FIGURE 3.
Etymology—The specific epithet “chiangraiensis” refers to the locality Chiang Rai Province (Thailand), where the holotype was
collected.
Holotype—MFLU 22-0256
Asexual morph: Conidiophores 50–150 × 2–7 μm (x = 117 × 4 μm, n = 20), simple, unbranched, septate, hyaline
to brown, micro- to semi-macro-nematous, sometimes geniculate towards upper region. Conidiogenous cells 7–11
× 3–7 (x = 9 × 5 μm, n = 20), polytretic, terminal or intercalary, sometimes proliferating sympodially, cylindrical to
sub-cylindrical, smooth to verruculose. Conidia 12–20 × 6–12 μm (x = 17 × 9 μm, n = 30), sublunate to ellipsoid or
obovoid, curved, sub-hyaline to pale brown, 1–2 distoseptate when immature, later becoming 2–3 euseptate and pale
brown to brown when mature, smooth to verruculose, swollen when mature, with the third cell from base becoming
prominent, partially enlarged along the horizontal axis, asymmetrically curved, darker, verrculose than apical and basal
septa, with apical and basal septa symmetrical. Hila inconspicuous to slightly conspicuous, slightly thickened 2–3 μm
(x = 2.5 μm) wide. Sexual morph: Undetermined.
Culture characteristics:—Colonies on PDA at 25 ℃ raised, filamentous at margin, with sparse aerial mycelia,
greyish brown to black, pale brown concentric rings, approximately 3.5 cm diam. at 7 days, reverse black. Hyphae
3–5 μm (x = 4 μm) wide, branched, septate, hyaline to pale brown when immature and later becoming brown, septate
hyphae.
Material examined:—Thailand, Chiang Rai, 20.0478N, 99.7619E, 863m, from forest soil, 23 September 2019.
W.A.E. Yasanthika (MFLU 22-0256, holotype); ex-type living culture MFLUCC 22-0091)
Notes—In this study, we introduce C. chiangraiensis (MFLU 22-0256) as a novel species isolated from soil in
Thailand based on both morphology and phylogeny. Multigene phylogeny indicates that C. chiangraiensis (MFLUCC
22-0091) forms an independent lineage sister to the clade comprising C. brachyspora (CBS 186.50) and C. simmonsii
(USJCC-0002) with 77% ML and 0.98 BYPP statistical support (FIGURE 1). The pairwise comparision of ITS,
GAPDH and tef1-α sequences of C. chiangraiensis with closely related species are shown in (TABLE 4)
Morphologically, C. chiangraiensis bears relatively smaller conidia. The conidial length-width ratio in C.
chiangraiensis is 1.7, while C. brachyspora and C. simmonsii show 1.9 and 2.6, respectively. Curvularia simmonsii
(21–27μm) and C. brachyspora (20–26 μm) shows distinct morphology in having longer conidia (Ferdinandez et
al. 2021) than C. chiangraiensis (12–20 μm). In addition, C. chiangraiensis can be distinguished by its relatively
shorter conidial lengths (12–20 μm) than C. aeria (18–26 μm) and C. homomorpha (25–42 μm) (Luttrell et al. 1959,
Sivanesan 1987, Almaguer et al. 2013, Tan et al. 2014, Ferdinandez et al. 2021). In C. chiangraiensis, conidial curvature
is prominent due to the third cell from base being partially enlarged along the horizontal axis and asymmetrically
curved with symmetrical apical and basal septa. In contrast, in C. brachyspora both central cells are approximately the
same size. Furthermore, conidia of C. aeria and C. simmonsii show slight curvature (Madrid et al. 2014, Kusai et al.
2016, Ferdinandez et al. 2021). Curvularia homomorpha has straight conidia in contrast to the curved conidia in C.
chiangraiensis (Sivanesan 1987, Tan et al. 2014). Conidiogenesis in C. chiangraiensis exhibits sympodial proliferation,
which is absent in C. brachyspora and C. simmonsii (Sivanesan 1987, Ferdinandez et al. 2021). Therefore, based
on both morphology and phylogenetic evidence, we introduce our species as C. chiangraiensis MFLUCC 22-0091)
isolated from forest soil in Thailand.
