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© 2020 Westerdijk Fungal Biodiversity Instute 1
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
Fungal Systemacs and Evoluon
doi.org/10.3114/fuse.2020.06.01
VOLUME 6
DECEMBER 2020
PAGES 1–24
INTRODUCTION
This study focuses on ve genera that form part of the Genera of
Fungi project (www.generaoungi.org; Crous et al. 2014a). The
overall intenon of this project is to revise and update the generic
names of fungi, to provide DNA sequence data for them and to
restudy or recollect their type species. In this study, we provide
DNA sequence data for the unusual and poorly known genus
Kramasamuha. Furthermore, the phylogenec posion of the
genus Melnikomyces is claried. We also resolve the taxonomy
and phylogeny of Thysanorea and related Minimelanolocus
species in the Herpotrichiellaceae. Addional new taxa are
introduced based on morphological and DNA sequence data.
MATERIALS AND METHODS
Isolates
Freshly collected leaves and twigs were placed in damp chambers
and treated as described by Castañeda-Ruiz et al. (2016).
Protocols used for the collecon and processing of soil samples
are described in Giraldo et al. (2012, 2019) and Groenewald et al.
(2018). Aer 1 wk of incubaon on 2 % malt extract agar (MEA)
supplemented with penicillin-G and streptomycin, individual
colonies were transferred to MEA plates without anbiocs
and incubated between 22–24 °C for 7–14 d, in order to obtain
axenic cultures.
Colonies were sub-cultured onto 2 % potato dextrose agar
(PDA), oatmeal agar (OA), MEA (Crous et al. 2019), autoclaved
pine needles on 2 % tap water agar (PNA) (Smith et al. 1996),
and incubated at 25 °C under connuous near-ultraviolet light
to promote sporulaon. Reference strains and specimens are
maintained at the Westerdijk Fungal Biodiversity Instute (CBS
Culture Collecon and herbarium, respecvely), Utrecht, The
Netherlands or Coleção Octávio de Almeida Drummond (COAD),
Viçosa, Brazil.
DNA isolaon, amplicaon and analyses
Genomic DNA was extracted from fungal colonies growing on
MEA using the Wizard® Genomic DNA puricaon kit (Promega,
Madison, WI), following the manufacturer’s protocol. The
primers V9G (de Hoog & Gerrits van den Ende 1998) or ITS5
(White et al. 1990) and LR5 (Vilgalys & Hester 1990) were used
to amplify part of the nuclear rDNA operon (ITS) spanning the 3’
end of the 18S nrRNA gene, the rst internal transcribed spacer
(ITS1), the 5.8S nrRNA gene, the second ITS region (ITS2) and
approximately 900 bp of the 5’ end of the 28S nrRNA gene. The
primers ITS4 (White et al. 1990) and LR0R (Vilgalys & Hester
1990) were used as internal sequence primers to ensure high
quality sequences over the enre length of the amplicon. Part
of the 18S small subunit nrRNA gene (SSU) was amplied and
sequenced for selected isolates using NS1 and NS4 (White et
al. 1990). Amplicaon condions followed those described by
Cheewangkoon et al. (2008). Part of the acn gene (act) was
The Genera of Fungi – G6: Arthrographis, Kramasamuha, Melnikomyces, Thysanorea, and Verruconis
M. Hernández-Restrepo1*, A. Giraldo1,2, R. van Doorn1, M.J. Wingeld3, J.Z. Groenewald1, R.W. Barreto4, A.A. Colmán4, P.S.C. Mansur4,
P.W. Crous1,2,3
1Westerdijk Fungal Biodiversity Instute, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
2Faculty of Natural and Agricultural Sciences, Department of Plant Sciences, University of the Free State, P.O. Box 339, Bloemfontein 9300, South
Africa
3Department of Genecs, Biochemistry and Microbiology, Forestry and Agricultural Biotechnology Instute (FABI), University of Pretoria, Pretoria,
0002, South Africa
4Departamento de Fitopatologia, Universidade Federal de Viçosa, 36570-900, Viçosa, Minas Gerais, Brazil
*Corresponding author: m.hernandez@wi.knaw.nl
Abstract: The Genera of Fungi series, of which this is the sixth contribuon, links type species of fungal genera to their
morphology and DNA sequence data. Five genera of microfungi are treated in this study, with new species introduced
in Arthrographis, Melnikomyces, and Verruconis. The genus Thysanorea is emended and two new species and nine
combinaons are proposed. Kramasamuha sibika, the type species of the genus, is provided with DNA sequence data
for rst me and shown to be a member of Helminthosphaeriaceae (Sordariomycetes). Aureoconidiella is introduced
as a new genus represenng a new lineage in the Dothideomycetes.
Key words:
DNA barcodes
fungal systemacs
ITS
LSU
new taxa
Corresponding editor:
U. Braun
Effectively published online: 5 February 2020.
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
2
amplied and sequenced for selected isolates using the primer
set Act1/Act4 (Voigt & Wöstemeyer 2000). The soware SeqMan
Pro v. 13.0.0 (DNASTAR, Madison, WI) and Geneious v. 11.0.4
(Kearse et al. 2012; hps://www.geneious.com) were used to
obtain consensus sequences of each isolate. Blast searches using
ITS and LSU sequences were performed for each isolate and the
closest matches were retrieved from GenBank and included in
the phylogenec analyses. Mulple sequence alignments for
individual genes were generated using the online version of
MAFFT (hp://ma.cbrc.jp/alignment/soware/). Subsequent
phylogenec analyses from individual and combined datasets
were conducted using Maximum-likelihood (ML) performed
on the CIPRES Science Gateway portal (Miller et al. 2012) using
RAxML v. 8.2.10 (Stamatakis 2014). The default parameters were
used, and bootstrap support (BS) was obtained using the rapid
bootstrapping algorithm with the automac halt opon. A BS ≥
95 % was considered as stascally signicant. Sequence data
were deposited in the GenBank/ENA public databases (Table 1)
and the alignments and trees in TreeBASE (hp://www.treebase.
org).
Morphology
Slide preparaons were mounted in lacc acid or water from
colonies sporulang on the media previously menoned.
Observaons were made with a Nikon SMZ1500 dissecng
microscope and with a Nikon Eclipse Ni compound microscope
using a DSRi2 digital camera (Nikon, Tokyo, Japan) and NIS-
Elements imaging soware v. 4.3. Colony characters and
pigment producon were noted aer 1–2 wk of growth on
MEA, PDA and OA incubated at 25 ºC under natural light.
Colony colours (surface and reverse) were determined using
the colour charts of Rayner (1970). Taxonomic noveles were
deposited in MycoBank (www.MycoBank.org; Crous et al.
2004).
RESULTS
Phylogeny
Three overview phylogenies were generated in this study. The
rst two of these were based on a paral alignment of LSU to
provide the phylogenec posion of the treated genera and
species within the Dothideomycetes (Fig. 1) and Sordariomycetes
(Fig. 2). A third analysis was implemented for selected
Herpotrichiellaceae genera based on a concatenated ITS/LSU/
SSU alignment (Fig. 3). Other phylogenec trees specic to the
treated species are discussed in the notes for those taxa.
The BLAST search results using the LSU and ITS sequences
for the isolates CBS 145943, CBS 145767, CBS 145768, and JW
showed that they were related to members of Cladoriellales
and Asterinales, Venturiales and Eremomycetales in the
Dothideomycetes. Consequently, the currently accepted taxa
in those orders and other orders in the Dothideomycetes were
included in our analyses (Fig. 1).
The overview phylogeny of Dothideomycetes (Fig. 1)
revealed that in the Arthrographis subclade (96 % BS), the
strains JW 49012, JW 190014, and JW 209002 grouped together
in a separate clade represenng a putave new species that is
described below.
The genus Aureoconidiella is introduced to accommodate
CBS 145943 that formed a lineage disnct from other genera,
families and orders included in the analysis. A new family and
order are introduced for this genus. Furthermore, the isolates
CBS 145767 and CBS 145768 nested in the Sympoventuriaceae
clade (91 % BS) within the Venturiales. They were closely related
but dierent to Melnikomyces vietnamensis and Verruconis
verruculosa, respecvely. These isolates are consequently
considered to represent putave new species in the genera
Melnikomyces and Verruconis that are introduced below.
The overview phylogeny of Sordariomycetes (Fig. 2) revealed
that four isolates, CBS 146133, CBS 146338, CBS 146339,
and COAD 2632 of Kramasamuha sibika grouped together
in a fully-supported terminal clade (100 % BS), related to
Helminthosphaeriaceae (97 % BS).
The combined analysis of the ITS/LSU/SSU (Fig. 3) revealed
that Minimelanolocus and Thysanorea cluster together in the
same clade (86 % BS). Several species treated so far as belonging
to Minimelanolocus, i.e. M. aquacus, M. asiacus, M. curvatus,
M. melanicus, M. obscurus, M. rosselianus, M. submersus, and
M. thailandensis, proved to be congeneric with Thysanorea and
therefore new combinaons are proposed to accommodate
them. In addion, CBS 145909 and CBS 145910 formed a
separate subclade (100 % BS) within Thysanorea, represenng
two putave new species.
Taxonomy
Aureoconidiellales Hern.-Restr. & Crous, ord. nov. MycoBank
MB833918.
Descripon: See descripon of Aureoconidiella.
Type family: Aureoconidiellaceae Hern.-Restr. & Crous
Aureoconidiellaceae Hern.-Restr. & Crous, fam. nov. MycoBank
MB833917.
Descripon: See descripon of Aureoconidiella.
Type genus: Aureoconidiella Hern.-Restr. & Crous
Aureoconidiella Hern.-Restr. & Crous, gen. nov. MycoBank
MB833915.
Etymology: Name refers to the golden brown colour of its
conidia.
Conidiophores macronematous, simple, septate, brown.
Conidiogenous cells integrated, terminal, polyblasc, with
thickened scars, brown to pale brown. Conidia globose to
subglobose with apiculate base, inially subhyaline, golden
brown at maturity, verrucose.
