Accepted Manuscript
Genera of phytopathogaenic fungi: GOPHY 3
Y. Marin-Felix, M. Hernández-Restrepo, I. Iturrieta-González, D. García, J. Gené,
J.Z. Groenewald, L. Cai, Q. Chen, W. Quaedvlieg, R.K. Schumacher, P.W.J. Taylor,
C. Ambers, G. Bonthond, J. Edwards, S.A. Krueger-Hadfield, J.J. Luangsa-ard, L.
Morton, A. Moslemi, M. Sandoval-Denis, Y.P. Tan, R. Thangavel, N. Vaghefi, R.
Cheewangkoon, P.W. Crous
PII:
S0166-0616(19)30008-9
DOI:
https://doi.org/10.1016/j.simyco.2019.05.001
Reference:
SIMYCO 89
To appear in:
Studies in Mycology
Please cite this article as: Marin-Felix Y, Hernández-Restrepo M, Iturrieta-González I, García D, Gené
J, Groenewald JZ, Cai L, Chen Q, Quaedvlieg W, Schumacher RK, Taylor PWJ, Ambers C, Bonthond
G, Edwards J, Krueger-Hadfield SA, Luangsa-ard JJ, Morton L, Moslemi A, Sandoval-Denis M, Tan YP,
Thangavel R, Vaghefi N, Cheewangkoon R, Crous PW, Genera of phytopathogaenic fungi: GOPHY 3,
Studies in Mycology, https://doi.org/10.1016/j.simyco.2019.05.001.
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ACCEPTED MANUSCRIPT
Genera of phytopathogaenic fungi: GOPHY 3
Y. Marin-Felix1,2*, M. Hernández-Restrepo1, I. Iturrieta-González2, D. García2, J. Gené2, J.Z.
Groenewald1, L. Cai3, Q. Chen3, W. Quaedvlieg4, R.K. Schumacher5, P.W.J. Taylor6, C. Ambers7, G.
Bonthond1,8, J. Edwards9,10, S.A. Krueger-Hadfield11, J.J. Luangsa-ard12, L. Morton13, A. Moslemi6, M.
Sandoval-Denis1,14, Y.P. Tan15, 16, R. Thangavel17, N. Vaghefi18, R. Cheewangkoon19, and P.W.
Crous1,20,21*
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Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; 2Mycology Unit, Medical School
and IISPV, Universitat Rovira i Virgili, Sant Llorenç 21, 43201 Reus, Spain; 3State Key Laboratory of Mycology, Institute of
Microbiology, Chinese Academy of Sciences, Beijing 100101, China; 4Royal Van Zanten, P.O. Box 265, 1430 AG, Aalsmeer, The
Netherlands; 5Hölderlinstraße 25, 15517 Fürstenwalde/Spree, Germany; 6Faculty of Veterinary and Agricultural Sciences,
University of Melbourne, Melbourne, VIC 3010, Australia; 7P.O. Box 631, Middleburg, VA 20118, USA; 8Benthic Ecology,
GEOMAR Helmholtz Centre for Ocean Research Kiel, Hohenbergstraße 2, 24105 Kiel, Germany; 9Agriculture Victoria Research,
Department of Jobs, Precincts and Regions, AgriBio Centre, Bundoora, Victoria 3083, Australia; 10School of Applied Systems
Biology, La Trobe University, Bundoora, Victoria 3083, Australia; 11Department of Biology, University of Alabama at Birmingham,
1300 University Blvd, CH464, Birmingham, AL, USA, 35294; 12Plant Microbe Interaction Research Team, Integrative Crop
Biotechnology and Management Research Group, Bioscience and Biotechnology for Agriculture, NSTDA 113, Thailand Science
Park Phahonyothin Rd., Khlong Nueng, Khlong Luang, Pathum Thani 12120, Thailand; 13P.O. Box 5607, Charlottesville, VA
22905, USA; 14Faculty of Natural and Agricultural Sciences, Department of Plant Sciences, University of the Free State, P.O. Box
339, Bloemfontein 9300, South Africa; 15Department of Agriculture and Fisheries, Biosecurity Queensland, Ecosciences Precinct,
Dutton Park 4012, QLD, Australia; 16Microbiology, Department of Biology, Utrecht University, Utrecht, Netherlands. 17Plant
Health and Environment Laboratory, Ministry for Primary Industries, P.O. Box 2095, Auckland 1140, New Zealand; 18Centre for
Crop Health, University of Southern Queensland, Queensland 4350, Australia; 19Department of Entomology and Plant Pathology,
Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200, Thailand; 20Department of Biochemistry, Genetics &
Microbiology, Forestry & Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, South Africa 21Wageningen
University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The
Netherlands.
*Correspondence: Y. Marin-Felix, y.marin@wi.knaw.nl; P.W. Crous, p.crous@wi.knaw.nl
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Abstract: This paper represents the third contribution in the Genera of Phytopathogenic Fungi (GOPHY) series. The series provides
morphological descriptions, information about the pathology, distribution, hosts and disease symptoms for the treated genera, as
well as primary and secondary DNA barcodes for the currently accepted species included in these. This third paper in the GOPHY
series treats 21 genera of phytopathogenic fungi and their relatives including: Allophoma, Alternaria, Brunneosphaerella, Elsinoe,
Exserohilum, Neosetophoma, Neostagonospora, Nothophoma, Parastagonospora, Phaeosphaeriopsis, Pleiocarpon, Pyrenophora,
Ramichloridium, Seifertia, Seiridium, Septoriella, Setophoma, Stagonosporiopsis, Stemphylium, Tubakia and Zasmidium. This study
includes three new genera, 42 new species, 23 new combinations, four new names, and three typifications of older names.
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Key words: DNA barcodes, Fungal systematics, new taxa.
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Taxonomic novelties: New genera: Arezzomyces Y. Marín & Crous, Globoramichloridium Y. Marín & Crous, Wingfieldomyces Y.
Marín & Crous; New species: Allophoma pterospermicola Q. Chen & L. Cai, Alternaria aconidiophora Iturrieta-González, Dania
García & Gené, Alternaria altcampina Iturrieta-González, Dania García & Gené, Alternaria chlamydosporifera Iturrieta-González,
Dania García & Gené, Alternaria curvata Iturrieta-González, Dania García & Gené, Alternaria fimeti Iturrieta-González, Dania
García & Gené, Alternaria inflata Iturrieta-González, Dania García & Gené, Alternaria lawrencei Iturrieta-González, Dania García
& Gené, Alternaria montsantina Iturrieta-González, Dania García & Gené, Alternaria pobletensis Iturrieta-González, Dania García
& Gené, Alternaria pseudoventricosa Iturrieta-González, Dania García & Gené, Brunneosphaerella roupeliae Crous, Elsinoe
picconiae Crous, Elsinoe veronicae Crous, Thangavel & Y. Marín, Neosetophoma aseptata Crous, R.K. Schumach. & Y. Marín,
Neosetophoma phragmitis Crous, R.K. Schumach. & Y. Marín, Neosetophoma sambuci Crous, R.K. Schumach. & Y. Marín,
Neostagonospora sorghi Crous & Y. Marín, Parastagonospora novozelandica Crous, Thangavel & Y. Marín, Parastagonospora
phragmitis Crous & Y. Marín, Phaeosphaeriopsis aloes Crous & Y. Marín, Phaeosphaeriopsis aloicola Crous & Y. Marín,
Phaeosphaeriopsis grevilleae Crous & Y. Marín, Phaeosphaeriopsis pseudoagavacearum Crous & Y. Marín, Pleiocarpon
livistonae Crous & Quaedvl., Pyrenophora avenicola Y. Marín & Crous, Pyrenophora cynosuri Y. Marín & Crous, Pyrenophora
novozelandica Y. Marín & Crous, Pyrenophora pseudoerythrospila Y. Marín & Crous, Pyrenophora sieglingiae Y. Marín & Crous,
Pyrenophora variabilis Hern.-Restr. & Y. Marín, Septoriella germanica Crous, R.K. Schumach. & Y. Marín, Septoriella hibernica
Crous, Quaedvl. & Y. Marín, Septoriella hollandica Crous, Quaedvl. & Y. Marín, Septoriella pseudophragmitis Crous, Quaedvl. &
Y. Marín, Setophoma brachypodii Crous, R.K. Schumach. & Y. Marín, Setophoma pseudosacchari Crous & Y. Marín, Stemphylium
rombundicum Moslemi, Y.P. Tan & P.W.J. Taylor, Stemphylium truncatulae Moslemi, Y.P. Tan & P.W.J. Taylor, Stemphylium
waikerieanum Moslemi, Jacq. Edwards & P.W.J Taylor, Vagicola arundinis Phukhams., Camporesi & K.D. Hyde, Zasmidium
thailandicum Crous; New combinations: Arezzomyces cytisi (Wanas. et al.) Y. Marín & Crous, Globoramichloridium indicum
(Subram.) Y. Marín & Crous, Phaeosphaeria phoenicicola (Crous & Thangavel) Y. Marín & Crous, Pyrenophora poae (Baudyš) Y.
Marín & Crous, Pyrenophora wirreganensis (Wallwork et al.) Y. Marín & Crous, Seiridium cupressi (Nattrass et al.) Bonthond,
Sandoval-Denis & Crous, Seiridium pezizoides (de Not.) Crous, Septoriella agrostina (Mapook et al.) Y. Marín & Crous,
Septoriella artemisiae (Wanas. et al.) Y. Marín & Crous, Septoriella arundinicola (Wanas. et al.) Y. Marín & Crous, Septoriella
arundinis (W.J. Li et al.) Y. Marín & Crous, Septoriella bromi (Wijayaw. et al.) Y. Marín & Crous, Septoriella dactylidis (Wanas. et
al.) Y. Marín & Crous, Septoriella elongata (Wehm.) Y. Marín & Crous, Septoriella forlicesenica (Thambug. et al.) Y. Marín &
Crous, Septoriella garethjonesii (Thambug. et al.) Y. Marín & Crous, Septoriella italica (Thambug. et al.) Y. Marín & Crous,
Septoriella muriformis (Ariyaw. et al.) Y. Marín & Crous, Septoriella rosae (Mapook et al.) Y. Marín & Crous, Septoriella
subcylindrospora (W.J. Li et al.) Y. Marín & Crous, Septoriella vagans (Niessl) Y. Marín & Crous, Wingfieldomyces cyperi (Crous
& M.J. Wingf.) Y. Marín & Crous, Zasmidium ducassei (R.G. Shivas et al.) Y. Marín & Crous; New names: Pyrenophora nisikadoi
Y. Marín & Crous, Septoriella dactylidicola Y. Marín & Crous, Septoriella neoarundinis Y. Marín & Crous, Septoriella
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neodactylidis Y. Marín & Crous; Typification: Epitypification: Ascochyta chrysanthemi F. Stevens, Pestalotia unicornis Cooke &
Ellis, Rhynchosphaeria cupressi Nattrass et al.
INTRODUCTION
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Genera of Phytopathogenic Fungi (GOPHY) is a series of papers with the main focus to provide a stable
platform for the taxonomy of phytopathogenic fungi. All genera included here are associated with plant
disease, but note that many species treated are not well-known plant pathogens, or Koch’s postulates
remain to be completed for them. This series links to a larger initiative known as the “The Genera of
Fungi project” (www.GeneraOfFungi.org, Crous et al. 2014a, 2015a, Giraldo et al. 2017), which aims to
revise the generic names of all currently accepted fungi (Kirk et al. 2013). Specific aims were detailed by
Marin-Felix et al. (2017), when this series was launched. One of the most important aims is to resolve
generic and species concepts of the fungi studied, since many taxa have been shown to represent species
complexes, or to comprise poly- or paraphyletic genera (Crous et al. 2015b). Other issues to resolve
include the fact that type material for many genera and species has not been designated or is missing, and
that the vast majority of these taxa were described before the DNA era (Hibbett et al. 2011) and thus lack
DNA barcodes (Schoch et al. 2012). Therefore, another important aim is to generate DNA barcodes of
type species and type specimens in order to fix the application of these names. Moreover, in cases where
no type material has been preserved, taxa need to be recollected, epi- or neotypes designated, and
registered in MycoBank to ensure traceability of the nomenclatural act (Robert et al. 2013). Finally, it is
necessary to designate a single scientific name for fungi (Crous et al. 2015b) for which sexual-asexual
links have been resolved.
Two issues of GOPHY have already been published, in which 41 genera were treated, including a
total of two new genera, 46 new species, 15 new combinations and 10 typifications of older names
(Marin-Felix et al. 2017, 2019). In this third contribution, a further 21 genera are treated, resulting in the
clarification of their taxonomy and classification, and the introduction of three new genera, 42 new
species, 23 new combinations, four new names and the typification of three older names.
For submissions to future issues in the GOPHY series, mycologists are encouraged to contact Pedro
Crous (p.crous@wi.knaw.nl) to ensure there is no overlap with activities arising from other research
groups. Preference will be given to genera that include novel species, combinations or typifications.
Generic contributions published in each issue will also be placed in the database displayed on
www.plantpathogen.org.