TABLE 4. Pairwise comparison of the bases of Curvularia chiangraiensis and closely related taxa.
New species Closely related species
Base pair differences (including gaps)
ITS GAPDH tef1-α
C. chiangraiensis
C. brachyspora 11/588 (93 gaps) 10/565 (62gaps) 3/892 (31gaps)
C. simmonsii 7/600 (99gaps) 9/590 (18gaps) 4/892 (39gaps)
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 31
FIGURE 3. Curvularia chiangraiensis (MFLU22-0256, holotype). a. Colony from above (on PDA). b. Colony from below (on PDA).
c. Sporulated colony with conidial attachments on the mycelium. d. Immature hyphae. e. Mature melanized hyphae. f–k. conidiogenesis.
l–p. Conidia. Scale bars: g = 25 μm, d–f, h–l = 20 μm, g, m–p= 10 μm.
YASANTHIKA ET AL.
32 • Phytotaxa 609 (1) © 2023 Magnolia Press
Curvularia chiangmaiensis Y. Marín, Senwanna & Crous, Mycosphere 8: 1565 (2017)
Index Fungorum number: IF822082, Faces of Fungi number: FoF 12887, FIGURE 4.
FIGURE 4. Curvularia chiangmaiensis (MFLU22-0252, new record) a. Colony from above (on PDA). b. Colony from below (on
PDA). c. Sporulated colony with conidial attachments on the mycelium. d. Immature hyphae e. Mature melanized hyphae. f. Hyaline
chlamydospores. g. Melanized chlamydospores h. Macronematous conidiogenesis on the conidiophore. i–m. Conidiogenesis. n–s.
Conidia. Scale bars: g = 25 μm, d–f, h–l = 20 μm, m–s= 10 μm
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 33
Asexual morph: Conidiophores 70–180 × 3.5–4 μm (x = 123 × 3.8 μm, n = 20), single, straight or flexuous, septate,
sometimes geniculate in the upper part, micronematous to macronematous, mononematous, pale brown to brown, paler
towards apex. Conidiogenous cells polytretic, proliferating sympodially, cylindrical to sub-cylindrical, hyaline to pale
brown, smooth-walled, terminal, or intercalary. Conidia 14–28 × 9–13 (x = 21 × 11 μm, n = 25) μm, ellipsoidal to
obovoid, sometimes curved, slightly verruculose, hyaline to pale brown, (2–)3-distoseptate, hila slightly conspicuous,
1.5–2.5 μm (x = 2). Sexual morph: Undetermined.
Culture characteristics:—Colonies on PDA at 25 ℃, flat, irregular margin, cottony, yellowish brown to black
surface, 3.5 cm diam. after 7 days, reverse black center, brownish black at margin. Hyphae 1.5–4 μm (x = 3 μm) wide,
often branched, rarely guttulate, smooth to rough-walled, sometimes geniculate, hyaline when immature, and later
become pale brown (melanized) to brown, septate. Chlamydospores guttulated, initially hyaline and becoming pale
brown to brown (melanized) when mature.
Known hosts and substrates:—Zea mays, Saccharum officinarum, and on soil (Marin-Felix et al. 2017b, Raza
et al. 2019, this study)
Known distribution:—China and Thailand (Marin-Felix et al. 2017b, Raza et al. 2019, this study).
Material examined:—Thailand, Krabi, 8°4’11 “N 98°53’25” E, 181m, on forest soil, 28 October 2019, W.A.E.