Type species: Aureoconidiella foliicola Hern.-Restr. & Crous
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
3
Table 1. GenBank accession numbers of taxa included in this study.
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Dothideomycetes
Aliquandospite khaoyaiensis CBS 118232 Thailand Twig –GU301796.1 –Schoch et al. (2009)
SS3028 Thailand –EF175649.1 –Campbell et al. (2007)
Alternaria tenuissima CBS 918.96 UK Dianthus chinensis –KC584311.1 –Woudenberg et al. (2013)
Alysidiella parasica CBS 120088 South Africa Leaves Eucalyptus sp. –DQ923525.1 –Summerell et al. (2006)
Apiosporina collinsii CBS 118973 Canada Amelanchier alnifolia –GU301798.1 –Schoch et al. (2009)
Arthrographis arxii CBS 203.78 India Dung of herbivore GQ272638.1 AB213426.1 HG316563.1/- Murata et al. (2005), Kang et al. (2010),
Giraldo et al. (2014a)
Arthrographis chlamydospora CBS 135936 USA Human urine HG004554.1 HG004543.1 HG316560.1/- Giraldo et al. (2014a)
Arthrographis curvata CBS 135933 USA Human nails HG004557.1 HG004539.1 HG316557.1/- Giraldo et al. (2014a)
CBS 135934 Spain River sediment HG004556.1 HG004542.1 HG316558.1/- Giraldo et al. (2014a)
Arthrographis globosa UTHSC 11-757 USA Bronchial wash HG004553.1 HG004541.1 HG316561.1/- Giraldo et al. (2014a)
Arthrographis grakisi JW 22011 = CBS 145529 The Netherlands Soil MN794359 MN794336 MN816497 This study
JW 22015 The Netherlands Soil MN794360 MN794337 MN816498 This study
JW 22019 The Netherlands Soil MN794361 MN794338 MN816499 This study
JW 49011 The Netherlands Soil MN794362 MN794339 MN816500 This study
JW 49012 The Netherlands Soil MN794363 MN794340 MN816501 This study
JW 180011 The Netherlands Soil MN794364 MN794341 MN816502 This study
JW 190014= CBS 145530 The Netherlands Soil MN794365 MN794342 MN816503 This study
JW 199018 The Netherlands Soil MN794366 MN794343 MN816504 This study
JW 209002 The Netherlands Soil MN794367 MN794344 MN816505 This study
JW 209003 The Netherlands Soil MN794368 MN794345 MN816506 This study
Arthrographis kalrae CBS 693.77 India Sputum AB116536.1 AB116544.1 HG316544.1/- Xi et al. (2004), Giraldo et al. (2014a)
JW 21004 The Netherlands Soil MN794369 MN794346 MN816507 This study
JW 21008 = CBS 145527 The Netherlands Soil MN794370 MN794347 MN816508 This study
JW 21029 The Netherlands Soil MN794371 MN794348 MN816509 This study
Arthrographis longispora CBS 135935 USA Human foot HG004555.1 HG004540.1 HG316559.1/- Giraldo et al. (2014a)
JW 22007 = CBS 145528 The Netherlands Soil MN794372 MN794349 MN816510 This study
Asterina chrysophylli VIC 42823 Brazil Leaves Henrieea
succosa
–KP143738.1 –Guamosim et al. (2015)
Asterina melastomas VIC 42822 Brazil Leaves Miconia sp. –NG_057055.1 –Guamosim et al. (2015)
Asterotexis cucurbitacearum PMA M141224 Panama Sechium edule –HQ610510.1 –Unpublished
VIC 24814 Brazil Leaves Cucurbita pepo –NG_057054.1 –Guamosim et al. (2015)
Aulographina eucalyp CPC 12986 Australia Eucalyptus cloeziana –HM535600.1 –Cheewangkoon et al. (2012)
Aureoconidiella foliicola CBS 145943 South Africa Leaves Syzygium
cordatum
MN794373 MN794350 –This study
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
4
Table 1. (Connued).
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Bezerromyces brasiliensis URM7411 Brazil Tacinga inamoena –KX518623.1 –Bezerra et al. (2017)
Bezerromyces pernambucoensis URM7412 Brazil Tacinga inamoena –KX518624.1 –Bezerra et al. (2017)
Blastacervulus eucalyp CBS 124759 Australia Eucalyptus robertsonii
subsp. hemisphaerica
–GQ303302.1 –Cheewangkoon et al. (2009)
Blastacervulus robbenensis CBS 124780 Cyprus Eucalyptus sp. –HM628777.1 –Cheewangkoon et al. (2012)
Botryosphaeria dothidea CBS 115476 Switzerland Prunus sp. –DQ377852.1 –Crous et al. (2006b)
Brachiosphaera tropicalis E192 – – – EF175653.1 –Campbell et al. (2007)
Byssosphaeria jamaicana SMH 1403 – – – GU385152.1 –Mugambi & Huhndorf (2009a)
Byssosphaeria salebrosa SMH 2387 – – – GU385162.1 –Mugambi & Huhndorf (2009a)
Capnodium coeae CBS 147.52 Zaire Berry Coea robusta –MH868489.1 –Vu et al. (2019)
Cladoriella eucalyp CBS 115899 South Africa Leaves Eucalyptus –EU040224.1 –Crous et al. (2007b)
Cladoriella kinglakensis CPC 32730 Australia Leaves Eucalyptus –MG386126.1 –Crous et al. (2017)
Cladoriella paleospora CBS 124761 Australia Leaves Eucalyptus –GQ303303.1 –Cheewangkoon et al. (2009)
Cladoriella rubrigena CBS 124760 Australia Leaves Eucalyptus –MH874921.1 –Vu et al. (2019)
Cladoriella xanthorrhoeae CBS 143398 Australia Leaves Xanthorrhoea sp. –NG_059054.1 –Crous et al. (2017)
Cladosporium halotolerans CBS 127371 Cuba Human –MH875988.1 –Vu et al. (2019)
Cladosporium variabile CBS 121636 USA Spinacia oleracea –MH874684.1 –Vu et al. (2019)
Clavaspora thailandica MFLUCC 17-2237 Thailand Hevea brasiliensi –MH062960.1 –Unpublished
MFLUCC 10-0107 Thailand Dead stems –NG_058863.1 –Boonmee et al. (2014)
Dibotryon morbosum N/A USA Prunus sp. –EF114694.1 –Winton et al. (2007)
Diplodia mula CBS 431.82 The Netherlands Dead branches Fraxinus
excelsior
–DQ377863.1 –Crous et al. (2006b)
Dissoconium aciculare CBS 204.89 Germany Astragalus –GU214419.1 –Crous et al. (2009a)
Eremomyces bilateralis CBS 781.70 USA Dung of pack rat HG004552.1 HG004545.1 HG316562.1/- Giraldo et al. (2014a)
Fusicladium pini CBS 463.82 The Netherlands Needle Pinus sylvestris –EU035436.1 –Crous et al. (2007c)
Fusicladium ramoconidii CBS 462.82 The Netherlands Needle Pinus sp. –EU035439.1 –Crous et al. (2007c)
Gibbera conferta CBS 191.53 Switzerland Vaccinium uliginosum –GU301814.1 –Schoch et al. (2009)
Gloniopsis arciformis GKM L166A – – – GU323211.1 –Schoch et al. (2009)
Glonium circumserpens CBS 123342 Tasmania Wood –FJ161208.1 –Boehm et al. (2009)
Glonium circumserpens CBS 123343 Tasmania Saxicolous on limestone –FJ161200.1 –Boehm et al. (2009)
Helicomyces roseus CBS 283.51 Switzerland Dead bark –AY856881.1 –Tsui et al. (2006)
Herpotrichia juniperi AFTOL-ID 1608 Switzerland Juniperus nana –DQ678080.1 –Schoch et al. (2009)
Heteroconium eucalyp CBS 120122 Uruguay Leaves Eucalyptus dunnii –DQ885893.1 –Crous et al. (2006a)
Hysterium angustatum CBS 123334 USA Bark Pinus rigida –FJ161207.1 –Boehm et al. (2009)
Hysterium pulicare ANM1455 USA – – GQ221904.1 –Mugambi & Huhndorf (2009b)
Hysteropatella clavispora CBS 247.34 USA Salix sp. –AY541483.1 –Lumbsch et al. (2005)
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
5
Table 1. (Connued).