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MATERIAL AND METHODS
Isolates and morphological analysis
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Descriptions of the new taxa and typifications are based on cultures obtained from the collection at the
American Type Culture Collection, Manassas, Virginia, USA (ATCC), the Queensland Plant Pathology
Herbarium, Brisbane, Australia (BRIP), the Westerdijk Fungal Biodiversity Institute, Utrecht, The
Netherlands (CBS), the working collection of P.W. Crous (CPC), housed at the Westerdijk Fungal
Biodiversity Institute (WI), the Chinese General Microbiological Culture Collection Center, Beijing,
China (CGMCC), the Facultat de Medicina i Ciències de la Salut, University Rovira i Virgili, Reus, Spain
(FMR), and the Victorian Plant Pathogen Herbarium, Bundoora, Australia (VPRI). For fresh collections,
we followed the procedures previously described in Waksman et al. (1922), Crous et al. (1991) and
Calduch et al. (2004). Colonies were transferred to different media, i.e. cornmeal agar (CMA), 2 % malt
extract agar (MEA), potato carrot agar (PCA), 2 % potato dextrose agar (PDA), synthetic nutrient-poor
agar (SNA), oatmeal agar (OA), water agar (WA) (Crous et al. 2019b), pine needle agar (PNA; Smith et
al. 1996), and incubated under different conditions to induce sporulation. Requirements of media and
conditions of incubation are specified for each genus. Reference strains and specimens are maintained at
ATCC, BRIP, CBS, CPC, CGMCC, FMR and VPRI.
Vegetative and reproductive structures were mounted in 100 % lactic acid or Shear’s solution either
directly from specimens or from colonies sporulating on CMA, MEA, OA, PCA, PDA, PNA, SNA or
WA. For cultural characterisation, isolates were grown and incubated on different culture media and
temperatures as indicated for each genus. Colour notations were rated according to the colour charts of
Kornerup & Wanscher (1978) for Alternaria, and Rayner (1970) for all other genera. Taxonomic
novelties were deposited in MycoBank (www.MycoBank.org; Crous et al. 2004).
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DNA isolation, amplification and analyses
RESULTS
Allophoma Q. Chen & L. Cai, Stud. Mycol. 82: 162. 2015. Fig. 1.
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Fungal DNA was extracted and purified directly from the colonies or host material as specified for each
genus. Primers and protocols for the amplification and sequencing of gene loci, and software used for
phylogenetic analyses can be found in the bibliographies provided for each genus. Phylogenetic analyses
consisted of Maximum-Likelihood (ML) and Bayesian Inference (BI). ML was inferred as described in
Hernández-Restrepo et al. (2016b), or by using MEGA v. 6.0 (Tamura et al. 2013). BI was carried as
described by Hernández-Restrepo et al. (2016b), or by using MrBayes on XSEDE v. 3.2.6 on the CIPRES
portal (www.phylo.org). Sequence data generated in this study were deposited in GenBank and the
alignments and trees in TreeBASE (http://www.treebase.org).
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Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Didymellaceae.
Type species: Allophoma tropica (R. Schneid. & Boerema) Q. Chen & L. Cai, basionym: Phoma tropica
R. Schneid. & Boerema, Phytopathol. Z. 83: 361. 1975. Isotype and ex-isotype strain: CBS H-7629,
CBS 436.75 = DSM 63365.
DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): rpb2, tub2. Table 1. Fig. 2.
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Conidiomata pycnidial, globose to flask-shaped, ovoid, superficial or (semi-)immersed, solitary or
confluent, ostiolate, sometimes with an elongated neck; conidiomatal wall pseudoparenchymatous,
multi-layered. Conidiogenous cells phialidic, hyaline, smooth-walled, ampulliform to doliiform,
sometimes flask-shaped or isodiametric. Conidia hyaline, smooth- and thin-walled, aseptate or 1septate, variable in shape and size, i.e. ovoid, oblong, ellipsoidal to cylindrical, or slightly allantoid,
mostly guttulate. Chlamydospores uni- or multicellular (pseudosclerotioid and dictyosporous), solitary
or in chains, intercalary or terminal, smooth-walled, brown, where multicellular variable in shape and
size. Swollen cells (pseudo-chlamydospores) pale brown, terminal or intercalary, solitary or in
clusters, variable in size and shape, commonly in aerial mycelia. Sexual morph unknown (adapted
from Chen et al. 2015, Babaahmadi et al. 2018).
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Culture characteristics: Colonies on OA white when young, grey to olivaceous or dull green, brown,
floccose to woolly, sometimes with rosy-buff tinges near the colony margins or yellow pigment in the
sterile sectors, margins regular.
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Optimal media and cultivation conditions: OA or sterile pine needles placed on OA under nearultraviolet light (12 h light, 12 h dark) to promote sporulation at 25 °C.
Distribution: Worldwide.
Hosts: Wide host range, occurring as pathogens or saprobes, on Araceae, Fabaceae, Gesneriaceae,
Myrtaceae, Papaveraceae, Piperaceae, Primulaceae, Rosaceae, Rubiaceae, Solanaceae, Sterculiaceae,
Verbenaceae and other hosts, including humans.
Disease symptoms: Dieback, tissue necrosis, leaf spots, stem rot, leaf blotch, but also saprobic or
isolated from other substrates and environments, e.g. air from karst caves and human infections.
Notes: The genus Allophoma was introduced by Chen et al. (2015) to accommodate five previously
described Phoma species, namely Al. labilis (syn. Pho. labilis), Al. minor (syn. Pho. minor), Al.
piperis (syn. Pho. piperis), Al. tropica (syn. Pho. tropica) and Al. zantedeschiae (syn. Pho.
zantedeschiae), and a new species Al. nicaraguensis. Another four species have been described in the
subsequent years, i.e. Al. cylindrispora (Valenzuela-Lopez et al. 2018), Al. hayatii (Babaahmadi et al.
2018), Al. oligotrophica (Chen et al. 2017) and Al. pterospermicola sp. nov. in the present study.
Differentiating Allophoma from related phoma-like genera based on morphology alone is sometimes
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complicated. Furthermore, Allophoma species are morphologically similar and hard to differentiate
from one another. Therefore, molecular data are essential for accurate identification of species within
this genus, with ITS, LSU, tub2 and rpb2 being the loci selected for this purpose (Chen et al. 2015,
2017, Valenzuela-Lopez et al. 2018). No sexual morph of this genus has been observed to date.
These fungi are generally found in soil, air and regarded as saprobes or as the causal organisms of
various diseases of different herbaceous and woody plants, such as some ornamental plants, coffee,
etc., and even human eye lesions (Boerema et al. 2004, Aveskamp et al. 2010, Chen et al. 2015, 2017,
Babaahmadi et al. 2018, Valenzuela-Lopez et al. 2018).
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References: Boerema et al. 2004 (morphology and pathogenicity), Aveskamp et al. 2010, Chen et al.
2015, 2017, Babaahmadi et al. 2018, Valenzuela-Lopez et al. 2018 (morphology, phylogeny and
pathogenicity).
Allophoma pterospermicola Q. Chen & L. Cai, sp. nov. MycoBank MB828313. Fig. 3.
Etymology: Name reflects Pterospermum, the host genus from which it was collected.
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Conidiomata pycnidial, solitary or aggregated, globose to subglobose, brown, glabrous or with a few
hyphal outgrowths, superficial, 60–330 × 67–280 µm, with 1–5 ostioles, sometimes elongated as a
long neck, up to 150 m long, papillate; conidiomatal wall pseudoparenchymatous, 3–5-layered, 12–
20 µm thick, composed of isodiametric cells. Conidiogenous cells phialidic, hyaline, smooth-walled,
ampulliform to doliiform, 6–10 × 3–6 µm. Conidia oval to oblong, occasionally bacilliform, smoothand thin-walled, hyaline, aseptate, 3–5.5 × 1.5–2 µm, with 1–2 minute guttules. Conidial matrix
cream.
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Culture characteristics: Colonies on OA, 33–40 mm diam after 1 wk, margins regular, floccose to
woolly, white, pale brownish grey, with a pale salmon concentric ring, pale salmon near the margins,
black pycnidia visible; reverse concolourous. Colonies on MEA, 20–25 mm diam after 1 wk, margins
regular, aerial mycelium sparse, olivaceous; reverse concolourous. Colonies on PDA, 20–30 mm diam
after 1 wk, margins regular, floccose to woolly, olivaceous, white near the margins; reverse dull green,
white near the margins. Application of NaOH results in a pale brownish olivaceous discolouration of
the agar.
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Typus: China, Guangxi, Nonggang National Nature Reserve, on diseased leaves of Pterospermum
xylocarpum (Sterculiaceae), Jun. 2017, Z.Y. Ma (holotype HMAS 247983, dried culture, ex-holotype
living culture CGMCC 3.19245 = LC 12185).
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Additional materials examined: China, Guangxi, Nonggang National Nature Reserve, on diseased leaves of Pterospermum
xylocarpum (Sterculiaceae), Jun. 2017, Z.Y. Ma, LC 12183; ibid. LC 12184; Guangxi, Jingxi, Gulongshan, on diseased leaves
of Maesa montana (Primulaceae), Jun. 2017, Z.Y. Ma, LC 12181; ibid. LC 12182.
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Notes: Allophoma pterospermicola represents the first report of a species in the family Didymellaceae
on the two host genera Pterospermum (Sterculiaceae) and Maesa (Primulaceae). This species is
closely related to Al. minor, but differs in producing longer conidiogenous cells [6–10 × 3–6 µm in Al.
pterospermicola vs. 4–5.5(–6.2) × 3–4.5(–4.7) in Al. minor] and slightly narrower conidia [3–5.5 ×
1.5–2 µm in Al. pterospermicola vs. (3–)3.5–4.5(–5) × 1.8–2.5(–3) m in Al. minor]. In addition, Al.
pterospermicola grows much slower on OA, MEA and PDA than Al. minor, and the latter species has
only been recorded on Syzygium aromaticum (Myrtaceae) (Aveskamp et al. 2010).
Authors: Q. Chen & L. Cai
Alternaria Nees, Das System der Pilze und Schwämme: 72. 1816 (1816–1817). Fig. 4.
For synonyms see Woudenberg et al. (2013).
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae.
Type species: Alternaria alternata (Fries) Keissler, basionym: Torula alternata Fr., Syst. Mycol.
(Lundae) 3: 500. 1832 (nom. sanct.); additional synonyms listed in Woudenberg et al. (2015). Neotype
designated by Simmons (1967): E.G.S. 11.050. Ex-epitype strain designated by de Hoog & Horré (2002):
CBS 916.96 = E.G.S. 34.016.
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DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, ATPase, gapdh, rpb2 and tef1. Table 2. Figs 5–7.
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Ascomata small, solitary to clustered, erumpent to almost superficial at maturity, dark brown, globose to
ovoid, apically papillate, ostiolate, smooth or setose at maturity, with a thin ascomatal wall; centrum
formed by a hamathecium with cellular pseudoparaphyses and asci in basal layer. Asci bitunicate,
fissitunicate, uni- or biseriate, (4–6–)8-spored, cylindrical to cylindro-clavate, straight or somewhat
curved, with a short furcate pedicel. Ascospores ellipsoid to fusoid, muriform, slightly constricted at
septa, 3–7-transverse septa, 1–2 series of longitudinal septa through the two original central segments,
end cells without septa, or with one longitudinal or oblique septum, or with a Y-shaped pair of septa,
yellow-brown, smooth-walled, without guttules. Conidiophores macronematous or semi-macronematous,
mononematous, simple or branched, pale brown or brown. Conidiogenous cells integrated, terminal
becoming intercalary, mono- or polytretic and sympodial, cicatrized. Conidia solitary or in simple or
branched chains, dry, ovoid, obovoid, cylindrical, narrowly ellipsoid or obclavate, beaked or non-beaked,
pale or medium olivaceous brown to brown, smooth-walled or verrucose, with transverse and with or
without oblique or longitudinal septa; septa can be thick, dark, an internal cell-like structure can be
formed. Species with meristematic growth are known (adapted from Ellis 1976, Woudenberg et al. 2013,
2014, Grum-Grzhimaylo et al. 2016).
Culture characteristics: Colonies effuse, grey, olivaceous brown, dark blackish brown or black;
mycelium immersed or partly superficial, composed of colourless, olivaceous brown or brown hyphae.
Optimal media and cultivation conditions: For morphological examinations the use of PCA and V-8 is
recommended, incubated at moderate temperatures (ca. 22–25 ºC) under near-ultraviolet light (8 h light,
16 h dark), without humidity control, for 5–7 d or more if necessary (Simmons 2007). We also
recommend microscopic examination of OA cultures due to the alterations observed on the conidial wall
when grown on PCA.
Distribution: Worldwide.
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Hosts: Mainly pathogens of a wide range of plant families, such as Apiaceae, Asteraceae, Brassicaceae,
Cyperaceae, Poaceae, Rosaceae, Rutaceae, Solanaceae, among others (Thomma 2003, Lawrence et al.
2016). Some are implicated as human pathogens (de Hoog et al. 2011).
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Disease symptoms: Most species are foliar pathogens, causing necrotic lesions as brown/black spots or
“target spot” with the fungus residing in the central area, but also inducing leaf blight; seed-borne species
may attack seedlings, resulting in damping-off, stem lesions or collar rot; sunken and dark lesions are
present in roots, tubers, stems and fruits infections; some rots and decay are typical symptoms of postharvest diseases (Laemmlen 2001, Thomma 2003, Lawrence et al. 2008). Phytotoxins are also produced
during the invasion process as virulence factors which affect a wide spectrum of plant species. Alternaria
toxins diffuse into host tissues resulting in a chlorotic or yellow halo around lesions, exacerbating the
severity of the symptoms (Singh et al. 2015).