Yasanthika (MFLU 22-0252); living culture, MFLUCC 22-0084
Notes—Curvularia chiangmaiensis (CPC 28829) is initially isolated from Zea mays from Chiang Mai, Thailand
(Marin-Felix et al. 2017b). In this study, we isolated C. chiangmaiensis (MFLUCC 22-0084) from forest soils in
Thailand. Our collection (MFLUCC 22-0084) has similar morphological characteristics to the type of C. chiangmaiensis
(CPC 28829) by having cylindrical to sub-cylindrical conidiogenous cells and curved, ellipsoidal to obovoid (2–
)3-distoseptate conidia (Marin-Felix et al. 2017b). The conidial length-width ratio in both C. chiangmaiensis (CPC
28829) and our collection (MFLUCC 22-0084) are similar (1.9) (Marin-Felix et al. 2017b). However, our collection
has comparatively shorter conidiophores (76–177 μm vs 2000 μm) (Marin-Felix et al. 2017b). These intra-species
morphological variations can result from host specialization (Francisco et al. 2019). According to the multigene
phylogenetic results herein, our collection (MFLUCC 22-0084) grouped with C. chiangmaiensis strains (CPC28829,
LC12044, LC12030) with 94% ML, 1.00 BYPP bootstrap support. Therefore, based on morphological and phylogenetic
evidence, we introduce our collection (MFLUCC 22-0084) as a new geographical record of C. chiangmaiensis from
Thailand. Interestingly, this is the first record of this species from the soil.
Verruconis soli Yasanthika, Tennakoon & Wanas., sp. nov.
Index Fungorum number: IF 558479, Faces of Fungi number: FoF 12888, FIGURE 5.
Etymology—The specific epithet “soli”, from Latin, refers to the soil, the substrate of which the holotype was collected.
Holotype—MFLU22-0257
Asexual morph: Conidiophores 20–62 × 1–3.5 µm (x = 39 × 2 μm, n = 20), micronematous, mononematous, solitary,
unbranched, cylindrical to sub-cylindrical, straight to slightly flexuous, hyaline to brown, smooth. Conidiogenesis cells
holoblastic, determinate, and rarely intercalary. Conidia 5–11 × 3–4.5 µm (x = 8 × 3.5 μm, n = 20), hyaline, obovoid,
smooth becoming sub-cylindrical or ellipsoidal, slightly verruculose thick and sometimes pale brown, 1-septate and
constricted at the septum at maturity. Sexual morph: Undetermined.
Culture characteristics:—Colonies on PDA at 25 °C after 7 days become 3.5 cm diam., irregular margined,
velvety, dark brown, raised, grayish brown at center, reverse dark brown. Mycelium hyaline, become pale brown to
brown at maturity, 1–3 µm (x = 1.5 µm), smooth to verruculose, septate.
Material examined:—Thailand, Chiang Rai, 20°4’35”N,100°5’43”E, 450 m, from forest soil, 7 January 2020,
W.A.E. Yasanthika (MFLU 22-0257, holotype); ex-type living culture MFLUCC 22-0082; (MFLU 22-0253, paratype),
ex-paratype living culture MFLUCC 22-0090.
Notes—Verruconis is aggregated with thermophilic species, and several have been reported from the soil, viz.,
V. gallopava from thermal soils, V. verruculosa from grassland soils, and V. thailandica from soils near the waterfall
(Zhang et al. 2018, Hernández-Restrepo et al. 2020, Shen et al. 2020). Herein, we introduce V. soli from soil in Thailand
with morpho-molecular justifications. Verruconis soli is phylogenetically adjacent to V. hainanensis (YMF 1.04165),
V. pseudotricladiata (YMF 1.04915), and V. mangrovei (NFCCI 4390, NFCCI-4389) (FIGURE 2). Though V. soli has
low bootstrap support in the phylogenetic tree, and pairwise comparisons of LSU and ITS sequences of closely related
species (TABLE 5) showed significant differences that are higher than 1% in each scenario, which is common to many
species in this genus. Further, they differ ecologically from V. soli as V. hainanensis and V. pseudotricladiata have been
reported from freshwater habitats (Qiao et al. 2019) while V. mangrovei reported from marine habitats (Hyde et al.
YASANTHIKA ET AL.