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Hysteropatella prosi H.B. 9934b Germany Malus domesca –KT876980.1 –Unpublished
Jahnula appendiculata SS2900 Thailand – – EF175654.1 –Campbell et al. (2007)
Jahnula aquaca R68-1 USA – – EF175655.1 –Campbell et al. (2007)
Jahnula bipileata F49-1 USA – – EF175657.1 –Campbell et al. (2007)
Jahnula seychellensis SS2113.1 Thailand – – EF175665.1 –Campbell et al. (2007)
Leptoxyphium fumago CBS 123.26 Indonesia Hibiscus liaceus –GU214430.1 –Crous et al. (2009a)
Macrophomina phaseolina CBS 227.33 –Zea mays –DQ377906.1 –Crous et al. (2006b)
Melanomma pulvis-pyrius CBS 124080 France Bark Salix caprea –GU456323.1 –Zhang et al. (2009a)
Melnikomyces thailandicus CBS 145767 Thailand Soil MN794374 MN794351 –This study
Melnikomyces vietnamensis CBS 136209 Vietnam Leaves –NG_058087.1 –Crous et al. (2014b)
Mycosphaerella puncformis CBS 113265 The Netherlands Dead leaves Quercus
robur
–DQ470968.1 –Spatafora et al. (2006)
Neocoleroa metrosideri PDD107531 New Zealand Metrosideros excelsa –NG_059638.1 –Johnston & Park (2016)
Neofusicoccum mangiferae CBS 118532 Australia Mangifera indica –NG_055730.1 –Crous et al. (2006b)
Neofusicoccum nonquaesitum CBS 126655 USA Umbellularia californica –NG_058258.1 –Yang et al. (2017)
Ochroconis constricta CBS 202.27 USA Soil –KF156147.1 –Samerpitak et al. (2014)
Ochroconis gamsii CBS 239.78 Sri Lanka Leaf Caryota plumosa –NG057992.1 –Samerpitak et al. (2014)
Patellaria cf. atrata BCC 28876 Thailand – – GU371828.1 –Schoch et al. (2009)
BCC 28877 Thailand – – GU371829.1 –Schoch et al. (2009)
Phaeocryptopus gaeumannii CBS 267.37 Germany Pseudotsuga menziesii –EF114698.1 –Winton et al. (2007)
Phaeotrichum benjaminii CBS 541.72 –Dung of rodent –AY779311.1 –Lumbsch et al. (2005)
Phoma herbarum CBS 567.63 USA Fruit Malus sylvestris –MH869982.1 –Vu et al. (2019)
Pirozynskiella laurisilvaca CBS 138109 Spain Leaves Laurus sp. –NG_058462.1 –Hernández-Restrepo et al. (2017)
Psiloglonium simulans CBS 206.34 USA Tilia sp. –FJ161178.1 –Boehm et al. (2009)
Rhexothecium globosum CBS 955.73 Egypt Desert soil MH860827.1 HG004544.1 –Giraldo et al. (2014a), Vu et al. (2019)
Rhydhysteron rufulum CBS 306.38 –Pistacia chinensis –FJ469672.1 –Schoch et al. (2009)
Schizothyrium pomi CBS 228.57 Italy – – EF134947.1 –Batzer et al. (2008)
CBS 486.50 The Netherlands Polygonum sachalinense –EF134948.1 –Batzer et al. (2008)
Scolecobasidiella avellanea CBS 772.73 Somalia Soil –EF204505.1 –Unpublished
Stemphylium herbarum CBS 191.86 India Leaf Medicago sava –JX681120.1 –Verkley et al. (2014)
Sympoventuria capensis CBS 120136 South Africa Leaf lier Eucalyptus sp. –NG_057984.1 –Samerpitak et al. (2014)
Teratosphaeria destructans CBS 111369 Indonesia Eucalyptus grandis –EU019287.2 –Crous et al. (2007a)
Teratosphaeria brillosa CBS 121707 South Africa Leaves Protea sp. –KF902075.1 –Quaedvlieg et al. (2014)
Teratosphaeria stellenboschiana CBS 116428 South Africa Leaf lier Eucalyptus sp. –EU019295.1 –Crous et al. (2007a)
Trichodelitschia bisporula CBS 262.69 The Netherlands Dung of Rabbit –GU348996.2 –Schoch et al. (2009)
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
6
Table 1. (Connued).
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Tubeua paludosa CBS 245.49 The Netherlands Corylus avellana –MH856510.1 –Vu et al. (2019)
Uwebraunia commune CBS 110747 South Africa Eucalyptus nitens –GQ852589.1 –Crous et al. (2009b)
Uwebraunia dekkeri CBS 111282 Zambia Eucalyptus globulus –GU214425.1 –Crous et al. (2009b)
Venturia inaequalis CBS 176.42 France – – GU348998.1 –Schoch et al. (2009)
Venturia populina CBS 256.38 Italy Populus canadensis –GU323212.1 –Schoch et al. (2009)
Verruconis calidiuminalis CBS 125818 Japan Hot spring euent –NG_057985.1 –Samerpitak et al. (2014)
Verruconis gallopava CBS 547.81 New Zealand – – KF156109.1 –Samerpitak et al. (2014)
CBS 437.64 USA Brain abscess Meleagris
gallopavo
–NG_58016.1 –Machouart et al. (2014)
Verruconis thailandica CBS 145768 Thailand Soil MN794375 MN794352 –This study
Verruconis verruculosa CBS 119775 Malaysia Root Hevea species –KF282668.1 –Machouart et al. (2014)
Xiliomyces brasiliensis URM7413 Brazil Tacinga inamoena –KX518625.1 –Bezerra et al. (2017)
Zasmidium cellare CBS 146.36 –Wall in wine cellar –EU041878.1 –Arzanlou et al. (2007)
Euroomycetes
Aculeata aquaca MFLUCC 11-0529 Thailand Submerged wood MG922575.1 MG922579.1 –/MG922571.1 Dong et al. (2018)
Capronia pilosella AFTOL-ID 657 – – DQ823099.1 DQ823106.1 –/DQ826737.1 James et al. (2006)
Cladophialophora carrionii CBS 160.54 Australia Man FJ358234.1 FJ358302.1 –/AF050262.1 Gueidan et al. (2008), Untereiner &
Naveau (1999)
Cladophialophora minourae CBS 556.83 Japan Decaying wood FJ358235.1 FJ358303.1 –/AY251087.1 Braun et al. (2003), Gueidan et al.
(2008)
Cladophialophora parmeliae CBS 129337 Portugal –JQ342182.1 – –/JQ342180.1 Diederich et al. (2013)
Cladophialophora sublis CBS 122642 The Netherlands Ice tea NG_058961.1 KX822283.1 –/NR_111363.1 Badali et al. (2008), Vasse et al. (2017)
Cyphellophora oxyspora CBS 698.73 Sri Lanka –KC455262.1 KC455305.1 –/KC455249.1 Réblová et al. (2013)
Cyphellophora sessilis CBS 243.85 The Netherlands Resin Picea abies EU514700.1 KC455308.1 –/EU514700.1 Untereiner et al. (2008), Réblová et al.
(2013)
Exophiala jeanselmei CBS 507.90 Uruguay Man FJ358242.1 FJ358310.1 –/NR_111129.1 Gueidan et al. (2008)
Exophiala nigra dH 12,296 – – FJ358244.1 FJ358312.1 –Gueidan et al. (2008)
Exophiala pisciphila CBS 537.73 USA Ictalurus punctatus MH872483.1 JN856018.1 –/AF050272.1 de Hoog et al. (2011), Untereiner &
Naveau (1999), Vu et al. (2019)
AFTOL-ID 669 – – DQ823101.1 DQ823108.1 –/DQ826739.1 Gueidan et al. (2008)
Exophiala salmonis AFTOL-ID 671 – – EF413609.1 EF413608.1 –Geiser et al. (2006)
CBS 157.67 Canada Salmo clarkii MH870616.1 JN856020.1 –/NR_121270.1 de Hoog et al. (2011), Schoch et al.
(2014), Vu et al. (2019)
Exophiala xenobioca CBS 115831 Germany Browncol FJ358246.1 FJ358314.1 –/AY857539.1 Gueidan et al. (2008)
Fonsecaea monophora CBS 102243 FJ358247.1 FJ358315.1 –/EU938579.1 Gueidan et al. (2008)
Melanoctona tectonae MFLUCC 12-0389 Thailand Tectona grandis KX258779.1 KX258780.1 –/KX258778.1 Unpublished
Phialophora americana AFTOL-ID 658 – – FJ358226.1 FJ358294.1 –Gueidan et al. (2008)
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
7
Table 1. (Connued).
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Phialophora verrucosa AFTOL-ID 670 – – EF413615.1 EF413614.1 –Geiser et al. (2006)
Rhinocladiella anceps AFTOL-ID 659 – – DQ823102.1 DQ823109.1 –/DQ826740.1 James et al. (2006)
Rhinocladiella anceps CBS 181.65 Canada Soil EU041862.1 AY554292.1 –/MH858534.1 Arzanlou et al. (2007)
Thysanorea asiaca MFLUCC 15-0237 China Submerged wood KR215610.1 KR215615.1 –/KR215604.1 Liu et al. (2015)
Thysanorea cantrelliae CBS 145909 USA Unidened twig MN794376 MN794353 –/MN794382 This study
Thysanorea curvata MFLUCC 15-0259 China Submerged wood KR215609.1 KR215614.1 –/KR215605.1 Liu et al. (2015)
Thysanorea lotorum CBS 235.78 USA Root Lotus corniculatus MH872892.1 – –/MH861130.1 Vu et al. (2019)
KUMCC 15-0206 China Submerged wood KX789215.1 – –/KX789212.1 Liu et al. (2015)
Thysanorea melanica MFLUCC 15-0415 China Submerged wood KR215613.1 KR215618.1 –/KR215608.1 Liu et al. (2015)
Thysanorea nonramosa MFLUCC 17-2378 Thailand Wood MH532970.1 – –/MH532971.1 Wang et al. (2019)
Thysanorea obscura MFLUCC 15-0416 China Submerged wood KR215611.1 KR215616.1 –/KR215606.1 Liu et al. (2015)
Thysanorea papuana CBS 212.96 Papua New Guinea –EU041871.1 – –/EU041814.1 Arzanlou et al. (2007)
MFLUCC 15-0966 Thailand Submerged wood MG922576.1 MG922580.1 –/MG922572.1 Dong et al. (2018)
Thysanorea rousseliana CBS 126086 Spain Dead branches Quercus
ilex
MH875246.1 – –/MH863784.1 Vu et al. (2019)
Thysanorea seiferi CBS 145910 USA Unidened twig MN794377 MN794354 –/MN794383 This study
Thysanorea thailandensis MFLUCC 15-0971 Thailand Submerged wood MG922577.1 MG922581.1 –/MG922573.1 Dong et al. (2018)
Thysanorea yunnanense MFLUCC 15-0414 Thailand Submerged wood KR215612.1 KR215617.1 –/KR215607.1 Liu et al. (2015)
Veronaea botryosa CBS 254.57 Italy Sansa olive EU041873.1 JN856021.1 –/EU041816.1 Arzanlou et al. (2007)
MFLUCC 11-0072 Thailand Submerged wood MG922574.1 MG922578.1 –/MG922570.1 Dong et al. (2018)
Veronaea compacta CBS 268.75 South Africa –EU041876.1 – –/EU041819.1 Arzanlou et al. (2007)
Veronaea japonica CBS 776.83 Japan Dead bamboo culm EU041875.1 – –/EU041818.1 Arzanlou et al. (2007)
Sordariomycetes
Anthostomella sp. SMH3101 USA – – AY780050.1 –Miller & Huhndorf (2005)
Camarops tubulina SMH4614 Denmark – – AY346266.1 –Huhndorf et al. (2004)
Camarops ustulinoides SMH1988 USA – – AY346267.1 –Huhndorf et al. (2004)
Chaetosphaeria ovoidea SMH2605 USA – – AF064641.1 –Fernandez et al. (1999)
Coniochaeta discoidea SANK 12878 – – – AY346297.1 –Huhndorf et al. (2004)
Coniochaedium savoryi TRTC 51980 – – – AY346276.1 –Huhndorf et al. (2004)
Cytospora ceratosperma AR3426 Austria Quercus robur –AF408387.1 –Castlebury et al. (2002)
Diaporthe phaseolorum FAU458 USA – – AY346279.1 –Huhndorf et al. (2004)
Echinosphaeria canescens JHC97-006 – – – KF765604.1 –Miller et al. (2014)
SMH4666 – – – KF765605.1 –Miller et al. (2014)
SMH4791 – – – AY436403.1 –Miller & Huhndorf (2004)
TL5730 – – – AY436404.1 –Miller & Huhndorf (2004)
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
8
Table 1. (Connued).