Notes: Alternaria is characterised mainly by its asexual morph with darkly pigmented multi-celled
conidia, which are typically dictyosporous, some phragmosporous, and arranged single or in chains on the
conidiophore. Some of these morphological features can also be observed in other closely related genera
such as Paradendryphiella (Woudenberg et al. 2013) or Stemphylium (Woudenberg et al. 2017).
However, Paradendryphiella mainly differs by its denticulate conidiogenous cells with prominent
conidial scars aggregated at the apex of simple or branched conidiophores, and Stemphylium by showing
percurrent conidiophores with apically swollen conidiogenous cells.
Extensive morphological investigations of the genus Alternaria were carried out by Emory G.
Simmons, which culminated with his monograph on Alternaria species identification (Simmons 2007).
Based on the sporulation patterns and conidial morphology, he described several Alternaria speciesgroups which were typified by representative species (Simmons 1992). In recent years, based on
molecular phylogenetic approaches using DNA sequence data, it has been shown that the main
morphological groups identified by Simmons represent monophyletic species groups. Lawrence et al.
(2013) provided the first strongly supported phylogenetic hypothesis among Alternaria lineages and
elevated several of those monophyletic species groups to the taxonomic status of sections, each with a
type species. Successive phylogenetic investigations added additional sections within the genus by
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synonymising genera such as Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, Embellisia,
Nimbya, Pseudoalternaria, Sinomyces, Teretispora, Ulocladium, Undiphilum and Ybotromyces
(Woudenberg et al. 2013, 2014, Lawrence et al. 2016). Therefore, the genus Alternaria currently
comprises close to 280 species, most of them classified in 27 sections. Taxonomic traits and species
composition of all Alternaria sections are summarised in Lawrence et al. (2016).
Considering, however, the overlap of morphological traits among Alternaria sections/species and that
the culture conditions can greatly influence the morphology of these fungi, molecular identification is
practically mandatory for the classification of Alternaria isolates. Although the ITS barcode is considered
a good phylogenetic marker to define sections, it has limited discriminatory power to distinguish species,
making multi-locus sequence analysis with several protein-coding loci essential for accurate species
identification. While Woudenberg et al. (2013), in addition to the nrDNA regions, used the combination
of gapdh, rpb2 and tef1 loci for redefining the genus, the combination of other phylogenetic markers have
since been analysed to determine relationships and species delineation in studies on a particular section;
i.e. ITS, Alt a-1, endoPG, gapdh, OPA10-2, rpb2 and tef1 for section Alternaria (Woudenberg et al.
2015); ITS, ATPase, tef1 and gapdh for sections Infectoriae and Pseudoalternaria (Andersen et al. 2009,
Deng et al. 2018, Poursafar et al. 2018); ITS, Alt a-1, gapdh, rpb2 and tef1 for section Porri
(Woudenberg et al. 2014); and ITS, Alt a-1 and gapdh for section Sonchi (Lawrence et al. 2012, Deng et
al. 2014). Nevertheless, according to Lawrence et al. (2013) the plasma membrane ATPase, cmdA, and
Alt a-1 loci are the most informative markers for Alternaria species delimitation. However, considering
that the latter locus unreliably amplifies some species within sect. Infectoriae, they suggested that the
most suitable genetic markers for molecular identification at the species level are ATPase and cmdA genes
(Lawrence et al. 2013, 2016). Unfortunately, the latter marker has not been used for the phylogeny of any
of the above-mentioned sections.
Alternaria is a very successful pathogenic genus that causes disease on a great number of
economically important plants, causing large economic losses due to the number of plant species affected
and worldwide distributions of several Alternaria species (Meena et al. 2017). They are commonly
described causing stem canker, leaf blight or leaf spot on a large variety of crops, including cereals,
ornamentals, oil crops, vegetables such as broccoli, cauliflower, carrot, onion and potato, and fruits like
apple, citrus, pear and strawberry, among others. Species in section Alternaria, such as A. alternata, A.
arborescens or A. tenuissima, as well as others from sections Alternantherae, Brassiccicola, Crivellia,
Gypsophilae, Nimbya, Radicina or Sonchi, are frequently reported causing such diseases, but the largest
group of phytopathogens in the genus is concentrated in section Porri (Lawrence et al. 2016, Meena et al.
2017). The most relevant plant pathogens in this latter section are A. bataticola, A. porri, A. solani and A.
tomatophila (Woudenberg et al. 2014). Alternaria species also produce diverse phytotoxins, which affect
their host plants at different stages of pathogenesis (Thomma 2003, Lawrence et al. 2008, Meena et al.
2017). Some of these phytotoxins have been evaluated by the European Food Safety Authority as
potentially causing risks to human health (Meena et al. 2017).
In humans, Alternaria species are commonly associated with hypersensitivity pneumonitis, bronchial
asthma, allergic sinusitis and rhinitis. To a lesser extent, they have been also described as causing
paranasal sinusitis, ocular infections, onychomycosis, cutaneous and subcutaneous infections,
granulomatous pulmonary disease, soft palate perforation and disseminated disease (Pastor & Guarro
2008, de Hoog et al. 2011).
In several surveys of microfungi from Spanish regions with different climates and biodiversity,
samples of plant litter (leaves, bark and twigs) and dung of wild and farm herbivore animals (rabbits,
rodents, goats, cattle and horses) were collected. From these samples, we found 16 interesting Alternaria
isolates, belonging to sections Infectoriae, Pseudoalternaria, Chalastospora and Radicina. The multilocus phylogenetic analysis based on five above-mentioned gene markers showed that 10 of them were
undescribed species for the genus, and the others were identified as A. kourtkuyana, A. rosae and A.
malorum (Figs 5–7). Most of these novel species have been isolated from herbivore dung, which appear
to represent a reservoir of interesting Alternaria species which could represent potential plant pathogens.
References: Ellis 1976, Simmons 2007 (morphology); Laemmlen 2001, Thomma 2003, Lawrence et al.
2008, Meena et al. 2017 (plant infections); Pastor & Guarro 2008, de Hoog et al. 2011 (human
infections); Woudenberg et al. 2013, 2014, 2015, Grum-Grzhimaylo et al. 2016, Lawrence et al. 2016,
Poursafar et al. 2018 (morphology and phylogeny).
Alternaria aconidiophora Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829626.
Fig. 8.
Etymology: Name refers to the lack of conidiophores from vegetative hyphae.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 1–4 m wide, septate, branched,
hyaline to greyish, smooth-walled. Conidiophores absent. Conidiogenous loci inconspicuous on
vegetative hyphae, scarce. Conidia commonly solitary at centre of the colony, globose, ovoid, near
ellipsoid or obclavate, 12–31 × 7–12 m, with some darkened middle transverse septa, 1–5 transverse, 0–
1(–2) longitudinal or oblique septa per transverse segment, brown, smooth-walled. Secondary
conidiophores present, may be formed apically from the conidial body as a short extension often
geniculate, with one or two, terminal or subterminal conidiogenous loci. Sexual morph not observed.
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Culture characteristics: Colonies on PDA reaching 64 mm diam after 1 wk at 25 °C, flat, cottony at
centre, slightly radially folded towards the periphery, aerial mycelium abundant, margins regular; surface
white (1A1); reverse yellowish white (4A2). On PCA attaining 54 mm diam, flat, aerial mycelium scarce,
margins regular; surface greyish green to greenish grey (1D3/1B1); reverse greenish grey (1C2/1B1). On
OA reaching 61 mm diam, flat, aerial mycelium scarce, margins regular; surface and reverse colourless.
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
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Typus: Spain, Catalonia, Alta Ribagorça, Vall de Boí, isolated from forest leaf litter, Dec. 2017, J. Gené
(holotype CBS H-23891, culture ex-type CBS 145419 = FMR 17111).
Notes: Alternaria aconidiophora together with A. fimeti, both species introduced here from herbivore
dung, are placed in an unsupported clade in Alternaria section Infectoriae (Fig. 5). Morphologically, the
latter differs from A. aconidiophora in having conspicuous sporulation with well-differentiated
conidiophores and verrucose conidia up to 44 m long. The conidia of A. aconidiophora are smoothwalled and 12–31 m long.
Alternaria altcampina Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829627. Fig. 9.
Etymology: Name refers to the region of Alt Camp (Catalonia) from where the fungus was collected.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 2–4 m wide, branched, pale
yellowish brown to brown, septate, smooth-walled to verruculose. Conidiophores macronematous, arising
laterally or terminally from aerial hyphae, erect to slightly flexuous, unbranched, occasionally branched,
up to 10-septate, 12–88 × 3–4 m, brown becoming pale towards apex, smooth-walled, with 1 terminal
and up to 3 subterminal conidiogenous loci. Conidia in branched chains, occasionally solitary, ovoid,
obclaviform, ellipsoidal or somewhat cylindrical, 9–43 × 6–8 m, with darkened middle transverse septa,
(1–)2–3(–6) transverse, 0–1 longitudinal or oblique septa in up to 4 of the transverse segments, usually
inconspicuous, pale yellowish to yellowish brown, verrucose. Secondary conidiophores commonly
formed apically as a beak from conidial body, or as a lateral conidiogenous loci from body cells bearing
conidia in short chains. Sexual morph not observed.
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Culture characteristics: Colonies on PDA reaching 73 mm diam after 1 wk at 25 °C, flat, densely
floccose, aerial mycelium abundant, margins fimbriate; surface olive brown to blond (4D3/4C4), white at
the periphery; reverse yellowish brown to orange-grey (5E4/5B2). On PCA attaining 66 mm diam, flat,
granular, aerial mycelium scarce, margins regular; surface dark green (30F8); reverse dull green (30E4).
On OA reaching 70 mm diam, flat, loosely floccose at centre, aerial mycelium scarce, margins regular;
surface dark green (28F4); reverse dull green (29E3).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
Typus: Spain, Catalonia, Alt Camp, isolated from goat dung, Mar. 2017, I. Iturrieta-González, M.
Guevara-Suarez & J. Guarro (holotype CBS H-23892, culture ex-type CBS 145420 = FMR 16476).
Notes: Based on the phylogeny of ITS, ATPase and gapdh, A. altcampina is classified in Alternaria
section Pseudoalternaria (Fig. 6). It is closely related to the recently described species A.
parvicaespitosa, which was isolated from harvested blueberry fruit (California, USA), and A. kordkuyana,
isolated from symptomatic wheat heads of Triticum aestivum (Kordkuy, Iran). Alternaria parvicaespitosa
differs in having smaller conidia (10–25 × 7–12 m) with smooth to slightly punctulate outer walls
(Gannibal & Lawrence 2016), and A. kordkuyana by its larger conidia [30–50(−60) × 7–11 m] and
shorter conidiophores (10–40 × 3–4 m) (Poursafar et al. 2018).
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Alternaria chlamydosporifera Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829628.
Fig. 10.
Etymology: Name refers to the production of abundant chlamydospores in culture.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 3–6 m wide, septate, branched,
pale brown to brown, smooth-walled. Conidiophores macronematous, arising directly from aerial hyphae,
erect to slightly flexuous, occasionally geniculate at apex, 1–4-septate, unbranched or scarcely branched,
14–140 × 3–5 m, dark brown, verruculose, with 1–2 conidiogenous loci. Conidia mostly solitary,
occasionally in short chains with up to two conidia, ellipsoidal or ovoid, occasionally subglobose, 12–41
× 7–20 m, with darkened middle transverse septa, 1–3(–4) transverse, and 0–1(–2) longitudinal septa per
transverse segments, brown to dark brown, verruculose. Secondary conidiophores can be formed apically
from conidial body as a beak, geniculate, with 1–3 terminal or lateral conidiogenous loci, bearing solitary
or short chains of conidia. Chlamydospores abundant, immersed, intercalary, irregular shape, rarely
broadly ellipsoidal or clavate, muriform, sometimes showing central constriction, 60–91 × 32–57 m,
dark brown to black. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching 82 mm diam after 1 wk at 25 °C, flat, densely
floccose, aerial mycelium abundant, margins regular; surface greyish brown (5E3); reverse black to
greyish brown (5E3). On PCA attaining 68 mm diam, flat, with granular appearance by the presence of
abundant chlamydospores, aerial mycelium scarce, margins regular; surface dark green (29F5); reverse
dark green (30F8). On OA reaching 71 mm diam, flat, loosely floccose at centre, slightly granular
towards the periphery, aerial mycelium scarce, margins slightly lobate; surface dark green (29F4); reverse
dark green (29F4).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
Typus: Spain, Huesca, Baells, isolated from rabbit dung, Apr. 2018, G. Sisó & D. García (holotype CBS
H-23893, culture ex-type CBS 145421 = FMR 17360).
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Notes: Alternaria chlamydosporifera belongs to Alternaria section Radicina (Fig. 7). It is included in a
well-supported clade (95 % BS) with A. petroselini, A. selini and A. smyrnii, which are pathogens of
Apiaceae (Lawrence et al. 2016). Alternaria petroselini and A. selini can be easily differentiated by the
lack of chlamydospores in culture and their larger (50–66 m in A. petroselini vs. 48–65 m in A. selini)
and usually ellipsoidal conidia (Simmons 1995). Although A. smyrnii, the closest relative to A.
chlamydosporifera, has been described as producing sclerotial knots in culture that are able to form fertile
conidiophores, its conidia are considerably longer (67–96 m) (Simmons 1995) than those of A.
chlamydosporifera (12–41 m long). In addition, we have never observed conidiophores associated with
the chlamydospores of the latter species.