34 • Phytotaxa 609 (1) © 2023 Magnolia Press
2020). Verruconis hainanensis, V. pseudotricladiata, and V. soli (MFLUCC 22-0082) are mainly distinguishable by
their conidiophores and conidial morphology. Verruconis hainanensis and V. pseudotricladiata have macronematous
conidiophores, whereas V. soli (MFLUCC 22-0082) has micronematous conidiophores (Qiao et al. 2019). In addition,
V. soli (MFLUCC 22-0082) has sub-cylindrical or ellipsoidal, 1-septate mature conidia and obovoid immature conidia,
while V. hainanensis has fusiform, 3-septate conidia that are rostrate at the apical cell (Qiao et al. 2019). Verruconis
pseudotricladiata has staurosporic, branched or unbranched conidia (cylindrical-clavate and 2–4-septate while
branched conidia are Y-, or T-shaped) (Qiao et al. 2019). Furthermore, the conidia of V. hainanensis and unbranched
conidia in V. pseudotricladiata have a slight constriction at the median septum, but V. soli (MFLUCC 22-0082) have a
prominent constriction at the septum and septation is absent at immature stages in conidia. Verruconis mangrovei has
been recorded as a sexual morph (Hyde et al. 2020); therefore, we could not compare the morphological characteristics
with our collection. The base pair differences of ITS sequences between V. mangrovei and V. soli were 31% (with
gaps). Therefore, we accommodate V. soli (MFLUCC 22-0082) isolated from forest soils in Thailand as a new species
with morpho-molecular support.
FIGURE 5. Verruconis soli (MFLU22-0257, holotype) a. Colony from above (on PDA). b. Colony from below (on PDA). c. Sporulated
colony. d. Melanized hyphae. e. Hyaline hyphae f–l. Conidiogenesis. m–r. Conidia. Scale bars: e = 15 μm, d, f–k = 10 μm, l–r = 5 μm.
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 35
TABLE 5. Pairwise comparison of the bases of Verruconis soli and closely related taxa.
New species Closely related species Base pair differences (including gaps)
ITS LSU
V. soli V. pseudotricladiata 248/615 (41 gaps) 37/844 (5gaps)
V. hainanensis 168/602 (35gaps) 24/829 (3gaps)
V. mangrovei 180/569 (24gaps) 41/778 (4gaps)
Verruconis thailandica Giraldo López & Crous, Fungal Systematics and Evolution 6: 21 (2020)
Index Fungorum number: IF833679, Faces of fungi number: FoF09618, FIGURE 6.
FIGURE 6. Verruconis thailandica (MFLU22-0258, new record) a. Colony from above (on PDA). b. Colony from below (on PDA).
c. Sporulated colony with conidial attachments on the mycelium. d. Immature hyphae e. Mature septate hyphae. f. Conidiogenesis
synnematous. g–j. Conidiogenesis mononematous conidiophores. k–p Conidia. Scale bars: d, e = 20 μm, f–l, n = 10 μm, m, o, p = 5 μm.
YASANTHIKA ET AL.
36 • Phytotaxa 609 (1) © 2023 Magnolia Press
Asexual morph: Conidiophores 1–2 μm (x = 1.8 μm, n=20) wide, macronematous, mononematous and synnematous,
erect, straight, or slightly bent, simple, sub-cylindrical, hyaline to pale brown, smooth-walled, conidiogenesis terminal
or intercalary, sympodial. Conidiogenous cells 2–3.5 × 1.5–3 μm (x = 3 × 1.8 μm, n = 20), monoblastic, discrete, sub-
cylindrical to ampulliform. Conidia 5–7 × 2.2–3.1 μm (x = 5.8 × 2.6 μm, n = 20), broadly ellipsoidal or rarely oval,
hyaline to pale brown, 1-septate, constricted at the septum, verrucose, thick-walled, protuberant hilum. Sexual morph:
Undetermined.
Culture characteristics:—Colonies on PDA at 25 °C after 7 days become 4 cm diam., filamentous margined,
hairy, and raised, brown to grayish brown at centre, flat, glabrous, dark brown at periphery, reverse dark brown,
diffusible brownish pigment. Mycelium hyaline, aseptate, smooth at immature become septate, pale brown, 1–3 (x =
2.5) μm diam. hyphae.
Known hosts and substrates:—soil (Hernández-Restrepo et al. 2020, this study).
Known distribution:—Thailand (Hernández-Restrepo et al. 2020, this study)
Material examined:—Thailand, Chiang Rai Province, 19°53’2” N, 100°5’37” E, 440m, from forest soil, 7
January 2020, W.A.E. Yasanthika (MFLU 22-0258); living culture, MFLUCC 22-0083.