Tax a Isolates1Country Substrate GenBank accession numbers2References
ITS LSU act/SSU
Eutypa sp. SMH3580 Panama Branch –AY346280.1 –Huhndorf et al. (2004)
Fusarium ambrosium SMH1999 – – – AY780077.1 –Miller & Huhndorf (2005)
Helminthosphaeria carpathica SMH3903 – – – KF765606.1 –Miller et al. (2014)
Helminthosphaeria cf. stuppea JF04120 – – – KF765611.1 –Miller et al. (2014)
TL11998 – – – KF765612.1 –Miller et al. (2014)
Helminthosphaeria clavariarum SMH4609 Denmark Clavulina cristata –AY346283.1 –Huhndorf et al. (2004)
Helminthosphaeria corciorum JF04225 – – – KF765607.1 –Miller et al. (2014)
Helminthosphaeria odonae ANM928 – – – KF765610.1 –Miller et al. (2014)
Helminthosphaeria tomaculum SMH2485 – – – KF765613.1 –Miller et al. (2014)
Helminthosphaeria triseptata JF04015 – – – KF765614.1 –Miller et al. (2014)
Hilberina caudata SMH1542 – – – KF765615.1 –Miller et al. (2014)
Hilberina munkii SMH1531 – – – KF765616.1 –Miller et al. (2014)
Kramasamuha sibiki CPC 35619 = CBS 146338 Australia Leaves Lophostemon
confertus
MN794378 MN794355 –This study
CPC 36725 = CBS 146339 Malaysia Needles Pinus
tecunumanii
MN794379 MN794356 –This study
CBS 146133 = CPC 36153 South Africa Leaves Syzygium
cordatum
MN794380 MN794357 –This study
COAD 2632 Brazil Leaves Hypericum
innodorum
MN794381 MN794358 –This study
Lasiosphaeria ovina SMH1538 – – – AF064643.1 –Fernandez et al. (1999)
Neurospora crassa MUCL 19026 – – – AF286411.1 –Untereiner et al. (2001)
Ruzenia spermoides ANM163 – – – KF765618.1 –Miller et al. (2014)
SMH4606 – – – AY436422.1 –Miller & Huhndorf (2004)
SMH4655 – – – KF765619.1 –Miller et al. (2014)
Sporoschisma hemipsila SMH2125 – – – AY346292.1 –Huhndorf et al. (2004)
Synaptospora plumbea ANM963 – – – KF765620.1 –Miller et al. (2014)
SMH3962 – – – KF765621.1 –Miller et al. (2014)
Valsonectria pulchella SMH1193 – – – AY346304.1 –Huhndorf et al. (2004)
1 BCC: BIOTEC Culture Collecon, Naonal Center for Genec Engineering and Biotechnology (BIOTEC), Bangkok, Thailand; CBS: Culture Collecon of the Westerdijk Fungal Biodiversity Instute, Utrecht,
The Netherlands; COAD: Coleção Octávio de Almeida Drummond, Viçosa, Brazil; CPC: Culture Collecon of Pedro Crous, Utrecht, The Netherlands; JW: Johanna Westerdijk Culture Collecon, Utrecht, The
Netherlands; MFLUCC: Mae Fah Luang University Culture Collecon, Chiang Rai, Thailand; UTHSC: Fungus Tesng Laboratory of the University of Texas Health Science Center at San Antonio, USA. For other
acronyms see references.
2 LSU: Large subunit of the nrDNA; SSU: Small subunit of the nrDNA; ITS: internal transcribed spacer regions of the nrDNA and intervening 5.8S nrDNA; act: paral acn gene. Accession numbers of sequences
newly generated in this study are indicated in bold.
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
9
Melanommataceae
Gloniaceae
Hysteriaceae
Pleosporaceae
Didymellaceae
Botryosphaeriaceae
Tubeufiaceae
Eremomycetaceae
Bezerromycetaceae
Patelleriaceae
Asterotexiaceae
Incertae sedis
Schizothyriaceae
Dissoconiaceae
Aliquandostipitaceae
HYSTERIALES
MYTILIDINALES
PLEOSPORALES
BOTRYOS-
PHAERIALES
TUBEUFIALES
BEZERRO-
MYCETALES
JAHNULALES
ASTERO-
TEXIALES
CAPNODIALES
PATELLERIALES
I
EREMO-
MYCETALES
INCERTAE SEDIS
JW190014
Jahnula bipileata F49-1
Hysterium pulicare ANM1455
Eremomyces bilateralis CBS 781.70
Arthrographis globosa UTHSC 11-757
Aliquandostipite khaoyaiensis SS3028
Uwebraunia dekkeri CBS 111282
Gloniopsis arciformis GKM L166A
Diplodia mutila CBS 431.82
Brachiosphaera tropicalis E192-1
Arthrographis curvata CBS 135934
Botryosphaeria dothidea CBS 115476
Aliquandostipite khaoyaiensis CBS 118232
Rhytidhysteron rufulum CBS 306.38
Asterotexis cucurbitacearum VI 24814
Rhexothecium globosum CBS 955.73
Jahnula appendiculata SS2900
Byssosphaeria salebrosa SMH 2387
Asterotexis cucurbitacearum PMM 0141224
Arthrographis longispora CBS 135935
Herpotrichia juniperi AFTOL-ID 1608
Dissoconium aciculare CBS 204.89
Pirozynskiella laurisilvatica CBS 138109
Phoma herbarum CBS 567.63
Neofusicoccum nonquaesitum CBS 126655
Arthrographis kalrae CBS 693.77
Alternaria tenuissima CBS 918.96
Tubeufia paludosa CBS 245.49
Neofusicoccum mangiferae CBS 118532
Macrophomina phaseolina CBS 227.33
Schizothyrium pomi CBS 486.50
Hysterium angustatum CBS 123334
Xiliomyces brasiliensis URM 7413
Uwebraunia communis CBS 110747
Schizothyrium pomi CBS 228.57
JW209002
Helicomyces roseus CBS 283.51
Glonium circumserpens CBS 123343
Psiloglonium simulans CBS 206.34
Byssosphaeria jamaicana SMH 1403
Arthrographis arxii CBS 203.78
Hysteropatella prostii H-9934b
Jahnula aquatica R68-1
Melanomma pulvis-pyrius CBS 124080
Hysteropatella clavispora CBS 247.34
Glonium circumserpens CBS 123342
Pleospora herbarum var. herbarum CBS 191.86
Bezerromyces brasiliensis URM 7411
Bezerromyces pernambucoensis URM 7412
JW49012
Jahnula seychellensis SS2113-1
Arthrographis grakistii sp. nov.
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96
100
95
85
100
98
88
100
96
96
75
94
98
97
100
90
90
99
100
99
100
76
100
100
80
100
Fig. 1. Maximum composite likelihood tree obtained from the RAxML analysis of the LSU sequence alignment of selected Dothideomycetes. Bootstrap
support values above 70 % are shown at the nodes. Families and orders are indicated with coloured blocks to the right of the tree. Taxonomic
noveles described in this study are indicated in boldface. The tree was rooted to Yarrowia hollandica (CBS 4855).
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
10
Teratosphaeriaceae
Cladosporiaceae
Capnodiaceae
Asterinaceae
Patelleriaceae
Cladoriellaceae
Sympoventuriaceae
Venturiaceae
Phaeotrichaceae
Mycosphaerellaceae
0.2
CAPNODIALES
PATELLARIALES
II
ASTERINALES
CLADORIELLA-
LES
VENTURIALES
PHAEOTRICHA-
LES
Fusicladium ramoconidii CBS 462.82
Aulographina eucalypti CPC 12986
Verruconis calidifluminalis CBS 125818
Blastacervulus eucalypti CBS 124759
Melnikomyces thailandicus sp. nov. CBS 145767
Blastacervulus robbenensis CBS 124780
Ochroconis constricta CBS 202.27
Leptoxyphium fumago CBS 123.26
Venturia inaequalis CBS 176.42
Cladoriella rubrigen CBS 124760
Saccharomyces cerevisiae DAOM 216365
Phaeocryptopus gaeumannii CBS 267.37
Scolecobasidiella avellanea CBS 772.73
Cladosporium halotolerans CBS 127371
Verruconis gallopava CBS 547.81
Yarrowia hollandica CBS 4855
Cladoriella xanthorrhoeae CBS 143398
Patellaria cf. atrata BCC 28876
Asterina chrysophylli VI 42823
Clavatispora thailandica MFLUCC 10-0107
Venturia populina CBS 256.38
Neocoleroa metrosideri PDD 107531
Apiosporina collinsii CBS 118973
Asterina melastomatis VI 42822
Teratosphaeria destructans CBS 111369
Teratosphaeria fibrillosa CBS 121707
Capnodium coffeae CBS 147.52
Clavatispora thailandica MFLUCC 17-2237
Trichodelitschia bisporula CBS 262.69
Cladosporium variabile CBS 121636
Patellaria cf. atrata BCC 28877
Cladoriella paleospora CBS 124761
Melnikomyces vietnamensis CBS 136209
Alysidiella parasitica CBS 120088
Verruconis thailandica sp. nov. CBS 145768
Ochroconis gamsii CBS 239.78
Cladoriella kinglakensis CPC 32730
Fusicladium pini CBS 463.82
Cladoriella eucalypti CBS 115899
Apiosporina morbosa N/A
Sympoventuria capensis CBS 120136
Zasmidium cellare CBS 146.36
Phaeotrichum benjaminii CBS 541.72
Gibbera conferta CBS 191.53
Verruconis verruculosa CBS 119775
Teratosphaeria stellenboschiana CBS 116428
Ramularia punctiformis CBS 113265
Heteroconium eucalypti CBS 120122
Verruconis gallopava CBS 437.64
87
97
100
74
100
100
92
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100
80
91
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92
92
83
100
82
91
92
99
100
91
99
82
100
100
84
100
99
99
100
91
93
100
100
100
Aureoconidiella foliicola gen. et sp. nov. CBS 145943 Aureoconidiellaceae fam. nov. AUREOCONIDIELLALES
ORD. NOV.