Alternaria curvata Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829628. Fig. 11.
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Etymology: Name refers to the presence of curved conidia.
Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 2–6 m wide, septate, branched,
hyaline to yellowish brown to brown, smooth-walled to verruculose. Conidiophores macronematous,
arising laterally or terminally from aerial hyphae, erect to slightly flexuous, usually unbranched, up to 14septate, 23–80 × 3–5 m, brown to dark brown, smooth-walled, with a terminal or occasionally a subterminal conidiogenous loci. Conidia forming branched chains, with up to 5 conidia in unbranched part,
ovoid or nearly ellipsoidal, often slightly curved, 13–47(–70) × 4–16 m, with darkened middle
transverse septa, (0–)1–5(–7) transverse, and 0–2(–3) longitudinal or oblique septa per transverse
segment, brown to dark brown, verrucose to tuberculate. Secondary conidiophores can be formed apically
or laterally from the conidial body as a short extension bearing conidia in short chains. Sexual morph not
observed.
Culture characteristics: Colonies on PDA reaching 63 mm diam after 1 wk at 25 °C, flat, densely
floccose, aerial mycelium abundant, margins regular; surface white to dull green (1A1/30D4); reverse
dark green to olive yellow (30F8/2D6), white at the periphery. On PCA attaining 62 mm diam, flat,
loosely floccose, aerial mycelium scarce, margins regular; surface olive (3F4); reverse dark green to grey
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(29F4/29B1). On OA reaching 61 mm diam, scarce aerial mycelium towards the periphery, margins
regular; surface greyish green (30E5), with greyish mycelium tufts at centre; reverse dull green (29E4).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park, isolated from goat dung, Oct. 2017, G. Sisó
& C. González-García (holotype CBS H-23894, culture ex-type CBS 145422 = FMR 16901).
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Notes: Alternaria curvata was included in the section Infectoriae, forming an unsupported basal clade
together with A. montsantina and A. pseudoventricosa, both introduced here, and A. ventricosa (Fig. 5).
Morphologically, the former two species differ from A. curvata in lacking curved conidia. Alternaria
ventricosa has asymmetrical conidia, due to the hyperplasia and hypertrophy of cells, especially on one
side of the conidia (Roberts 2007). Other morphologically similar species are A. fimeti and A. triticina,
which also have curved conidia. However, A. triticina is phylogenetically more distant and its conidia are
strongly inequilateral and wider (up to 22 m) (Simmons 2007) than those of A. curvata (4–16 m wide).
Alternaria fimeti can be differentiated from A. curvata by the production of longer conidiophores (22–182
m in A. fimeti vs. 23–80 m in A. curvata) and the absence or scarce development of secondary
conidiophores.
Alternaria fimeti Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829630. Fig. 12.
Etymology: Name refers to the substrate where the species was isolated, herbivore dung.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 2–5 m wide, septate, branched,
hyaline to subhyaline to pale yellowish, verruculose. Conidiophores semi- to macronematous, arising
laterally or terminally from aerial hyphae, erect to slightly flexuous, unbranched (can be slightly branched
on OA), up to 9-septate, 22–182 × 1–5 m, pale brown, smooth-walled, with 1 terminal conidiogenous
locus. Conidia solitary or in short chains of up to six conidia, ovoid, obpyriform or obclavate, some
slightly curved, 9–44 × 5–14(–23) m, with darkened middle transverse septa, 0–5 transverse (up to 7 in
OA), and 0–1(–2) longitudinal or oblique septa per segment, brown, verrucose. Secondary conidiophores
only scarcely produced on OA as apical or lateral extension from conidial body, up to 25 m long. Sexual
morph not observed.
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Culture characteristics: Colonies on PDA reaching 66 mm diam after 1 wk at 25 °C, flat, densely
floccose, aerial mycelium abundant, margins fimbriate; surface yellowish grey to yellowish white
(3C2/3A2); reverse yellowish brown to light yellow (5E8/4A5). On PCA attaining 65 mm diam, flat,
slightly floccose at centre, aerial mycelium scarce, margins regular; surface olive-brown (4F5); reverse
olive-brown (4F8/4E4). On OA reaching 64 mm diam, flat, slightly floccose, scarce aerial mycelium,
margins regular; surface dull green (30E5) with grey floccose area; reverse dull green (30E4).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
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Typus: Spain, Catalonia, Priorat, Montsant Natural Park, Arbolí, isolated from small rodent dung, Feb.
2018, I. Iturrieta-González, E. Carvalho & J. Gené (holotype CBS H-23895, culture ex-type CBS 145423
= FMR 17110).
Note: Alternaria fimeti is placed in a clade of section Infectoriae together with A. aconidiophora (see
notes of this latter species).
Alternaria inflata Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829631. Fig. 13.
Etymology: Name refers to the presence of swollen cells in the conidial body.
Asexual morph on PCA: Mycelium superficial and immersed. Hyphae of 1–3 m wide, septate, branched,
hyaline to pale brown, smooth-walled to verruculose. Conidiophores arising laterally or terminally from
aerial hyphae, erect to slightly flexuous, semi- to macronematous, up to 10-septate, commonly
unbranched, 9–73(–105) × 2–5 m, pale brown to brown, smooth-walled, with one terminal
conidiogenous loci or up to three subterminal conidiogenous loci. Conidia solitary or in short chains with
up to four conidia, broadly ellipsoidal or ovoid, 13–41 × 5–14 m, often with some swollen cells
protruding the conidium outline, some with darkened middle transverse septa, (1–)2–3(–5) transverse
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septa, and 0–2 longitudinal or oblique septa per transverse segment, brown, verruculose. Secondary
conidiophores scarcely produced, as an apical extension up to 15 m long, bearing conidia in short
chains. Chlamydospores present, consisting of intercalary, thick-walled, brown swollen cells, up to 8 × 6
m, arranged in chains or in clusters. Sexual morph not observed.
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Culture characteristics: Colonies on PDA reaching 62 mm diam after 1 wk at 25 °C, flat, aerial mycelium
abundant, floccose, margins fimbriate; surface white (1A1); reverse greyish yellow to yellowish white
(4C6/4A2). On PCA attaining 67 mm diam, flat, scarce aerial mycelium, margins regular; surface dull
green to grey (30E4/30B1); reverse dark green to grey (30E4/30B1). On OA reaching 61 mm diam, flat,
loosely floccose, margins regular; surface pale grey (1B1); reverse pale grey (1B1).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
Typus: Spain, Catalonia, Conca de Barberà, Poblet, isolated from rabbit dung, Mar. 2017, J. Guarro & I.
Iturrieta-González (holotype CBS H-23896, culture ex-type CBS 145424 = FMR 16477).
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Notes: Our phylogeny shows that A. inflata belongs to section Pseudoalternaria (Fig. 6). It clustered in a
well-supported clade (86 % BS) with A. altcampina, A. kordkuyana and A. parvicaespitosa, but formed a
single basal lineage representative of a distinct species. Alternaria inflata can be differentiated from all
the species in the section by the production of chlamydospores and by the formation of broadly ellipsoidal
conidia, usually with swollen cells protruding from the conidial body. In addition, A. altcampina also
differs in the production of secondary conidiophores, A. parvicaespitosa in its shorter conidiophores (up
to 70 m) and conidia (10–25 m) (Gannibal & Lawrence 2016), and A. kordkuyana in the production of
longer conidial chains [up 5–8(−10) conidia] and conidia measuring 30–50(−60) × 7–11 m with up to
seven transverse septa (Poursafar et al. 2018).
With the additions of A. altcampina and A. inflata section Pseudoalternaria now comprises seven
species. It is of note, however, that most of these taxa are only known from a single collection. In our
survey of asexual fungi from herbivore dung, we identified several Spanish isolates belonging to other
species in the section, namely A. kordkuyana and A. rosae (Fig. 6). Considering that most of the species
in section Pseudoalternaria are mainly associated with herbaceous plants of the families Brassicaceae,
Ericaceae, Poacea, or Rosaceae, it is not surprising to find these fungi in faeces of herbivorous animals.
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Alternaria lawrencei Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829632. Fig. 14.
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Etymology: Name in honour of Daniel P. Lawrence for his contribution to the taxonomy of the genus
Alternaria.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 1–3 m wide, septate, branched,
pale brown, smooth-walled. Conidiophores macronematous, solitary, arising directly from aerial hyphae,
erect to slightly flexuous, occasionally geniculate at apex, usually unbranched, up to 10-septate, 9–125 ×
3–4(–5) m, brown, smooth-walled, with 1–2 lateral or terminal conidiogenous loci; micronematous
conidiophores also present, reduced to intercalary conidiogenous cells with a single conidiogenous locus
on hyphae. Conidia solitary or in short chains, up to six conidia in the unbranched part, ovoid, obpyriform
or obclavate, 6–71 × 7–15 m, with darkened middle transverse septa, (1–)2–7(–9) transverse, and 0–2(–
3) longitudinal or oblique septa, pale brown to brown, verrucose to tuberculate. Secondary conidiophores
commonly formed apically on conidia as a geniculate extension with several conidiogenous loci, or as
lateral extensions from cells of conidial body, up to 35 m long, producing conidia solitary or in short
chains. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching 68 mm diam after 1 wk at 25 °C, low convex,
cottony, aerial mycelium abundant, margins regular; surface white (1A1); reverse yellowish brown to
greyish yellow (5E7/4B6). On PCA attaining 69 mm diam, low convex, slightly floccose, aerial
mycelium relatively abundant at centre, margins regular; surface yellowish grey to olive (2D2/1E3);
reverse dark green to olive (30F8/1E3). On OA reaching 63 mm diam, loosely floccose at centre, flat and
scarce aerial mycelium towards the periphery, margins regular and diffuse; surface olive (2F4) to olivegrey (2B1); reverse olive to yellowish grey (2F8/2D2).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
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Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park, isolated from goat dung, Oct. 2017, G. Sisó
& C. González-García (holotype CBS H-23897, culture ex-type CBS 145425 = FMR 17004).
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Notes: Although A. lawrencei is clearly placed in section Infectoriae, the multilocus analysis did not
reveal any phylogenetic relationship with any species included in the analysis (Fig. 4). It is of note,
however, that eight species of the section (i.e. A. cerasi, A. cesenica, A. dactylidicola, A. forlicesenensis,
A. hampshirensis, A. litorea, A. murispora and A. poaceicola) could not be included in our analysis due to
their limited molecular data. Nevertheless, A. cesenica, A. dactylidicola, A. forlicesenensis, A.
hampshirensis, A. murispora and A. poaceicola can be distinguished from A. lawrencei by the production
of a sexual morph (Ariyawansa et al. 2015b, Liu et al. 2015, Thambugala et al. 2017, Wanasinghe et al.
2018), A. cerasi by its inequilateral conidia (Potebnia 1907, Simmons 2007), and A. litorea by the
production of shorter primary conidiophores (40–50 m long) and smooth-walled conidia that are 22–32
m long (Simmons 2007).
Alternaria montsantina Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829633. Fig.
15.
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Etymology: Name refers to the place, Montsant Natural Park (Catalonia), where the fungus was collected.
Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 1–7 m wide, septate, branched,
usually forming hyphal coils, subhyaline to pale brown, smooth-walled to verruculose. Conidiophores
macronematous, arising laterally or terminally from aerial hyphae, erect to slightly flexuous, unbranched,
up to 15-septate, 12–137 × 3–6 m, often with geniculate apical portion containing intercalary and
terminal conidiogenous loci, brown, smooth-walled to verruculose. Conidia solitary or in short chains
with up to five conidia, subglobose, ovoid or obpyriform, 8–65 × 6–12 m, with 1–3(–11) transverse
septa, and 0–2 longitudinal or oblique septa, brown, verrucose to tuberculate. Secondary conidiophores
commonly produced apically as a long, often geniculate extension, up to 105 m long and 10-septate,
bearing terminal conidial chains. Sexual morph not observed.
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Culture characteristics: Colonies on PDA reaching 76 mm diam after 1 wk at 25 °C, flat, densely
floccose, aerial mycelium abundant, margins regular; surface pastel grey to greyish yellow (1C1/2C4) and
with a white edge; reverse blond to white (4C4/1A1). On PCA attaining 70 mm diam, flat, loosely
floccose at centre, aerial mycelium moderate, margins regular; surface olive brown to white (4D4/1A1);
reverse olive to white (3D4/1A1). On OA reaching 75 mm diam, flat, cottony, margins regular; surface
yellowish grey to olive (3D2/2F4) and white edge; reverse olive to white (2F4/1A1).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
EP
Typus: Spain, Catalonia, Priorat, Montsant Natural Park, Swamp of Siurana, isolated from an unidentified
twig, Feb. 2018, I. Iturrieta-González, E. Carvalho & J. Gené (holotype CBS H-23898, culture ex-type
CBS 145426 = FMR 17060).
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Notes: Alternaria montsantina is placed in a weakly supported basal clade of the section Infectoriae,
together with A. curvata, A. pseudoventricosa and A. ventricosa. Morphologically, this new species can
be distinguished from A. curvata and A. ventricosa by the absence of curved or inequilateral inflated
conidia. Alternaria montsantina differs from A. pseudoventricosa in the production of longer (12–137
m) and often geniculate primary and secondary conidiophores, bearing solitary conidia or arranged in
short chains (up to five conidia). Conidiophores in A. pseudoventricosa are 30–44 m long, and the
conidial chains include up to 19 conidia.