Notes—Verruconis thailandica (CBS 145768) was first isolated from soil near a waterfall in Nakhon Nayok
Province, Thailand (Hernández-Restrepo et al. 2020). Verruconis thailandica has a closer phylogenetic affinity to V.
verruculosa and V. terricola (Hernández-Restrepo et al. 2020) (FIGURE 1). Our collection is grouped with V. thailandica
isolates (CBS 145768, GUCC 18267) in a well-supported clade with 100% ML, 1.00 BYPP support, and related to the
isolate CBS 145768 with 97% ML and 1.00 BYPP. Our strain morphologically resembles V. thailandica (CBS 145768)
by having erect, straight or slightly bent, sub-cylindrical conidiophores and broadly ellipsoidal or rarely oval, 1-septate
conidia (Hernández-Restrepo et al. 2020). We observed mononematous and synnematous conidiophores in our isolate
(MFLUCC 22-0083), which were missing in the type species (CBS 145768). This morphological variation can be
due to the environmental adaptations of the organism (Francisco et al. 2019). Therefore, with morpho-molecular
justifications, we present the second report of V. thailandica from forest soils in Thailand.
Discussion
Soil is a heterogeneous environment. Due to its complexity, it paves the way for many researchers to aim at linking
the diversity, taxonomy, and ecology of belowground fungi (Groenewald et al. 2018, Delgado-Baquerizo et al. 2020).
Thus, the soil is a valuable source for exploring new fungal species. Many researchers have isolated numerous soil-
associated Dothideomycetes by different isolation methods, which is becoming an emerging trend (Hou et al. 2020).
Hitherto, the majority of studies in this field have primarily concentrated on the dilution method. This method involves
diluting a sample or substance to reduce its concentration, allowing for further analysis or experimentation (Yasanthika
et al. 2022). However, the potential of alternative methods, beyond the dilution method, such as baiting and soil plating
technique has received limited attention and investigation (Hou et al. 2020, Yasanthika et al. 2022). These alternative
methods have not been thoroughly studied or explored, leaving their capabilities and potential benefits relatively
unknown. Therefore, further research is needed to understand and evaluate the effectiveness of these alternative
methods. Further, the ITS region in high-throughput sequencing (HTS), nuclear ribosomal and protein-coding regions
obtained from pure cultures have been found to provide stronger support for species identification (Groenewald et al.
2018, Hou et al. 2020).
The ITS region is useful for determining Curvularia species at the generic level and separating them into
major clades in Curvularia (Tan et al. 2014, 2016, 2018, Manamgoda et al. 2015). However, species delineation of
closely related Curvularia species cannot be determined by the ITS region alone, and tef1-α considered as the least
phylogenetically informative region for Curvularia. GAPDH is considered an essential locus to resolve closely related
species in this genus (Tan et al. 2014, 2016, 2018, Manamgoda et al. 2015, Ferdinandez et al. 2021). Following
the previous studies, we employed a concatenated alignment of the ITS, GAPDH, and tef1-α regions to accurately
determine the phylogenetic placement of our species (Manamgoda et al. 2015, Marin-Felix et al. 2017, 2019, 2020).
By conducting pairwise comparisons, we observed a similar pattern to previous research, with GAPDH emerging as
the most informative region in comparison to ITS and tef1-α. This highlights the significance of including GAPDH in
the analysis for a comprehensive understanding of the phylogenetic relationships among closely related Curvularia
species. Recently, soil-inhabiting Curvularia taxa have been detected from HTS methods without any linked specimens
(Delgado et al. 2021). The detection of soil-inhabiting Curvularia taxa through HTS methods without linked specimens
NOVEL DOTHIDEOMYCETES SPECIES Phytotaxa 609 (1) © 2023 Magnolia Press • 37
challenging to validate the identity and assign proper taxonomic classifications. The absence of a physical specimen
hinders the ability to differentiate true Curvularia taxa from potential contaminants or misidentified sequences and
limits the availability of valuable ecological and distribution data.