Fig. 1. (Connued).
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
11
Sordariaceae
Chaetosphaeriaceae
Helminthosphaeriaceae
Boliniaceae
Lasiosphaericeae
Coniochaetaceae
Valsaceae
Diaporthaceae
Nectriaceae
Bionectriaceae
Xylariaceae
Diatrypaceae
Ruzenia spermoides SMH4606
Helminthosphaeria cf. stuppea JF04120
Hilberina caudata SMH1542
Helminthosphaeria triseptata JF04015
Echinosphaeria canescens SMH4666
Echinosphaeria canescens TL5730
Valsonectria pulchella SMH1193
Helminthosphaeria clavariarum SMH4609
Echinosphaeria canescens JHC97 006
Helminthosphaeria corticiorum JF04225
Synaptospora plumbea ANM963
Camarops tubulina SMH4614
Ruzenia spermoides SMH4655
Chaetosphaeria ovoidea SMH2605
Synaptospora plumbea SMH3962
Helminthosphaeria tomaculum SMH2485
Helminthosphaeria cf. stuppea TL11998
Helminthosphaeria odontiae ANM928
Fusarium ambrosium SMH1999
Coniochaeta discoidea SANK 12878
CBS 146133
Camarops ustulinoides SMH1988
Helminthosphaeria carpathica SMH3903
Orbilia auricolor AFTOL-ID 906
Orbilia vinosa AFTOL-ID 905
Eutypa sp SMH3580
Sporoschisma hemipsila SMH2125
Diaporthe phaseolorum FAU458
Coniochaetidium savoryi TRTC 51980
Hilberina munkii SMH1531
Ruzenia spermoides ANM163
Echinosphaeria canescens SMH4791
Cytospora ceratosperma AR3426
Neurospora crassa MUCL 19026
COAD 2632
Lasiosphaeria ovina SMH 1538
Anthostomella sp. SMH3101
CBS 146338
CBS 146339
Kramasamuha sibiki
0.06
100
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100
77
86
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100
97
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96
92
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76
100
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100
72
SORDARIALES II
HYPOCREALES
XYLARIALES
SORDARIALES I
CHAETOS-
PHAERIALES
BOLINIALES
CONIOCHAETA-
LES
DIAPORTHALES
Fig. 2. Maximum composite likelihood tree obtained from the RAxML analysis of the LSU sequence alignment of selected Sordariomycetes. Bootstrap
support values above 70 % are shown at the nodes. Families and orders are indicated with coloured blocks to the right of the tree. Included strains
described in this study are indicated in boldface. The tree was rooted to Orbilia vinosa (AFTOL-ID 905) and Orbilia auricolor (AFTOL-ID 906).
Aureoconidiella foliicola Hern.-Restr. & Crous, sp. nov.
MycoBank MB833916. Fig. 4.
Etymology: The epithet “foliicola” refers to its habitat on a dead
leaf.
Mycelium consisng of septate, smooth, brown, 1–2.5 µm wide
hyphae. Conidiophores macronematous, simple, septate, brown,
40–85 × 3–5.5 µm. Conidiogenous cells integrated, terminal,
polyblasc, with thickened scars, brown to pale brown, 25–53
× 3–4.5 µm. Conidia globose to subglobose with apiculate base,
inially subhyaline, golden brown at maturity, verrucose, 5–8
µm diam, base 1–2 µm wide.
Culture characteriscs: On MEA and OA surface coony aerial
mycelium Fawn, sepia to dark brick close to the agar, margin
euse, enre; reverse sepia to black.
Typus: South Africa, KwaZulu-Natal, Richards Bay, on living
leaves of Syzygium cordatum (Myrtaceae), Jun. 2016, M.J.
Wingeld (holotype CBS H-24099, culture ex-type CPC 36154 =
CBS 145943).
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
12
0.03
Cyphellophora sessilis CBS 243.85
Veronaea japonica CBS 776.83
Exophiala xenobiotica CBS 115831
“Minimelanolocus curvatus” MFLUCC 15-0259
Exophiala jeanselmei CBS 507.90
Veronaea botryosa MFLUCC 11-0072
Capronia pilosella AFTOL-ID 657
“Minimelanolocus submersus” KUMCC 15-0206
Rhinocladiella anceps CBS 181.65
“Minimelanolocus obscurus” MFLUCC 15-0416
“Minimelanolocus aquaticus” MFLUCC 15-0414
Exophiala nigra dH 12,296
Rhinocladiella anceps AFTOL-ID 659
Exophiala salmonis CBS 157.67
“Minimelanolocus melanicus” MFLUCC 15-0415
Exophiala salmonis AFTOL-ID 671
Fonsecaea monophora CBS 102243
Phialophora americana AFTOL-ID 658
Cyphellophora oxyspora CBS 698.73
Thysanorea seifertii sp. nov. CBS 145910
Phialophora verrucosa AFTOL-ID 670
Cladophialophora parmeliae CBS 129337
Melanoctona tectonae MFLUCC 12-0389
“Minimelanolocus asiaticus” MFLUCC 15-0237
Cladophialophora subtilis CBS 122642
Veronaea compacta CBS 268.75
Cladophialophora minourae CBS 556.83
Exophiala pisciphila AFTOL-ID 669
Veronaea botryosa CBS 254.57
Thysanorea papuana MFLUCC 15-0966
Thysanorea papuana CBS 212.96
“Minimelanolocus thailandensis” MFLUCC 15-0971
Aculeata aquatica MFLUCC 11-0529
“Pseudospiropes lotorum” CBS 235.78
Exophiala pisciphila CBS 537.73
Cladophialophora carrionii CBS 160.54
“Minimelanolocus nonramosus” MFLUCC 17-2378
“Minimelanolocus rousselianus” CBS 126086
97
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89
73
86
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100
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98
73
88
100
100
75
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99
100
Herpotrichiellaceae
Thysanorea
Thysanorea cantrelliae sp. nov. CBS 145909
Fig. 3. Maximum composite likelihood tree obtained from the RAxML analysis of the ITS/LSU/SSU sequence alignment of selected Herpotrichiellaceae.
Bootstrap support values above 70 % are shown at the nodes. Taxonomic noveles described in this study are indicated in boldface. The tree was
rooted to Cyphellophora oxyspora (CBS 698.73) and Cyphellophora sessilis (CBS 243.85).
Notes: This new lineage is introduced to accommodate a fungus
characterised by unbranched conidiophores, cicatrised and
sympodial conidiogenous cells with thickened scars, producing
sub-globose, verruculose, and golden brown conidia. Other
related lineages are those accommodang Asterinales and
Cladoriellales (Fig. 1). However, they dier from those in the
Aureoconidiellales based on the morphology of the asexual
morphs. The Asterinales is mainly characterised by taxa that are
coelomycetes with pycnothyrial conidiomata (Guamosim et
al. 2015, Jaklitsch et al. 2016). The Cladoriellales is a monotypic
order related to cladosporium-like hyphomycetous fungi with
conidia frequently remaining aached in long acropetal chains
(Crous et al. 2006c, 2017).
Authors: M. Hernández-Restrepo, P.W. Crous and M.J. Wingeld
Arthrographis Sigler & J.W. Carmich., Mycotaxon 4: 359. 1976.
Synonym: [Arthrographis G. Cochet, Annls Parasit. Hum. Comp.
17: 97. 1939. (Nom. inval., Art. 39.1)]
Vegetave hyphae septate, hyaline, smooth- and thin-walled.
Conidiophores macro- or micronematous, erect, simple or
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
13
poorly branched, hyaline, smooth-walled. Conidiogenous
hyphae simple or branched, thin-walled, forming septa
basipetally to form arthroconidia released by schizolyc
secession. Arthroconidia unicellular, cylindrical or cuboid,
straight, subhyaline, thick- and smooth-walled. Synasexual
morph trichosporiella-like, with conidia growing directly on
undierenated hyphae, sessile, lateral, terminal, globose,
subglobose or clavate, subhyaline, thin- and smooth-walled.
Sexual morph not observed.
Type species: Arthrographis kalrae (R.P. Tewari & Macph.) Sigler
& J.W. Carmich.
Arthrographis kalrae (R.P. Tewari & Macph.) Sigler & J.W.
Carmich., Mycotaxon 4: 360. 1976.
Basionym: Oidiodendron kalrae R.P. Tewari & Macph., Mycologia
63: 603. 1971 [as ‘kalrai’].