Alternaria pobletensis Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829634. Fig.
16.
Etymology: Name refers to the place, Poblet (Catalonia), from where the species was collected.
Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 2–5 m wide, branched, pale
brown, septate, smooth-walled. Conidiophores semi- to macronematous, solitary, arising directly from
aerial hyphae, erect to slightly flexuous, occasionally slightly geniculate at apex, unbranched or branched,
up to 8-septate, 14–82 × 4–5(–6) m, brown, smooth-walled, with 1–2 lateral or terminal conidiogenous
loci. Conidia commonly in short, scarcely branched chains, with up to seven conidia, obpyriform or
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obclavate, some ellipsoidal or subcylindrical, 8–50 × 5–20 m, (1–)3–7(–9) transverse septa, often middle
septa darker, and 0–1(–2) longitudinal or oblique septa per transverse segment, pale brown to brown,
smooth-walled or verruculose. Secondary conidiophores commonly produced apically as a short beak up
to 11 m long, or laterally from cells of conidial body, bearing conidia in short chains. Sexual morph not
observed.
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Culture characteristics: Colonies on PDA reaching 46 mm diam after 1 wk at 25 °C, flat, floccose at the
centre, velvety towards the periphery, aerial mycelium moderate, margins regular; surface olive (3F8),
whitish at the periphery; reverse black to yellowish brown (5D5). On PCA attaining 58 mm diam, flat,
velvety, margins regular; surface dark green to dull green (30F8/28D3); reverse dark green to dull green
(30F8/28D3). On OA reaching 55 mm diam, flat, loosely floccose, margins regular; surface greyish green
to dull green (29C3/29E4); reverse dark green to dull green (30F8/30E3).
Cardinal temperature for growth: Optimum 25 °C, maximum 35 °C, minimum 5 °C.
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Typus: Spain, Catalonia, Conca de Barberà, Poblet, isolated from unidentified herbivore dung, Mar.
2017, J. Guarro & I. Iturrieta-González (holotype CBS H-23899, culture ex-type CBS 145427 = FMR
16448).
Notes: Alternaria pobletensis clustered in section Chalastospora in a single branch clearly separated from
the other six species that currently comprise the section (Fig. 7). Other species of section Chalastospora
rarely produce conidia with longitudinal septa (Woudenberg et al. 2013); however, the conidia in A.
pobletensis usually have two or more longitudinal or oblique septa. Its closest relative is A. breviramosa.
This was originally described as Chalastospora ellipsoidea, found on Triticum (Poaceae) in Australia
(Crous et al. 2009a), but later its name was changed to avoid confusion with Alternaria ellipsoidea, an
already described species from section Gypsophilae (Woudenberg et al. 2013). Section Gypsophilae
contains all Alternaria species that occur on Caryophyllaceae (Lawrence et al. 2016). Alternaria
breviramosa differs from A. pobletensis by having shorter conidiophores (up to 25 µm), often reduced to
conidiogenous cells, and ellipsoidal, subcylindrical to fusoid conidia with only transverse septa (Crous et
al. 2009a).
D
Alternaria pseudoventricosa Iturrieta-González, Dania García & Gené, sp. nov. MycoBank MB829635.
Fig. 17.
TE
Etymology: Name refers to the apparent phylogenetic relationship to A. ventricosa.
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Asexual morph on PCA: Mycelium superficial and immersed. Hyphae 1–7 m wide, septate, branched,
hyaline to pale brown, smooth-walled. Conidiophores macronematous, arising laterally from aerial
hyphae, erect to slightly flexuous, up to 4-septate, unbranched, 30–45 × 4–6 m, brown, smooth-walled,
with one terminal conidiogenous locus. Conidia commonly in unbranched chains, with up to 19 conidia,
obpyriform or obclavate, 10–48(–66) × 5–14 m, with darkened middle transverse septa, 1–7 transverse,
0–1 longitudinal or oblique septa, brown to dark brown, verrucose to tuberculate. Secondary
conidiophores scarce, as a beak arising from the conidial body. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching 64 mm diam after 1 wk at 25 °C, flat, cottony at the
centre, floccose towards the periphery, margins regular; surface white (1A1); reverse yellowish white
(4A2). On PCA attaining 62 mm diam, flat towards the periphery, margins regular; surface dark green
(29F4), with tuft of white aerial mycelium at centre; reverse dark green to grey (29F8/29B1). On OA
reaching 67 mm diam, flat, loosely floccose, margins regular; surface dull green (29E4); reverse dull
green (29E4).
Cardinal temperature for growth: Optimum 25 °C, maximum 37 °C, minimum 5 °C.
Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park, isolated from horse dung, Oct. 2017, G. Sisó
& C. González-García (holotype CBS H-23900, culture ex-type CBS 145428 = FMR 16900).
Notes: Alternaria pseudoventricosa and A. ventricosa clustered in an unsupported monophyletic basal
clade in section Infectoriae. They can be differentiated by their conidial morphology. Conidia in A.
ventricosa are usually asymmetric, laterally swollen, and pale cinnamon brown (Roberts 2007). In
contrast, those of A. pseudoventricosa are obpyriform or obclavate and brown to dark brown.
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Authors: I. Iturrieta-González, D. García, M. Hernández-Restrepo & J. Gené
Brunneosphaerella Crous, Stud. Mycol. 64: 31. 2009. Fig. 18.
Classification: Dothideomycetes, Dothideomycetidae, Capnodiales, Mycosphaerellaceae.
DNA barcodes (genus): LSU and ITS.
DNA barcodes (species): chs, rpb2 and tef1. Table 3.
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Type species: Brunneosphaerella protearum (Syd. & P. Syd.) Crous, basionym: Leptosphaeria
protearum Syd. & P. Syd. Epitype and ex-epitype strain designated by Crous et al. (2011): CBS H20335, CBS 130597 = CPC 16338.
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Ascomata amphigenous, immersed to semi-immersed, black, single, gregarious, substomatal, pyriform
or globose with a papillate, periphysate ostiole; ascomatal wall consisting of three strata of slightly
compressed textura angularis, an outer stratum of dark brown, thick-walled cells, becoming paler in
the central stratum, and hyaline, thin-walled in the inner stratum. Pseudoparaphyses absent. Asci
clavate to cylindro-clavate, often curved, tapering to a pedicel, narrowing slightly to a rounded apex
with an indistinct ocular chamber, 8-spored, bitunicate with fissitunicate dehiscence. Ascospores
biseriate, fusiform, broader at the apical end, initially hyaline and 1-septate, becoming yellow-brown
and 3-septate at maturity, slightly constricted at median to supra-median septum (adapted from Crous
et al. 2009b).
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Culture characteristics: Colonies on OA spreading, flat, with sparse to moderate aerial mycelium, lobate
and smooth, rarely feathery, margins. On OA surface pale luteous, smoke grey with submerged iron-grey
margins, or olivaceous grey with iron-grey, pale olivaceous grey and smoke grey patches. On PDA
surface olivaceous grey, sometimes with pale olivaceous grey to smoke grey patches, or smoke grey
with iron-grey margins; reverse iron-grey. On MEA surface pale olivaceous grey, smoke grey, dirty
white with patches of smoke grey, or smoke grey with dirty white and olivaceous grey patches and
submerged iron-grey margins; reverse iron-grey or olivaceous grey.
TE
Optimal media and cultivation conditions: MEA, OA, PDA and SNA at 25 °C under near-ultraviolet
light to promote sporulation.
Distribution: Africa, mainly reported from South Africa. Also reported from Pacific Islands (Hawaii)
and Europe (Portugal and Spain).
EP
Hosts: Protea spp. (Proteaceae).
Disease symptoms: Leaf spots and Brunneosphaerella leaf blight.
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Notes: Brunneosphaerella was introduced by Crous et al. (2009b) to accommodate Leptosphaeria
protearum, which is a major leaf spot and blight pathogen of Protea spp. causing severe losses in
plantations of South African Protea spp. wherever they are cultivated. Morphologically,
Brunneosphaerella is distinct from Leptosphaeria in that ascospores are always brown at maturity,
and asexual morphs have brown, percurrently proliferating conidiogenous cells. A new species
isolated from leaves of Protea repens in South Africa, B. jonkershoekensis, was included in the genus
when it was introduced (Crous et al. 2009b). This species appears to be a serious pathogen of Pr.
repens in the Western Cape Province of South Africa. Subsequently, Crous et al. (2011) described the
third species known from the genus, B. nitidae. This was isolated from the same area as B.
jonkershoekensis, but B. nitidae was isolated from leaves of Pr. nitida, causing leaf spots on this host.
Thus, the genus comprises four species, all of which were isolated from species of Protea in South
Africa. The ITS sequences of the four species are highly similar. However, these can be easily
delimited based on the chs, rpb2 and tef1 sequences.
References: Crous et al. (2009b, 2011), Videira et al. 2017 (morphology and phylogeny).
Brunneosphaerella roupeliae Crous, sp. nov. MycoBank MB829609. Fig. 19.
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Etymology: Name refers to Protea roupeliae, the host species from which it was collected.
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Leaf spots amphigenous, sub-circular, 5–20 mm diam, medium brown, with raised, dark brown
border. Ascomata pseudothecial, amphigenous, black, immersed to erumpent, globose, to 250 µm
diam, with apical ostiole; ascomatal wall of 2–3 layers of brown cells of textura angularis. Asci
aparaphysate, fasciculate, bitunicate, subsessile, ellipsoid-fusoid, straight to slightly curved, 8-spored,
65–110 × 11–15 µm. Ascospores bi- to triseriate, overlapping, guttulate, thick-walled, straight to
slightly curved, obovoid with obtuse ends, widest in middle of apical cell, 3-septate, constricted at
median septum, tapering towards both ends, but more prominently towards lower end, (19–)22–23(–
25) × 5(–6) µm. Ascospores germinating from both ends, becoming brown and verruculose,
constricted at primary septum, with germ tubes parallel to the long axis, ascospore becoming 7–9 µm
diam.
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Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium, and even,
lobate margins, reaching 10 mm diam after 2 wk at 25 °C. On MEA surface pale olivaceous grey,
reverse olivaceous grey; on PDA surface and reverse olivaceous grey, and on OA surface pale luteous.
Typus: South Africa, KwaZulu-Natal Province, Drakensberg, Monks Cowl, on leaves of Protea
roupeliae (Proteaceae), 18 Jan. 2010, A. Wood, HPC 1522 (holotype CBS H-23847, culture ex-type
CPC 32914 = CBS 144602).
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Notes: Brunneosphaerella roupeliae was isolated from the same host genus in South Africa as the
other three species of the genus, Protea. The ITS and LSU sequences located this species in the genus
Brunneosphaerella since both sequences showed more than 99 % of nucleotide similarity with the extype strains of the other three species. The rpb2 sequence showed a nucleotide similarity of 95.86 %
with the ex-epitype strain of B. protearum, 95.45 % with the ex-epitype strain of B. jonkershoekensis,
and 94.5 % with the ex-type strain of B. nitidae. The tef1 sequence showed a nucleotide similarity of
95.66 % with the ex-epitype strain of B. protearum and the ex-type strain of B. nitidae, and 94.44 %
with the epitype strain of B. jonkershoekensis. Brunneosphaerella roupeliae produces the shortest
ascospores of the genus [(19–)22–23(–25) µm in B. roupeliae vs. (25–)27–34(–37) µm in B.
jonkershoekensis vs. (20–)24–28(–30) µm in B. nitidae vs. (20–)23–26(–30) µm in B. protearum].
TE
Authors: P.W. Crous, J.Z. Groenewald & Y. Marin-Felix
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Elsinoe Racib., Parasit. Alg. Pilze Java's (Jakarta) 1: 14. 1900. Figs 20, 21.
Synonyms: Sphaceloma de Bary, Ann. Oenol. 4: 165. 1874.
Manginia Viala & Pacottet, C. r. hebd. Séanc. Acad. Sci., Paris 139: 88. 1904.
Melanobasidium Maubl., Bull. Soc. mycol. Fr. 22: 69. 1906.
Plectodiscella Woron., Mykol. Zentbl. 4: 232. 1914.
Isotexis Syd., in Sydow & Petrak, Annls mycol. 29: 261. 1931.
Melanobasis Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 224. 1931.
Melanodochium Syd., Annls mycol. 36: 310. 1938.
Bitancourtia Thirum. & Jenkins, Mycologia 45: 781. 1953.
Kurosawaia Hara, List of Japanese Fungi, 4th Edn: 172. 1954.
Uleomycina Petr., Sydowia 8: 74. 1954.
Melanophora Arx, Verh. K. ned. Akad. Wet., tweede sect. 51: 43. 1957.
Classification: Dothideomycetes, Dothideomycetidae, Myriangiales, Elsinoaceae.
Type species: Elsinoe canavaliae Racib. Type or reference material not available.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, rpb2, tef1. Table 4. Fig. 22.