Curvularia exhibits similar morphological characteristics to Bipolaris, Pyrenophora, Drechslera, and Exserohilum
species (Manamgoda et al. 2015, Marin-Felix et al. 2019, 2020). However, conidial curvature in Curvularia differs
from Bipolaris by having inordinately enlarged intermediate cells that influence its conidial curvature, while the
conidial curvature occurs throughout the conidial length in Bipolaris. Regardless of this, both genera have species
with intermediate conidial morphology (Manamgoda et al. 2014, 2015, Marin-Felix et al. 2020). Besides, the asexual
morph of Curvularia resembles Pyrenophora; thus, only the sexual morph can be used to distinguish species between
these two genera (Marin-Felix et al. 2020). Therefore, molecular analyses are essential in species delineation among
these genera (Tan et al. 2014, Marin-Felix et al. 2019, 2020).
Many soil-dwelling Sympoventuriaceae species have been recorded in Ochroconis, Scolecobasidium, and
Verruconis (Samerpitak et al. 2016, Qiao et al. 2019, Hernández-Restrepo et al. 2020, Shen et al. 2020). Based
on previous studies, asexual morphs are more reliable than sexual morphs for species delimitation in Venturiales
with specific conidial, conidial apparatus, and conidiogenesis characteristics (Sivanesan 1978, Schubert et al. 2003,
Crous et al. 2007, Shen et al. 2020). Ochroconis and Verruconis exhibit rhexolytic conidial liberation, a rare fungal
characteristic that can be used for species delimitation in Sympoventuriaceae (Samerpitak et al. 2015, 2016). Verruconis
show oligotrophic nature and distribute at low nutritive conditions, such as clean water and cave walls (Samerpitak et
al. 2016). Verruconis thailandica and V. heavae have previously been identified in Thailand (Huanraluek et al. 2019,
Hernández-Restrepo et al. 2020).
Members of Pleosporales and Venturiales show saprobic lifestyles as the ancestral state. However, both
Curvularia and Verruconis have plant pathogens switched from the saprobic state, clustering at the terminal nodes
in phylogenetic trees, which is a common evolutionary characteristic at the class level (Dothideomycetes) (Crous et
al. 2009, Samerpitak et al. 2016, Marin-Felix et al. 2019, Haridas et al. 2020, Shen et al. 2020, Jayawardena et al.
2021). The identification of this evolutionary switch has significant implications for Dothideomycetes taxonomy and
evolutionary studies. The morpho-molecular information presented in this study fills an important gap in the field
and contributes to the overall data availability. This enhanced dataset will prove valuable for future investigations
into the taxonomy and evolutionary dynamics of Dothideomycetes. By incorporating these findings, researchers can
gain a deeper understanding of the evolutionary processes and relationships within this fungal class. However, it is
important to note that more exploratory studies are required to fully uncover the biodiversity of soil fungi. These
studies can provide additional data on the morphology and phylogeny of novel Dothideomycetes species found in soils
worldwide. Such efforts will contribute to a more comprehensive understanding of the ecological roles and adaptations
of Dothideomycetes in soil ecosystems. More exploratory studies are needed to uncover the soil fungal biodiversity
and provide data on the morphology and phylogeny of novel Dothideomycetes species from soils worldwide.
Acknowledgment
Dhanushka Wanasinghe would like to thank CAS President’s International Fellowship Initiative (grant number
2021FYB0005), the National Science Foundation of China (NSFC) under the project code 32150410362 and the
Postdoctoral Fund from Human Resources and Social Security Bureau of Yunnan Province. Danushka S. Tennakoon
would like to thank Post-Doctoral Fellowship 2022 for Reinventing Chiang Mai University (R000030885). W.A. Erandi
Yasanthika would like to thank the Mushroom Research Foundation (Thailand), Theisis support grant from Mae Fah
Luang University; reference number 7702(6)/580 and Impact of climate change on fungal diversity and biogeography
in the Greater Mekong Sub region (grant number: RDG6130001). D. Jayarama Bhat gratefully acknowledges the
financial support provided under the Distinguished Scientist Fellowship Programme (DSFP), at King Saud University,
Riyadh, Saudi Arabia. Abhaya Balasuriya is thanked for English editing.
YASANTHIKA ET AL.
38 • Phytotaxa 609 (1) © 2023 Magnolia Press
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