Synonym: [Arthrographis langeronii G. Cochet (as ‘langeroni’),
Annls Parasit. hum. comp. 17: 97. 1939. (Nom. inval., Art. 39.1)]
Descripons and illustraons: Sigler & Carmichael (1976, 1983),
Giraldo et al. (2014a).
Specimens examined: The Netherlands, Utrecht, isolated from
soil, 2017, E. Kieviet (JW 21004 = CBS 145527), ibid. JW 21008,
ibid. JW 21029.
Arthrographis grakisi Giraldo López & Hern.-Restr., sp. nov.
MycoBank MB833677. Fig. 5.
Etymology. Named aer Ewan Grakist, who collected the soil
sample. This species was discovered as part of a Cizen Science
project in the Netherlands.
Fig. 4. Aureoconidiella foliicola gen. et sp. nov. (CBS 145943). A. Conidiophores and conidia. B. Conidiophore with conidiogenous cell. C, D.
Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars: A = 20 μm, all others = 10 μm.
Fig. 5. Arthrographis grakisi sp. nov. (CBS 145530). A. Poorly branched conidiophores and arthroconidia. B, C. Trichosporiella-like synasexual morph.
Scale bars = 10 μm.
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
14
Vegetave hyphae septate, hyaline, smooth- and thin-
walled, 1.5–2 μm wide. Conidiophores semi-macronematous
or micronematous, erect, simple, hyaline, smooth-walled.
Conidiogenous hyphae simple or branched, 1.5–2 μm wide, thin-
walled, forming septa basipetally to form arthroconidia released
by schizolyc secession. Arthroconidia unicellular, cylindrical or
cuboid, straight, 2–4 × 2–3 μm, subhyaline, thick- and smooth-
walled. Synasexual morph trichosporiella-like with conidia
growing directly on undierenated hyphae, sessile, somemes
with a subcylindrical to clavate intercalary cell, lateral, terminal,
globose, subglobose or clavate, 3–5 × 2–3 μm, subhyaline, thin-
and smooth-walled. Sexual morph not observed.
Culture characteriscs: Colonies at 25 °C aer 14 d: on OA
reaching 13–14 mm, at, glabrous to occose, surface and
reverse bu. On MEA and PDA reaching 11–13 mm and
12–17 mm, respecvely; at or raised, dusty to coony at
centre, glabrous toward the periphery, bu to honey, reverse
uncoloured. No growth at 37 °C.
Typus: The Netherlands, Utrecht Province, Wijk bij Duurstede,
from garden soil, 2017, E. Grakist (holotype CBS H-23912,
culture ex-type CBS 145530 = JW 190014).
Addional materials examined: The Netherlands, Utrecht Province,
IJsselstein, from garden soil, 2017, J. Brus (JW 209002, JW 209003);
from garden soil, 2017, R. de Bruyn (JW 180011); Utrecht, from garden
soil, 2017, M. Wickham (JW 199018); Zeeland Province, Vlissingen,
from garden soil, 2017, N. Penabad (CBS 145529 = JW 22011, JW 22015,
JW 22019); Gelderland Province, Zaltbommel, from garden soil, 2017,
K. & T. de Man (JW 49011, JW 49012).
Notes: Based on a BLAST search using the ITS and LSU loci
several soil isolates (JW isolates listed in Table 1) were idened
as belonging to Arthrographis. In order to conrm their identy
at the species level, a combined analysis of the LSU/ITS/act loci
was performed, including all members from Eremomycetaceae.
The ML tree (Fig. 6) showed that three isolates (JW 21004,
JW 21008 and JW 21029) grouped with the type species of A.
kalrae (CBS 693.77) and one (JW 22007) with A. longispora (CBS
135935). However, most of the isolates formed a well-supported
clade that represents the new species A. grakisi.
The phylogenec analyses showed that A. grakisi is closely
related to A. longispora (Figs 1, 6). The laer species, however,
has longer and narrower arthroconidia [5–10(–13) × 1–1.5 μm
in A. longispora vs. 2–4 × 2–3 μm in A. grakisi] and does not
produce the trichosporiella-like synasexual morph in culture
(Giraldo et al. 2014a).
Morphologically, A. grakisi resembles A. kalrae and A.
curvata in having cylindrical arthroconidia and a trichosporiella-
like synasexual morph. Arthrographis kalrae and A. curvata are
able to grow at 37 °C (Sigler & Carmichael 1983, Giraldo et al.
2014a), while A. grakisi does not grow at this temperature.
Authors: A. Giraldo López and M. Hernández-Restrepo
Kramasamuha Subram. & Vial, Canad. J. Bot. 51: 1128. 1973.
Conidiophores erect, exuous, solitary to fasciculate, arising
from a swollen basal cell, which appears lobed due to rhizoids;
medium brown, smooth, mul-septate, simple or branched,
giving rise to parallel spes, becoming paler towards apex,
terminang in an acute conidiogenous cell. Conidiogenous
cells monoblasc, pale brown, ampulliform, straight to curved,
tapering to a truncate apex, thin-walled, solitary or in clusters,
integrated or discrete, terminal and intercalary. Conidia solitary,
smooth, septate, obovoid to pyriform; second cell from base
thick-walled, dark brown, somewhat swollen, basal and apical
cell subhyaline, with short narrow separang cell at base as
remnant from conidiogenous cell.
Type species: Kramasamuha sibika Subram. & Vial
Kramasamuha sibika Subram. & Vial, Canad. J. Bot. 51: 1129.
1973. Fig. 7.
Typus: India, Tamil Nadu, Chingleput district, Vandalur, on dead
leaves of Gymnosporia emarginata (Celastraceae), 23 Apr. 1971,
B.P.R. Vial, Herb. MUBL 2153 (not seen).
Occurring on leaf lier. Conidiophores erect, exuous, solitary
to fasciculate, arising from a swollen basal cell, 10–14 µm diam,
which appears lobed due to rhizoids; medium brown, thin-
walled, smooth, mul-septate, septa 17–30 µm apart, up to 550
µm tall, 3–5 µm diam, unbranched or branched, giving rise to
parallel spes, becoming paler towards apex, terminang in an
acute conidiogenous cell. Conidiogenous cells monoblasc, pale
brown, ampulliform, straight to curved, tapering to a truncate
apex, thin-walled, solitary or in clusters of 2–4, integrated or
discrete, terminal and intercalary, 4–8 × 3–4 µm. Conidia solitary,
smooth, (1–)2(–3)-septate, blastoconidia, obovoid to pyriform,
apex obtuse, (18–)25–27(–34) × (10–) 11(–12) µm; second cell
from base thick-walled, dark brown, somewhat swollen, basal
and apical cell subhyaline, with short narrow separang cell at
base as remnant from conidiogenous cell, 1–1.5 × 1 µm.
Culture characteriscs: Colonies on OA with scarce aerial
mycelium, coony to velvety, grey olivaceous, submerged
mycelium vinaceous, margin euse, irregular, reverse vinaceous.
Conidiophores, conidiogenous cells, and conidia very similar to
those observed in natural substrate. Conidiogenous cells 4.5–9 ×
2.5–4 µm. Conidia 14–32.5 × 9–14 µm, (0–)2(–3)-septate.
Materials examined: Australia, New South Wales, Mallanganee,
on leaves of Lophostemon confertus (Myrtaceae), 17 Apr. 2018,
A.J. Carnegie, CBS 146338. Brazil, state of Minas Gerais, Viçosa, on
Hypericum innodorum (Hypericaceae) leaves bearing necroc spots
caused by Seimatosporium hypericinum (Pinaceae), 4 Jul. 2017,
A.A. Colmán (VIC 47176, COAD 2632). Malaysia, on needles of Pinus
tecunumanii, Oct. 2018, M.J. Wingeld, CBS 146339. South Africa,
KwaZulu-Natal, Richards Bay, on living leaves of Syzygium cordatum
(Myrtaceae), Jun. 2016, M.J. Wingeld, CBS 146133 = CPC 36153.
Notes: Kramasamuha resembles the genus Garnaudia and
some species of Endophragmiella in having conidia with a
short narrow separang cell at the base as remnant from
the conidiogenous cell. However, species in these genera
can be disnguished by the arrangement and colour of the
conidiogenous cells. In Kramasamuha they are pale brown and
solitary or in clusters along the conidiophores (Subramanian &
Vial 1973). In Garnaudia, the conidiogenous cells are brown,
and vercillate in terminal branches (Borowska 1977), while
in Endophragmiella the conidiogenous cells are hyaline and
mainly solitary and terminal (Hughes 1979, Seifert et al. 2011).
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
15
Kramasamuha is a monotypic genus originally described from
India on Gymnosporia emarginata (Subramanian & Vial 1973).
However, the specimens examined here were from dierent
substrates and connents, i.e. Lophostemon confertus in
Australia, Hypericum×innodorum in Brazil, Pinus tecunumanii in
Malaysia, and Syzygium cordatum in South Africa and can thus
not serve as an epitype. This species has also been recorded
from leaves on Feijoa sellowiana in New Zealand, and on Psidium
guajava in Western Samoa (Landcare database 2019).
This is the rst me that DNA sequence data has become
available for K. sibika and shows that Kramasamuha is related to
Helminthosphaeriaceae (Sordariomycetes, Fig. 2). Asexual morphs
in Helminthosphaeria have been recognised as diplococcium-like
(Samuels et al. 1997, Réblová 1999) with trec conidiogenous
Eremomycetaceae, Eremomycetales
0.02
JW 22011
Arthrographis longispora JW 22007
Arthrographis kalrae JW 21008
Eremomyces bilateralis CBS 781.70T
JW 49011
JW 49012
JW 22015
Arthrographis kalrae JW 21029
Arthrographis kalrae JW 21004
JW 199018
JW 22019
JW 209002
Arthrographis longispora CBS 135935T
Arthrographis curvata CBS 135933T
Arthrographis curvata CBS 135934
Arthrographis kalrae CBS 693.77T
JW 209003
Arthrographis grakistii sp. nov.