Ascostromata solitary, aggregated, or gregarious, wart-like, or as small distinctively coloured
elevations, or pulvinate, immersed to semi-immersed, globose to subglobose, white, pale yellow or
brown, soft, multi-loculate, locules scattered in upper part of ascostromata; ascostromatal wall
composed of pseudoparenchymatous cells of textura globulosa to textura angularis; locules with few
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to numerous asci inside each locule, ostiolate; ostiole minute. Periphyses absent. Asci 8-spored,
bitunicate, fissitunicate, saccate to globose, with a minute pedicel, and ocular chamber. Ascospores
irregularly arranged, oblong or fusiform with slightly acutely rounded ends, with 2–3 transverse septa,
hyaline, smooth-walled, lacking a sheath. Acervuli or sporodochia subepidermal,
pseudoparenchymatous. Conidiophores hyaline to pale brown, polyphialidic. Conidiogenous cells
formed directly from the upper cells of the pseudoparenchyma, mono- to polyphialidic, integrated or
discrete, determinate, hyaline to pale brown, without visible periclinal thickening. Conidia hyaline,
smooth-walled, aseptate, ellipsoidal, guttulate (adapted from Fan et al. 2017).
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Culture characteristics: Colonies on MEA, slow growing, raised, irregular, erumpent, folded or
cerebriform, smooth and irregular margins, with sparse to moderate white to grey aerial mycelium. On
MEA, surface white to pale luteous, cinnamon, sepia, apricot, saffron with or without purplish grey in
centre, brown with apricot margins, rosy buff in centre with cinnamon margins, livid red, scarlet red with
diffuse red pigment in agar, or iron-grey; reverse umber, ochreous, iron-grey, dark vinaceous, or centre
scarlet and orange with cinnamon margins.
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Optimal media and cultivation conditions: MEA, OA, PDA, SNA and WA at 22 °C under nearultraviolet light (12 h light, 12 h dark).
Distribution: Worldwide.
Hosts: Wide range of hosts, including some economically important crops such as avocado, cassava,
citrus, grapevines, ornamentals such as poinsettias, field crops and woody hosts.
Disease symptoms: Scab, leaf and fruit spot and anthracnose disease.
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Notes: Elsinoe comprises plant pathogenic species that cause scab and spot anthracnose on a wide
range of hosts, including some economically important crops and ornamentals. The disease symptoms
that these species produce are easily recognisable, being known as “signature-bearing diseases”, for
the cork-like appearance of older infected tissues with scab-like appearance. Also, these can produce
other disease symptoms often called anthracnose, as in the case of infected grapevines (Barrus &
Horsfall 1928, Jenkins 1947, Farr et al. 1989, Pan 1994, Phillips 1994, Gottwald 1995). However, the
use of this name is confusing since it is used much broader to include diseases caused by
Colletotrichum. Although many species of Elsinoe causing scab disease have been described, only
few of them cause important diseases (Holliday 1980), having the main impact on the appearance of
the harvested product and its market acceptability rather than on crop productivity (Swart et al. 2001).
Species of Elsinoe seem to be host-specific since 77 of the 81 species accepted in the present study
occur on only one host species or genus.
Elsinoe and its asexual morph, Sphaceloma, were recently reviewed by Fan et al. (2017). In that
study, 26 new combinations were proposed for the species originally placed in Sphaceloma.
Moreover, eight new species were introduced and 13 epitypes were designated. Based on phylogenetic
data, Fan et al. (2017) accepted 75 species in the genus. However, E. banksiae, which was described
by Swart et al. (2001), and three species described by Crous et al. (2016a), i.e. E. eelemani, E.
eucalyptigena and E. preissianae, were not included in that study. Our phylogenetic analysis
corroborated the placement of these species in the main well-supported clade representing the genus
Elsinoe, except for E. eucalyptigena, whose placement remains unknown and, therefore it is not
considered an accepted species of Elsinoe in the present study. Moreover, another three new species
have been subsequently described by Crous et al. (2018), i.e. E. banksiigena, E. elaeocarpi and E.
leucopogonis. However, in our phylogenetic analysis, the two first species were not located in the
Elsinoe s. str. clade, and are thus excluded from the genus at present. Therefore, hitherto a total of 79
species are accepted, plus the new species described in the present study. Unfortunately, there are no
cultures and molecular data of the type species of the genus, E. canavaliae, which needs to be
epitypified in order to clarify its phylogenetic position.
References: Fan et al. 2017 (morphology and phylogeny).
Elsinoe picconiae Crous, sp. nov. MycoBank MB829611. Fig. 23.
Etymology: Name refers to Picconia, the host from which this fungus was collected.
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Conidiomata sporodochial or acervular on leaves, medium brown, up to 250 µm diam; conidiomatal
wall composed of textura angularis. Conidiophores subcylindrical to doliiform, hyaline to pale
brown, smooth-walled, 0–1-septate, unbranched, 10–15 × 3–4 µm. Conidiogenous cells polyphialidic,
hyaline, smooth-walled, subcylindrical to doliiform, 5–8 × 3–4 µm. Conidia hyaline, aseptate,
ellipsoid, apex obtuse, base truncate, (4–)5–6(–7) × (2–)2.5 µm.
Culture characteristics: Colonies erumpent, spreading, with sparse aerial mycelium, folded surface,
and smooth, lobate margins, reaching 7 mm diam after 2 wk at 25 °C. On MEA surface rust, reverse
sienna; on PDA surface coral, reverse bay; on OA surface scarlet with diffuse scarlet pigment.
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Typus: Spain, Tenerife, Los Silos, leaf of Picconia excelsa (Oleaceae), 12 Mar. 2017, A. van Iperen,
HPC 2063 (holotype CBS H-23848, culture ex-type CBS 145026 = CPC 33648).
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Notes: Elsinoe picconiae is related to E. freyliniae, E. oleae and E. salicina. However, E. picconiae
can be easily distinguished by its narrower conidia [(2–)2.5 µm vs. (2.5–)3–4 µm in E. freyliniae, 3–6
µm in E. oleae, and (2.5–)3–4.5(–5) µm in E. salicina]. Moreover, E. picconiae is the first species of
the genus isolated from Picconia excelsa.
Elsinoe veronicae Crous, Thangavel & Y. Marín, sp. nov. MycoBank MB829610. Fig. 24.
Etymology: Name refers to Veronica, the host from which this fungus was collected.
Conidiomata sporodochial, erumpent, 80–200 µm diam, based on a pale brown stroma, giving rise to
densely aggregated conidiophores. Conidiophores hyaline to pale brown, smooth-walled, subcylindrical,
1–2-septate, 15–30 × 2.5–3.5 µm, unbranched or branched above. Conidiogenous cells integrated,
subcylindrical, hyaline, smooth-walled, 7–10 × 2.5–3.5 µm, polyphialidic. Conidia solitary, aggregating
in mucoid mass, aseptate, hyaline, smooth-walled, guttulate, ellipsoid, apex subobtuse, tapering at base to
truncate hilum, (4–)5–5.5(–6.5) × 2.5(–3) µm.
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Culture characteristics: Colonies erumpent, spreading, surface folded with moderate aerial mycelium,
and even, lobate margins, reaching 25 mm diam after 2 wk. On MEA surface brick, reverse cinnamon; on
PDA surface brick to scarlet, reverse brick in centre, scarlet in outer region; on OA surface scarlet with
diffuse scarlet pigment.
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Typus: New Zealand, Auckland, St. John, Morrin Road, on Veronica sp. (Scrophulariaceae), 2013, R.
Thangavel (holotype CBS H-23865, culture ex-type CBS 145362 = CPC 34137 = T17_00408D).
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Notes: Elsinoe veronicae is closely related to E. othonnae. Although the ITS sequences of both species
showed more than 99 % nucleotide similarity, the rpb2 sequences showed only 96.95 % similarity.
Morphologically, these are also similar differing mainly in their conidial size [(4–)5–5.5(–6.5) × 2.5(–3)
µm in E. veronicae vs. (5–)6–7 × (2.5–)3(–4) µm in E. othonnae]. Moreover, E. veronicae was found on
Veronica (Scrophulariaceae) in New Zealand, while E. othonnae has only been reported on Othonna
(Asteraceae) in South Africa (Crous et al. 2015c).
Authors: Y. Marin-Felix, R. Thangavel & P.W. Crous
Exserohilum K.J. Leonard & Suggs, Mycologia 66: 290. 1974. Fig. 25.
Synonyms: Setosphaeria K.J. Leonard & Suggs, Mycologia 66: 294. 1974.
Luttrellia Khokhr. & Gornostaĭ (as ‘Lutrellia’; non Luttrellia Shearer), Vodorosli, Griby i Mkhi Dal’nego
Vostoka [Algae, Fungi and Mosses of the Soviet Far-East] (Vladivostok): 80. 1978.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae.
Type species: Exserohilum turcicum (Pass.) K.J. Leonard & Suggs, basionym: Helminthosporium
turcicum Pass. Ex-epitype and ex-epitype strain designated by Hernández-Restrepo et al. (2018): CBS H23323, CBS 690.71.
DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, gapdh, rpb2. Table 5. Fig. 26.
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Ascomata superficial, immersed or erumpent, globose to ellipsoid, unilocular, dark brown to black, with
or without a neck, ostiolate, with simple rigid setae around the ostiolar apex and on the upper half of the
ascoma where they are often mixed with hyaline, filiform, septate hyphae; ascomatal wall composed of
pseudoparenchymatous cells, dark brown and thick-walled on the outside, but with more or less hyaline
cells towards the inside, cells of textura angularis. Pseudoparaphyses filiform, hyaline, septate, branched,
anastomosing. Asci arising from a basal cushion of thin-walled pseudoparenchymatous cells, bitunicate,
1–8-spored, cylindrical to cylindrical-clavate, short or moderately long-stalked, thick-walled, with an
apical nasse and fissitunicate dehiscence. Ascospores fusoid, hyaline to pale brown, smooth-walled, 2–6
or rarely more transversely septate, constricted at the septa, surrounded by a hyaline mucilaginous sheath
which often extends some distance beyond the ends of the spore. Conidiophores macronematous,
mononematous, septate, cylindrical, olivaceous brown to brown, smooth-walled to verruculose, often
geniculate above. Conidiogenous cells integrated, terminal and intercalary, sympodial, mono- or
polytretic, cicatrized; conidiogenous nodes smooth to rough. Conidia fusiform, cylindrical or obclavate,
straight to curved, multi-distoseptate, with a protruding hilum (adapted from Hernández-Restrepo et al.
2018).
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Culture characteristics: Colonies on PDA brown or grey olivaceous to olivaceous black, sometimes
white, pale grey, hairy, cottony to powdery, margins fimbriate.
Optimal media and cultivation conditions: Sterilized Zea mays leaves placed on 1.5 % WA or PDA at 25
°C under near-ultraviolet light (12 h light, 12 h dark) to induce sporulation.
Distribution: Worldwide.
Host: Mainly pathogens of grasses, but some also on non-grass hosts. Other substrates where they can be
found include river sediments, soil, grains, plant debris, and humans. Exserohilum rostratum has been
reported as a human pathogen.
Disease symptoms: In plants: leaf blight, leaf spots, melting out, root rot, and foot rot, among others. In
humans: skin infections, keratitis, non-invasive allergies, invasive sinusitis, and disseminated infections.
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Notes: Exserohilum is differentiated from the closely related Bipolaris, Curvularia and Pyrenophora by
producing conidia with a protruding hilum (Leonard & Suggs 1974). Recently the taxonomy and
phylogeny of Exserohilum has been revisited by Hernández-Restrepo et al. (2018). Based on
morphological and molecular data 11 phylogenetic species are accepted (Fig. 26, Table 5). Three species
were excluded from the genus, namely Ex. novae-zelandiae relocated to Sporidesmiella, and Ex. paspali
and Ex. sorghicola to Curvularia, while another 15 species were retained in Exserohilum, although some
were doubtful. Species in Exserohilum are morphologically very variable and a molecular analysis is
required for a correct species identification.
The type species of the genus, Ex. turcicum (= Helminthosporium turcicum), was described from Italy
causing northern leaf blight of corn (Passerini 1876, Saccardo 1886). Other species attacking
economically significant crops include Ex. pedicellatum, causing root rot on maize and brown lesions on
wheat roots (Henry 1924, Sivanesan 1987), and Ex. rostratum, producing leaf spot on banana, maize and
wheat, foot rot in wheat, damping off of sugarcane seedlings, blackening and seed germination failure in
cereals (Drechsler 1923, Leonard 1976, Sivanesan 1987, Lin et al. 2011).
Previously, three different species, Ex. longirostratum, Ex. macginnisii and Ex. rostratum, were
recognised as human pathogens (McGinnis et al. 1986, Padhye et al. 1986, de Hoog et al. 2000, da Cunha
et al. 2012). However, a multi-locus phylogenetic analysis (Hernández-Restrepo et al. 2018),
demonstrated that they are actually the same phylogenetic species. Exserohilum rostratum has been
reported as an agent of phaeohyphomycosis and sometimes causing life-threatening infections in humans
(McGinnis et al. 1986, Padhye et al. 1986, Aquino et al. 1995, Adler et al. 2006). This species was
recently implicated in an outbreak of fungal meningitis associated with contaminated methylprednisolone
in the USA (Kainer et al. 2012).
References: Drechsler 1923, 1934, Luttrell 1963, Leonard & Suggs 1974, Sivanesan 1987 (taxonomy,
morphology and pathogenicity), Leonard 1976 (sexual/asexual connection), de Hoog et al. 2000, da
Cunha et al. 2012 (human pathogens), Hernández-Restrepo et al. 2018 (morphology, phylogeny, review).
Author: M. Hernández-Restrepo
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Neosetophoma Gruyter et al., Mycologia 102: 1075. 2010. Fig. 27.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
Type species: Neosetophoma samarorum (Desm.) Gruyter et al., basionym: Phoma samarorum Desm.
Epitype and ex-epitype strain designated by Gruyter et al. (2010): CBS H-20319, CBS 138.96.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 6. Fig. 29.
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DNA barcode (genus): LSU. Fig. 28.
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Ascomata ostiolate, globose to subglobose, solitary to gregarious, dark brown to black, immersed to
slightly erumpent or superficial, smooth; ascomatal wall composed of 2–4 layers of brown to reddishbrown or dark brown to black cells of textura angularis to textura prismatica. Hamathecium
comprising numerous, septate, cellular or filamentous pseudoparaphyses, embedded in a hyaline
gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical to clavate, short pedicellate, with
a furcate pedicel, apically rounded, with a minute or indistinct ocular chamber. Ascospores
overlapping, 1–3-seriate, hyaline or subhyaline when young, becoming pale yellow, pale brown or
yellowish brown to brown at maturity, 1–5-septate, straight to slightly curved, fusoid or narrowly
fusoid, with rounded or acute ends, constricted or not at the septum, enlarged at the second cell below
apex, guttulate, smooth-walled, without any mucilaginous sheath and appendages. Conidiomata
pycnidial, solitary to confluent, immersed or superficial, globose to subglobose or irregular, with
mycelial outgrowths, or confluent, unilocular, occasionally multi-locular, with papillate ostioles,
sometimes developing long necks, honey, olivaceous, olivaceous black, pale brown, brown, dark
brown, or black, with up to 10 layers of pseudoparenchymatal cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, extending percurrently at
apex, hyaline, doliiform to ampulliform, determinate, hyaline, smooth-walled. Conidia hyaline, slightly
yellowish or pale brown, 0–3(–4)-septate, ellipsoidal, cylindrical, subcylindrical, fusiform, or teardrop shape, straight to curved, usually attenuate at one end, or apex and base obtuse, or sometimes with
bluntly rounded to truncate base, continuous or constricted at the septa, smooth-walled, often guttulate
(asexual morph description adapted from Gruyter et al. 2010).
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Culture characteristics: Colonies flat, with a moderate amount of aerial mycelium. On PDA surface
fluffy, circular or irregular, margins entire or filiform, white, pale grey, grey, greenish grey, or mouse
grey; reverse yellowish, yellowish grey, greyish white, grey olivaceous, or dark brown. On MEA pale
grey to almost white, buff, or brown to dark brown; reverse dark brown, or buff with patches of grey
olivaceous.
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Optimal media and cultivation conditions: CMA, PDA, PNA and OA at 25 °C under continuous nearultraviolet light to promote sporulation.
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Distribution: Asia, Europe and North America.
Hosts: Pathogens or saprophytes found on a wide range of host including Cirsium arvense (Asteraceae),
Clematis vitalba (Ranunculaceae), Iris germanica (Iridaceae), Lunaria annuna (Brassicaceae), Malva
sp. (Malvaceae), Phlox paniculata (Polemoniaceae), Phragmites australis (Poaceae), Rosa canina
(Rosaceae), Sambucus spp., Viburnum opulus (Caprifoliaceae), and Urtica dioica (Urticaceae). Also
isolated from soil.
Disease symptoms: Leaf spots.
Notes: Neosetophoma was introduced by de Gruyter et al. (2010) to accommodate Phoma samarorum,
which is a pathogen causing leaf spots of grasses. Subsequently, 12 new species have been added to this
genus, all of which appear to be saprobes, except for Nph. iranianum, which was isolated from soil
(Karunarathna et al. 2017), and Nph. lunariae, which is endophytic (Hernández-Restrespo et al. 2016a).
This genus is characterised by globose to irregular conidiomata with papillate ostioles, and yellowish
to brownish conidia usually attenuated at one end, less frequent with apex and base obtuse or with a
bluntly rounded to truncate base. The sexual morph was observed for the first time by Tibpromma et al.
(2017), when Nph. garethjonesii was introduced. Subsequently, four new species producing a sexual
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morph have been described, of which only one also produces the asexual morph, namely Nph.
shoemakeri (Hyde et al. 2018).
References: de Gruyter et al. 2010, Quaedvlieg et al. 2013, Tibpromma et al. 2017, Hyde et al. 2018,
Wanasinghe et al. 2018 (morphology and phylogeny).
Neosetophoma aseptata Crous, R.K. Schumach. & Y. Marín, sp. nov. MycoBank MB829639. Fig.
30.
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Etymology: Name refers to its aseptate conidia, which have never been observed in the other species
of the genus.
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Conidiomata solitary, brown, erumpent, globose, 250–350 µm diam with 1–3 ostioles; conidiomatal wall
of 3–4 layers of brown cells of textura angularis. Conidiophores reduced to conidiogenous cells lining
inner cavity, hyaline, smooth-walled, subcylindrical to ellipsoid, phialidic with minute collarette, 4–8 ×
4–5 µm. Conidia solitary, aseptate, hyaline, smooth-walled, subcylindrical to ellipsoid, apex obtuse, base
truncate, (3.5–)4–5 × (1.5–)2 µm.
Culture characteristics: Colonies flat, spreading, surface folded, with moderate aerial mycelium and
even, lobate margins, reaching 50 mm diam after 2 wk. On MEA surface pale olivaceous grey, reverse
umber with diffuse umber pigment; on PDA surface olivaceous grey, reverse umber with diffuse umber
pigment; on OA surface saffron with patches of grey olivaceous.
Typus: Germany, near Berlin, moist meadow, on Viburnum opulus (Caprifoliaceae), 7 Jun. 2017, R.K.
Schumacher, HPC 2131, RKS 123 (holotype CBS H-23866, culture ex-type CBS 145363 = CPC 33919).
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Notes: Neosetophoma aseptata was located in a large, well-supported clade (96 % BS / 1 PP) comprising
Nph. clematidis, Nph. iranianum, Nph. lunariae, Nph. rosae and Nph. shoemakeri. Neosetophoma
aseptata can be easily distinguished by its aseptate conidia, being 1-septate in Nph. shoemakeri, 1–3septate in Nph. iranianum and Nph. rosae, 3-septate in Nph. clematidis, and (1–)3(–4)-septate in Nph.
lunariae. Moreover, Nph. aseptata produces the smallest conidia in the complex [(3.5–)4–5 × (1.5–)2 µm
in Nph. aseptata vs. 4–6 × 2–4 m in Nph. iranianum vs. 7.5–10.5 × 2.5–3 µm in Nph. shoemakeri vs. 8–
14 × 1.5–3 µm in Nph. rosae vs. 11–15 × 2–4 m in Nph. clematidis vs. (10–)14–17(–22) × (2.5–)3 m in
Nph. lunariae]. Neosetophoma clematidis produces the largest conidiomata in this complex, being up to
475 m diam (up to 300 m in Nph. aseptata and Nph. lunaria, up to 180 m in Nph. shoemakeri, up to
130 m in Nph. rosae, up to 120 m in Nph. iranianum).
Neosetophoma aseptata is the first species isolated from Viburnum. Neosetophoma samarorum and
Nph. sambuci are reported in Sambucus spp., which is a member of the same family, Caprifoliaceae.
Neosetophoma phragmitis Crous, R.K. Schumach. & Y. Marín, sp. nov. MycoBank MB829640. Fig.
31.
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Etymology: Name refers to Phragmites, the host from which this fungus was collected.
Conidiomata solitary, pycnidial, brown, globose, 180–200 µm diam, neck papillate with central ostiole;
conidiomatal wall of 3–4 layers of brown cells of textura angularis. Conidiophores reduced to
conidiogenous cells lining inner cavity, hyaline, smooth-walled, ampulliform, 5–8 × 2.5–3 µm,
proliferating percurrently at apex. Conidia solitary, pale brown, smooth-walled, aseptate, straight, apex
obtuse, base truncate, (3–)4–5(–6) × 2 µm.
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and feathery margins,
covering dish in 2 wk. On MEA surface and reverse ochreous; on PDA surface and reverse hazel; on OA
surface hazel.
Typus: Germany, near Berlin, on leaf sheath of Phragmites australis (Poaceae), 16 Apr. 2016, R.K.
Schumacher, HPC 1178 (holotype CBS H-23867, culture ex-type CBS 145364 = CPC 30680).
Notes: In the phylogenetic analysis based on ITS, LSU and rpb2 sequences, Nph. phragmitis was located
in an independent branch removed from the other species of the genus. This is the first species isolated
from Phragmitis australis (Poaceae). Neosetophoma poaceicola is the only species that was reported
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before on a member of the Poaceae, being isolated from a grass host. However, both species are not
related, and Nph. phragmitis only produces the asexual morph, while only the sexual morph was observed
in Nph. poaceicola.
Neosetophoma sambuci Crous, R.K. Schumach. & Y. Marín, sp. nov. MycoBank MB829641. Fig.
32.
Etymology: Name refers to the genus Sambucus, the host from which this fungus was collected.
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Conidiomata solitary, erumpent, brown, pycnidial, globose, 150–200 µm diam, with central ostiole;
conidiomatal wall of 3–4 layers of brown textura angularis. Conidiophores reduced to conidiogenous
cells lining the inner cavity, hyaline, smooth-walled, ampulliform, phialidic, 4–6 × 3–4 µm. Conidia
solitary, pale brown, smooth-walled, guttulate, subcylindrical, apex obtuse, base truncate, aseptate,
becoming 1-septate and swollen (ellipsoid) with age, (5–)7–8(–10) × (2–)2.5(–3) µm.
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Culture characteristics: Colonies flat, spreading, covering dish in 2 wk, with sparse to moderate aerial
mycelium. On MEA surface ochreous, reverse umber; on PDA surface and reverse olivaceous grey; on
OA surface ochreous.
Typus: Germany, near Berlin, on twig of Sambucus nigra (Caprifoliaceae), 11 Mar. 2016, R.K.
Schumacher, HPC 1072 (holotype CBS H-23868, culture ex-type CBS 145365 = CPC 30357).
Authors: Y. Marin-Felix & P.W. Crous
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Notes: In the phylogenetic analysis based on ITS, LSU and rpb2 sequences, Nph. sambuci was located in
a well-supported clade (97 % BS / 0.98 PP) together with Nph. garethjonesii, Nph. samarorum and Nph.
rosigena. Neosetophoma garethjonesii can be easily distinguished from the other species by only
producing a sexual morph. The other three species produce an asexual morph, and can be differentiated
by the size of their conidia [4–16 × 1.5–3 µm in Nph. samarorum vs. (5–)7–8(–10) × (2–)2.5(–3) µm in
Nph. sambuci vs. 4–6 × 1.5–2.5 µm in Nph. rosigena]. Moreover, conidia in Nph. sambuci are pale brown
while in Nph. samarorum they are slightly yellowish and in Nph. rosigena they are olivaceous brown.
Neosetophoma sambuci was isolated from twigs of Sambucus nigra (Caprifoliaceae). The only species
previously reported on this host was Nph. samarorum.
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Neostagonospora Quaedvl. et al., Stud. Mycol. 75: 364. 2013. Fig. 33.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
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Type species: Neostagonospora caricis Quaedvlieg et al. Holotype and ex-type strain: CBS H-21306,
CBS 135092.
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DNA barcode (genus): LSU. Fig. 28.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 7. Fig. 34.
Conidiomata immersed, pycnidial, globose, exuding a pale luteous to creamy conidial mass;
conidiomatal wall composed of 2–3 layers of pale brown cells of textura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells phialidic, hyaline, smooth-walled, aggregated,
lining the inner cavity, ampulliform to doliiform, tapering at apex with prominent periclinal
thickening. Conidia hyaline, smooth-walled, granular, thin-walled, narrowly fusoid-ellipsoidal to
subcylindrical, apex subobtusely rounded, base truncate, widest in middle, aseptate or transversely
euseptate, becoming constricted with age (adapted from Quaedvlieg et al. 2013).
Culture characteristics: Colonies flat, spreading, erumpent, circular or undulate, smooth to velvety, even
margins, with sparse to moderate aerial mycelium. On PDA, surface dirty white, greyish sepia to
isabelline; reverse pale white to pale pink, pale pink with white edge, luteous or olivaceous grey to pale
olivaceous grey.
Optimal media and cultivation conditions: PDA and sterilised Carex leaves or Anthriscus stem placed on
1.5 % water agar at 25 °C under continuous near-ultraviolet light to promote sporulation.
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Distribution: Africa (South Africa), Asia (Russia and Taiwan), Australia and Europe (Italy and the
Netherlands).
Hosts: Arrhenatherum elatius, Phragmites australis, Sorghum halepense and Spinifex littoreus
(Poaceae), Carex acutiformis (Cyperaceae) and Elegia cuspidata (Restionaceae).
Disease symptoms: Leaf spots.
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Notes: Neostagonospora was introduced by Quaedvlieg et al. (2013) to accommodate two taxa
associated with leaf spots on Carex acutiformis (Nst. caricis) and Elegia cuspidata (Nst. elegiae).
However, its pathogenicity remains unclear since Koch’s postulates have not been completed.
Subsequently, another foliicolous fungus was included in this genus, Nst. spinificis (Yang et al. 2016),
associated with green tissues and leaf spots of Spinifex littoreus. The most recently introduced species
are both saprobes on members of Poaceae: Nst. arrhenatheri and Nst. phragmitis (Thambugala et al.
2017). Further studies are needed to prove the pathogenicity of members included in this genus.