JW 190014T
Arthrographis globosa CBS 135397T
Arthrographis arxii CBS 203.78T
JW 180011
Arthrographis chlamydospora CBS 135396T
Rhexothecium globosum CBS 955.73T
100
100
100
100
98
100
100
100
98
86
Fig. 6. Maximum composite likelihood tree based on paral sequences from the LSU, ITS and act regions from all members of Eremomycetales. Colour
boxes (blue and green) indicate the generic clades. Bootstrap support values above 70 % are shown at the nodes. Ex-type strains are in boldface. T =
Ex-type.
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
16
Fig. 7. Kramasamuha sibika (CBS 146133). A–C. Conidiophores overview on OA. D–F. Conidiophores, conidiogenous cells and conidia. G. Conidiogenous
cells and conidia. H–J. Conidiogenous cells. K–N. Conidia. Scale bars: D–F = 25 μm, all others = 10 μm.
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
17
cells, diering from the monoblasc conidiogenous cells observed
in Kramasamuha (Seifert et al. 2011).
Authors: M. Hernández-Restrepo, P.W. Crous, M.J. Wingeld,
A.A. Colmán, P.S.C. Mansur and R.W. Barreto
Melnikomyces Crous & U. Braun, Persoonia 32: 263. 2014.
Mycelium consisng of brown, septate, branched, smooth,
thick-walled hyphae. Conidiophores subcylindrical, brown,
smooth, erect, straight or geniculate, reduced to conidiogenous
cells, or long, exuous, mulseptate. Conidiogenous cells
polyblasc, subcylindrical to subclavate, terminal or intercalary,
brown, smooth, developing a rachis with numerous dencle-
like loci. Conidia solitary, brown, verruculose, fusoid-ellipsoidal,
1-septate, ends sub-obtuse, released by rhexolyc secession.
Chlamydospores terminal, globose to subglobose, in short
chains, simple or branched, brown, smooth (modied from
Crous et al. 2014b).
Type species: Melnikomyces vietnamensis Crous & U. Braun
Melnikomyces thailandicus Giraldo López, sp. nov. MycoBank
MB833678. Fig. 8.
Etymology: Name refers to Thailand where the fungus was
collected.
Mycelium consisng of brown, septate, branched, smooth, thick-
walled, 2–2.5 μm diam hyphae. Conidiophores macronematous,
arising directly from vegetave hyphae, erect, straight or
slightly bent, simple, mulseptate, cylindrical, 14–37 × 2–3 μm,
brown, paler apex, thick and smooth-walled. Conidiogenous
cells integrated, terminal, polyblasc, brown to pale brown,
sympodial, with long open dencles; dencles cylindrical,
pale brown, up to 1 μm long. Conidia fusoid, ends subobtuse,
1-septate, solitary, subhyaline, smooth-walled, (8–)9.5–12(–
13) × (2–)2.5(–3) μm. Chlamydospores lateral, globose to
subglobose, in short and simple chains, light brown, thick-and
smooth-walled, 5.5–10 μm diam.
Typus: Thailand, Nakhon Nayok Province, Mueang Nakhon Nayok
district, Wang Takrai waterfall, N14.330023° E101.307168°, 64
m above sea level, from soil, 22 Jul. 2008, P.W. Crous (holotype
CBS H-24236, culture ex-type CBS 145767).
Notes: The monotypic genus Melnikomyces was introduced by
Crous and Braun (Crous et al. 2014b), based on M. vietnamensis
collected from dry leaves in Vietnam. This species was treated as
incertae sedis in the Chaetothyriales, Euroomycetes (Crous et
al. 2014b). However, the results of this study show that it resides
in the Sympoventuriaceae (Venturiales, Dothideomycetes),
together with other genera producing septate conidia from
denculate conidiogenous cells, such as Ochroconis and
Verruconis (Machouart et al. 2014, Samerpitak et al. 2014).
Melnikomyces thailandicus is the second species described in
the genus, which diers morphologically from M. vietnamensis
in having shorter conidiophores (10–60 μm long vs.14–37 μm
long) and longer and narrower smooth-walled conidia (8–13 ×
(2 –3) μm vs. 7–11 × 2.5–3.5 μm). Originally, M. vietnamensis
was described with two types of conidiophores (Crous et al.
2014b), but in M. thailandicus one of the conidiophore types
more closely resembles chlamydospores, as they appear to stay
aached to the hyphae.
Authors: A. Giraldo López and P.W. Crous
Thysanorea Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 80.
2007. emend.
Conidiophores micro- or macronematous, erect, simple or
apically branched, somemes proliferang percurrently in
the apex, brown, smooth. Conidiogenous cells terminal or
intercalary, polyblasc, smooth, brown at the base, paler
towards the apex, subcylindrical, clavate to doliiform, sympodial,
with crowded conidiogenous loci inconspicuous to slightly
prominent, refracve to somewhat obscure, slightly thickened.
Conidia solitary, oblong, obovoid, cylindrical, broadly fusiform
to subpyriform, pale brown, smooth, with a narrowly truncate
base and darkened hilum; conidial secession schizolyc.
Synasexual morph: Conidiophores erect, simple, brown, smooth.
Fig. 8. Melnikomyces thailandicus sp. nov. (CBS 145767). A–C. Conidiophores and denculated conidiogenous cells. D. Conidia. E. Chlamydospores.
Scale bars = 10 μm.
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
18
Conidiogenous cells terminal, discrete, phialidic, subglobose to
lageniform, with a balloon- to funnel-shaped collaree, brown,
oen in clusters at the apex of the conidiophores. Conidia
solitary, subglobose to obovate, unicellular, hyaline, guulate,
smooth.
Type species: Thysanorea papuana (Aptroot) Arzanlou et al.
Notes: Thysanorea was established for a genus similar to
Periconiella based on the branching paern of the conidiophores
(Arzanlou et al. 2007). However, recent studies have shown that
those branching paerns depend on culture condions, where
those on natural substrates or in young cultures are not as
prominently branched as previously described (Kirschner 2016,
Wang et al. 2019). The generic descripon is emended here to
include species with a phialidic synasexual morphs.
Phylogenecally, Thysanorea is closely related to
Minimelanolocus in the Chaetothyriales (Fig. 3). However,
the phylogenec posion of M. navicularis, the type species,
remains unknown since DNA sequence data are not available for
it, and the supposed phylogenec posion has been based on
other species (Liu et al. 2015, Wang et al. 2019).
Morphologically, M. navicularis is characterised by terminal
conidiogenous cells that produce navicular conidia with sub-
hyaline polar cells and darker central cells (Castañeda-Ruiz
1987). They consequently dier from those species placed in
Minimelanolocus based on DNA sequence data and in which
the conidiogenous cells are terminal and intercalary, and the
conidia are oblong, obovoid, cylindrical, broadly fusiform, and
uniformly pale brown (Liu et al. 2015, Wang et al. 2019). In
this regard, they would t beer with the generic concept of
Thysanorea (Arzanlou et al. 2007, Wang et al. 2019). Based on
these morphological dierences and phylogenec relaonships,
we propose new combinaons for those species that have been
shown as related to Thysanorea. The phylogenec placement of
other species for which DNA sequence data are not available,
including M. navicularis must sll be determined.
Thysanorea asiaca (H.Y. Su, et al.) Hern.-Restr. & Crous, comb.
nov. MycoBank MB833919.
Basionym: Minimelanolocus asiacus H.Y. Su, et al., Fungal Biol.
119: 1054. 2015.
Thysanorea cantrelliae Hern.-Restr., R. van Doorn & Crous, sp.
nov. MycoBank MB833914. Fig. 9.
Etymology: Named in honour of Sharon Cantrell, who was the
organizer of the IMC 11 (2018) in Puerto Rico. This fungus was
collected on a eld trip held during the IMC 11.
Mycelium composed of hyaline to pale brown, septate, smooth,
1–2 µm wide hyphae. Conidiophores semi-micronematous,
somemes reduced to conidiogenous cells, simple, erect,
straight or exuous, cylindrical, pale brown, smooth, 8–31 ×
2–3 µm. Conidiogenous cells holoblasc, polyblasc, mainly
terminal, integrated, sympodial, pale brown, 7–22 × 2–3
µm. Conidia solitary, fusiform to acicular, straight or curved,
(1–)3(–4)-septate, subhyaline to pale brown, smooth, 10–34 ×
1.5–3 µm, apex acute, base truncate.
Culture characteriscs: Colonies at 25 °C aer 14 d: on OA
reaching 22–25 mm, aerial mycelium moderate, coony
to occose, olivaceous black, margin euse enre; reverse
black. On MEA and PDA reaching 22–30 mm, aerial mycelium
abundant, coony to occose, olivaceous grey, black close to
the agar, margin euse, enre; reverse black.
Typus: USA, Puerto Rico, from unidened twig, 20 Jul. 2018,
M. Hernández-Restrepo (holotype CBS H-24100, culture ex-type
CBS 145909).
Notes: Thysanorea cantrelliae clustered in a separate clade
together with T. seiferi (Fig. 3). It can be disnguished from T.
seiferi and other species in the genus by its acicular conidia
(Fig. 9).
Thysanorea curvata (H.Y. Su et al.) Hern.-Restr. & Crous, comb.
nov. MycoBank MB833921.
Basionym: Minimelanolocus curvatus H.Y. Su et al., Fungal Biol.
119: 1055. 2015.
Thysanorea lotorum (Morgan-Jones) Hern.-Restr. & Crous,
comb. nov. MycoBank MB833922.
Basionym: Pseudospiropes lotorum Morgan-Jones, Mycotaxon
5: 481. 1977 [as ‘lotorus’]
Synonym: Nigrolenlocus lotorum (Morgan-Jones) R.F.
Castañeda & Heredia, Cryptog. Mycol. 22: 15. 2001.
Minimelanolocus submersus Z.L. Luo et al., Fungal Diversity 80:
143. 2016.
Thysanorea melanica (H.Y. Su, et al.) Hern.-Restr. & Crous, comb.
nov. MycoBank MB833923.
Basionym: Minimelanolocus melanicus H.Y. Su et al., Fungal Biol.
119: 1056. 2015.
Thysanorea nonramosa (X.D. Yu et al.) Hern.-Restr. & Crous,
comb. nov. MycoBank MB833924.
Basionym: Minimelanolocus nonramosus X.D. Yu et al., Mycol.
Progr. 18: 514. 2019.
Thysanorea obscura (Matsush.) Hern.-Restr. & Crous, comb.
nov. MycoBank MB833925.
Basionym: Pseudospiropes obscurus Matsush., Matsushima
Mycol. Mem. 3: 14. 1983.
Synonym: Minimelanolocus obscurus (Matsush.) R.F. Castañeda
& Heredia, Cryptog. Mycol. 22:10. 2001.
Thysanorea rousseliana (Mont.) Hern.-Restr. & Crous, comb.
nov. MycoBank MB833926.
Basionym: Helminthosporium rousselianum Mont., Ann. Sci.
Nat., Sér. 3, Bot., 12: 300. 1849.
Synonyms: Pleurophragmium rousselianum (Mont.) S. Hughes,
Canad. J. Bot. 36: 798. 1958.
Spiropes rousselianus (Mont.) de Hoog & Arx, Kavaka 1: 59.
1973.
Pseudospiropes rousselianus (Mont.) M.B. Ellis, More
Demaaceous Hyphomycetes: 221. 1976.
Minimelanolocus rousselianus (Mont.) R.F. Castañeda & Heredia,
Cryptog. Mycol. 22: 10. 2001.
Thysanorea seiferi Hern.-Restr., R. van Doorn & Crous, sp. nov.
MycoBank MB833920. Fig. 10.
Etymology: Named in honour of Prof. dr Keith A. Seifert, who was
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
19
Fig. 9. Thysanorea cantrelliae sp. nov. (CBS 145909). A. Conidiophore and conidium. B, C. Conidiophores. D–G. Conidiogenous cells. H, I. Conidia.
Scale bars = 10 μm.
President of the Internaonal Mycological Associaon during
the IMC 11 (2018) in Puerto Rico. This fungus was collected on a
eld trip during the IMC 11.
Mycelium composed of hyaline to pale brown, septate, smooth,
1–2 µm wide hyphae. Conidiophores mononematous, simple,
erect, straight or exuous, cylindrical, brown, paler towards
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
20
the apex, smooth, 30–133 × 2–3 µm. Conidiogenous cells
holoblasc, polyblasc, terminal or intercalary, integrated,
sympodial, pale brown to brown, 7.5–46.5 × 2–3 µm. Conidia
solitary, subcylindrical to clavate or oblong, straight or slightly
curved, (0–)1–3(–4)-septate, pale brown, smooth, 7–15 × 1.5–3
µm, apex rounded, base darkened and truncated. Synasexual
morph: Conidiophores micro- or macronematous, erect, straight
or exuous, cylindrical, brown, smooth, 10–51 × 2–4 µm.
Conidiogenous cells enteroblasc, phialidic, arranged around the
apex of the conidiophore, brown, subglobose to ampulliform,
2–4(–6) × 2–3 µm, with a balloon- to funnel-shaped collaree,
1–3 × 1–3 µm. Conidia solitary, subglobose to obovate, unicellular,
hyaline, guulate, smooth, 1–2 × 1–1.5 µm, base truncated.
Culture characteriscs: Colonies at 25 °C aer 14 d: on OA
reaching 22–25 mm, aerial mycelium moderate, coony to
occose, grey olivaceous to olivaceous black, margin euse,
enre; reverse black. On MEA and PDA reaching 18–20 mm,
aerial mycelium moderate to abundant, coony to occose,
olivaceous grey, black close to the agar, margin euse, enre;
reverse black.
Typus: USA, Puerto Rico, from unidened twig, 20 Jul. 2018,
M. Hernández-Restrepo (holotype CBS H-24101, culture ex-type
CBS 145910).
Notes: Some of the conidia in T. seiferi resemble those of T.
obscura in being 3-septate. However, conidia in T. seiferi are
smaller than those of T. obscura (7–15 × 1.5–3 μm vs. 20–31
× 5–8 μm, Castañeda-Ruiz et al. 2001). Thysanorea seiferi
is the only species in the genus known to produce a phialidic
synasexual morph.
Fig. 10. Thysanorea seiferi sp. nov. (CBS 145910). A. Conidiophores and conidia. B, C. Conidiogenous cells and conidia. D. Conidia. E–J. Synasexual
morph. E, F. Conidiophores. G–I. Conidiogenous cells and conidia. H. Conidia. Scale bars: G–J = 5 μm, all others = 10 μm.
© 2020 Westerdijk Fungal Biodiversity Instute
Genera of Fungi – G6
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
21
Thysanorea thailandensis (W. Dong et al.) Hern.-Restr. & Crous,
comb. nov. MycoBank MB833927.
Basionym: Minimelanolocus thailandensis W. Dong et al., Mycol.
Progr. 17: 622. 2018.
Thysanorea yunnanensis Hern.-Restr. & Crous, nom. nov.
MycoBank MB833928.
Replaced synonym: Minimelanolocus aquacus H.Y. Su et al.,
Fungal Biol. 119: 1049. 2015 [non Thysanorea aquaca W. Dong,
H. Zhang & K.D. Hyde, 2018].
Etymology: The name refers to the Chinese Province of Yunnan
where the fungus was collected.
Authors: M. Hernández-Restrepo, R. van Doorn and P.W. Crous
Verruconis Samerp. et al., Fungal Diversity 65: 117. 2014.
Mycelium consisng of septate, pale brown, smooth and thick-
walled hyphae. Conidiophores dierenated, erect, straight
or slightly bent, unbranched, pale brown. Conidiogenous cells
mostly polyblasc, subcylindrical to narrowly mucronate,
producing conidia sympodially on long open dencles; dencles
cylindrical, pale brown, scaered at the apical third of the
conidiogenous cell. Conidia two-celled, ellipsoidal, cylindrical
or clavate, brown, verrucose or smooth-walled, released by
rhexolyc secession. Sexual morph unknown.
Type species: Verruconis gallopava (W.B. Cooke) Samerp. & de
Hoog
Verruconis thailandica Giraldo López & Crous, sp. nov.
MycoBank MB833679. Fig. 11.
Etymology: The name refers to Thailand where the fungus was
collected.
Mycelium consisng of septate, pale brown, smooth, thick-
walled, 2–2.5 μm diam hyphae. Conidiophores dierenated,
arising directly from vegetave hyphae, erect, straight or slightly
bent, simple, 0–1-septate, subcylindrical, (3.6–)4.1–7.1(–9)
× (1.3–)2(–2.3) μm, pale brown, thick- and smooth-walled,
producing conidia sympodially on long open dencles; dencles
cylindrical, pale brown, up to 1 μm long. Conidia abundant on
OA and PCA, scarce on PDA, two-celled, broadly ellipsoidal with
a protuberant hilum, constricted at the septum, (5–)5.8(–7) ×
(2.2–)2.6(–3.1) μm, brown, verrucose, thick-walled, somemes
with a wing-like gelanous brown sheath, released by rhexolyc
secession. Sexual morph not observed.
Cultural characteriscs: Colonies at 25 °C aer 14 d: on OA
and PCA, at, woolly at centre, glabrous at periphery, top and
reverse sepia. On PDA raised, felty, top and reverse olivaceous
with ochreous diusible pigment.
Typus: Thailand, Nakhon Nayok Province, Mueang Nakhon Nayok
district, Wang Takrai waterfall, N14.330023° E101.307168°, 64
m above sea level, from soil, 22 Jul. 2008, P.W. Crous (holotype
CBS H-24237, culture ex-type CBS 145768).
Notes: The genus Verruconis (Sympoventuriaceae, Venturiales,
Dothideomycetes) was established to accommodate
thermophilic species segregated from Ochroconis ( O. gallopava
and O. calidiuminalis) and Scolecobasidium ( S. verruculosum),
which produce septate conidia from sympodially proliferang
conidiophores, released by rhexolyc secession (Samerpitak et
al. 2014). These species have been isolated from hot spring
water, warm euents or as soil saprobes (Yarita et al. 2007,
Samerpitak et al. 2014, Giraldo et al. 2014b). However, the type
species, V. gallopava has been reported as an opportunisc
pathogen of humans and causing infecons in other warm-
blooded animals, mainly birds (Revankar & Suon 2010, de
Hoog et al. 2011).
Verruconis thailandica is phylogenecally related to V.
verruculosa (Fig. 1), but the two species can be disnguished by
the length of their conidiophores (up to 9 μm long in V. thailandica
vs. up to 45 μm long in V. verruculosa) and conidia (up to 7 μm
long in V. thailandica vs. up to 9 μm long in V. verruculosa;
Samerpitak et al. 2014). Recently, three new species were
added to the genus, namely V. panacis from Panax notoginseng
(Zhang et al. 2018), V. hainanensis and V. pseudotricladiata from
submerged decaying leaves (Qiao et al. 2019). These species
clustered in a separate clade phylogenecally distant from V.
thailandica. They can be disnguished by the presence of four-
celled conidia in V. panacis and V. hainanensis, and branched
Y-shaped conidia in V. pseudotricladiata (Zhang et al. 2018, Qiao
et al. 2019).
Authors: A. Giraldo López and P.W. Crous
Fig. 11. Verruconis thailandica sp. nov. (CBS 145768). A, B. Colonies on PDA and OA, respecvely, at 25 °C aer 14 d. C–E. Denculated conidiogenous
cells. F. Conidia with gelanous brown sheath (arrow). Scale bars = 10 μm.
© 2020 Westerdijk Fungal Biodiversity Instute
Hernández-Restrepo et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
22
ACKNOWLEDGEMENTS
The authors thank the technical sta, A. van Iperen and T. Merkx for
their valuable assistance with cultures.
Conict of interest: The authors declare that there is no conict
of interest.
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