Neostagonospora is similar to Stagonospora since both produce pycnidial conidiomata with
euseptate, hyaline, fusoid-ellipsoidal to subcylindrical conidia, but Neostagonospora is distinguished
by having conidiogenous cells that are phialidic, with prominent periclinal thickening (Quaedvlieg et
al. 2013).
In our phylogenetic analysis based on ITS and LSU sequences of members of the family
Phaeosphaeriaceae (Fig. 28), Nst. artemisiae, which was the most recently described species
(Wanasinghe et al. 2018), is not included in the clade that represents the genus Neostagonospora, being
located in the Septoriella clade. Therefore, this species is excluded from Neostagonospora and
transferred to Septoriella (see Septoriella below).
References: Quaedvlieg et al. 2013, Yang et al. 2016, Thambugala et al. 2017 (morphology and
phylogeny).
Neostagonospora sorghi Crous & Y. Marín, sp. nov. MycoBank MB829612. Fig. 35.
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Etymology: Name refers to the genus Sorghum, the host from which this fungus was collected.
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Conidiomata solitary, erumpent, pycnidial, brown, globose, 180–200 µm diam, with central ostiole;
conidiomatal wall of 2–3 layers of brown textura angularis. Conidiophores reduced to conidiogenous
cells lining the inner cavity. Conidiogenous cells hyaline to pale brown, smooth-walled, ampulliform,
phialidic with visible periclinal thickening and collarette, 5–8 × 4–5 µm. Conidia solitary, aseptate,
ellipsoid with obtuse ends, straight, guttulate, (4–)5–6 × (2–)2.5(–3) µm.
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Culture characteristics: Colonies flat, spreading with moderate aerial mycelium and even, lobate margins,
covering dish in 2 wk. On MEA surface hazel to isabelline, reverse isabelline; on PDA surface isabelline
with scarlet outer margins, reverse brown vinaceous; on OA surface isabelline with patches of scarlet.
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Typus: Australia, Western Australia, Denmark, Mount Lindesay Walk trail, on Sorghum halepense
(Poaceae), 19 Sep. 2015, P.W. Crous, HPC 697A (holotype CBS H-23869, culture ex-type CBS 145366
= CPC 29239).
Notes: Neostagonospora sorghi is the first species of the genus reported from Sorghum, and occurring in
Australia. It is closely related to Nst. caricis, which is associated with leaf spots on Carex acutiformis.
However, Nst. sorghi can be distinguished from this species and from other species of the genus by its
aseptate conidia. Moreover, Nst. sorghi produces smaller conidia than Nst. caricis [(4–)5–6 × (2–)2.5(–3)
µm in Nst. sorghi vs. (10–)13–16(–19) × (3–)3.5(–4) µm in Nst. caricis].
Authors: Y. Marin-Felix & P.W. Crous
Nothophoma Q. Chen & L. Cai, Stud. Mycol. 82: 212. 2015. Fig. 36.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Didymellaceae.
Type species: Nothophoma infossa (Ellis & Everh.) Q. Chen & L. Cai, basionym: Phoma infossa Ellis &
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Everh., J. Mycol. 4: 102. 1888. Neotype and ex-neotype strain designated by Aveskamp et al. (2009):
CBS H-20145, CBS 123395.
DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): rpb2, tub2. Table 8. Fig. 37.
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Conidiomata pycnidial, globose to elongated, or irregular, superficial or immersed into the agar,
solitary or confluent, ostiolate, sometimes with an elongated neck; conidiomatal wall
pseudoparenchymatous, multi-layered, outer wall pigmented. Conidiogenous cells phialidic, hyaline,
smooth-walled, ampulliform to doliiform, sometimes flask-shaped. Conidia hyaline but incidentally
brown, smooth- and thin-walled, aseptate, ovoid or ellipsoidal, eguttulate or guttulate.
Chlamydospores elongated barrel-shaped, olivaceous brown, in chains. Sexual morph unknown
(adapted from Chen et al. 2015, Crous et al. 2017b).
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Culture characteristics: Colonies on OA yellow/green to olivaceous grey/brown, dull green, or
translucent, aerial mycelium tenuous, sometimes margins irregular and whitish, flattened or effused,
compact, floccose.
Optimal media and cultivation conditions: OA or sterile pine needles placed on OA under nearultraviolet light (12 h light, 12 h dark) to promote sporulation at 25 °C.
Distribution: Worldwide.
Hosts: Wide host range, mainly occurring as pathogens, and also endophytes or saprobes, on
Amaryllidaceae, Anacardiaceae, Fabeceae, Fagaceae, Haemodoraceae, Malvaceae, Oleaceae,
Rosaceae, Rhamnaceae, Rutaceae. Also isolated from other substrates and environments, such as soil,
fungi and human infections.
Disease symptoms: Leaf spots, stem cankers, brown spot of fruits, shoot canker.
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Notes: Nothophoma was one of the genera established recently in order to delineate a more natural
classification for the Ascochyta-Didymella-Phoma complex (Chen et al. 2015). Currently this genus
comprises nine species, including five Phoma species previously classified in Phoma, and four species
that were recently proposed (Crous et al. 2016b, 2017b, Valenzuela-Lopez et al. 2018). Within
Nothophoma morphological differences between species are insignificant, and phylogenies based on
multi-locus sequence data are primarily used to distinguish species.
Species in this genus are seed- and soil-borne endophytes or pathogens mainly causing leaf spots
and stem canker of cultivated crops and plants, such as groundnut and cotton. Some species are
mycophylic on other fungi or occur in soil, as well as in the respiratory secretion of a patient with
pneumonia or in a human bronchial wash sample (Boerema et al. 2004, Aveskamp et al. 2009, 2010,
Chen et al. 2015, Crous et al. 2016b, 2017b, Valenzuela-Lopez et al. 2018).
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References: Boerema et al. 2004 (morphology and pathogenicity), Aveskamp et al. 2010, Chen et al.
2015, Crous et al. 2016b, 2017b, Valenzuela-Lopez et al. 2018 (morphology and phylogeny).
Authors: Q. Chen & L. Cai
Parastagonospora Quaedvl. et al., Stud. Mycol. 75: 362. 2013. Fig. 38.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
Type species: Parastagonospora nodorum (Berk.) Quaedvlieg et al., basionym: Depazea nodorum
Berk. Reference strain: CBS 110109.
DNA barcode (genus): LSU. Fig. 28.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 9. Fig. 39.
Ascomata perithecial, immersed, globose, becoming depressed, dark brown to black, with central ostiole
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with upper region slightly papillate; ascomatal wall thin- or thick-walled, composed of 2–6 layers of
brown cells of textura angularis. Pseudoparaphyses filiform, hyaline, septate. Asci bitunicate, clavate,
cylindrical, narrowly fusoid or curved, shortly stipitate, thick-walled, 8-spored. Ascospores fusoid or
ellipsoidal, hyaline or subhyaline to pale brown, smooth-walled, transversely 3-euseptate, cells above
central septum often broader than the lower ones, with acute rounded ends, constricted or not at each
septum, sometimes with distinct oil droplets in each cell. Conidiomata pycnidial, brown to black,
erumpent or immersed to semi-immersed, subepidermal, globose to subglobose, ampulliform, or
obpyriform, with central papillate ostiole, exuding creamy or pinkish conidial mass; conidiomatal wall
composed of 2–4 layers of brown cells of textura angularis, or composed of 1–5 outer layers of dark
brown cells and 1–3 inner layers of hyaline cells of textura angularis, or composed of an outer layer of
brown to dark brown cells of textura globosa and an inner layer of pale brown to hyaline cells of textura
angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, hyaline,
smooth-walled, aggregated, lining the inner cavity, ampulliform to subcylindrical, broadly cylindrical or
broadly conical, with percurrent proliferation near apex. Conidia hyaline or subhyaline, smooth-walled,
thin- or thick-walled, cylindrical, subcylindrical or fusiform, granular to multi-guttulate, with obtuse or
subobtuse apex and truncate base, rarely rounded at both ends, straight to gently curved, sigmoid,
transversely 1–9-euseptate, sometimes constricted at the septa (adapted from Quaedvlieg et al. 2013).
Culture characteristics: Colonies flat, with aerial mycelium, white to pink, olivaceous, grey or vinaceous
buff.
Optimal media and cultivation conditions: Sterilised Carex leaves placed on 1.5 % WA at 25 °C under
continuous near-ultraviolet light to promote sporulation.
Distribution: Worldwide.
Hosts: Pathogens or saprophytes of grass (Poaceae). Species of Parastagonospora are directly or
indirectly responsible for significant annual crop losses worldwide on wheat, barley and rye.
Disease symptoms: Leaf, glume and node spots.
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Notes: Parastagonospora was recently introduced in order to accommodate a clade of several common
and serious cereal pathogens that had been previously been placed in the genera Septoria/Stagonospora
or Leptosphaeria/Phaeosphaeria (Quaedvlieg et al. 2013). This genus differs from Stagonospora
mainly in the sexual morph, being phaeosphaeria-like in Parastagonospora and didymella-like in
Stagonospora.
In the phylogenetic analysis based only on the ITS and LSU sequences of representative members of
the family Phaeosphaeriaceae (Fig. 28), all the species of Parastagonospora were located in a wellsupported clade (0.98 PP), except for P. phoenicicola. The ex-type strain of this latter species clustered
in well-supported clade (84 % BS) representing the genus Phaeosphaeria. Therefore, a new combination
is proposed for this taxon.
In our phylogenetic analysis (Fig. 39), P. cumpignensis, P. dactylidis and P. minima grouped in the
same well-supported clade (79 % BS / 0.96 PP) without significant phylogenetic distance. The three
species all have been isolated from Thailand on dead stems of Dactylis (Li et al. 2015, 2016a). Only ITS
sequences are available for P. cumpignensis and P. dactylidis, and nucleotide similarity for this locus for
all three species is 100 %. Therefore, these three species are reduced to synonymy. The same problem is
found in P. forlicesenica, which is one of the most recently described species in the genus (Thambugala
et al. 2017). Based on ITS, P. forlicesenica shares a nucleotide similarity of 99.8% with P. avenae.
Therefore, further studies should be done to confirm if P. forlicesenica represents a separate species or
should be synonymized with P. avenae.
Species of Parastagonospora are pathogens or saprophytes of grasses, being directly or indirectly
responsible for significant annual crop losses worldwide. Parastagonospora avenae causes minor leaf
blotch of barley and rye, while it is considered an important pathogen of oats (Cunfer 2000).
Parastagonospora nodorum is known primarily as a major necrotrophic pathogen of wheat that causes
leaf and glume blotch, but also infects barley, on which it is considered as not economically important
(Cunfer 2000, Oliver et al. 2012).
References: Cunfer 2000 (pathology and morphology), Oliver et al. 2012 (pathology, genomics and
host resistence), Quaedvlieg et al. 2013, Li et al. 2015, Thambugala et al. 2017 (morphology and
phylogeny).
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Parastagonospora dactylidis W.J. Li et al., Mycosphere 6: 691. 2015.
Synonyms: Parastagonospora minima W.J. Li, et al., Mycosphere 6: 691. 2015.
Parastagonospora cumpignensis Tibpromma et al., Fungal Diversity 78: 48. 2016.
Typus: Italy, Province of Arezzo, Passo della Consuma, on dead stem of Dactylis sp. (Poaceae), 19
Jun. 2012, Erio Camporesi (holotype MFLU 15-0693, culture ex-type MFLUCC 13-0375 = ICMP
20774 = KUMCC15-0131).
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Additional materials: Italy, Province of Arezzo, Passo della Consuma, on dead stem of Dactylis sp.
(Poaceae), 19 Jun. 2012, Erio Camporesi, MFLUCC 13-0376 = ICMP 20776 = KUMCC15-0132;
ibid., Campigna, Santa Sofia, ForlìCesena Province, on dead stem of Dactylis glomerata (Poaceae),
23 Jun. 2012, Erio Camporesi, MFLUCC 13-0573.
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Notes: In our phylogenetic analysis (Fig. 39), the ex-type strains of P. cumpignensis (MFLUCC 130573), P. dactylidis (MFLUCC 13-0375) and P. minima (MFLUCC 13-0376) grouped in the same
well-supported clade (79 % BS / 0.96 PP) without significant phylogenetic distance. The ITS sequences
of the three species showed a nucleotide similarity of 100 %. Moreover, all of them were isolated from
from Thailand on dead stems of Dactylis (Li et al. 2015, 2016a). Therefore, the three species are
herewith reduced to synonymy.
Parastagonospora novozelandica Crous, Thangavel & Y. Marín, sp. nov. MycoBank MB829668.
Fig. 40.
Etymology: Name refers to New Zealand, the country where this fungus was isolated.
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Culture nearly sterile, with only a few conidiomata observed. Conidiomata solitary, pycnidial, dark
brown, globose, 180–200 µm diam, with central ostiole; conidiomatal wall with 3–6 layers of pale brown
textura angularis. Conidiophores reduced to conidiogenous cells lining the inner cavity. Conidiogenous
cells hyaline, smooth, ampulliform to subcylindrical, 6–8 × 2.5–5 µm, proliferating percurrently at apex.
Conidia solitary, hyaline to pale olivaceous, smooth, guttulate, subcylindrical, straight, apex subobtuse,
base truncate, 1-septate, (9–)11–13(–16) × (2–)2.5(–3) µm.
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Culture characteristics: Colonies flat, spreading, reaching 60 mm diam after 2 wk, with moderate aerial
mycelium, and even, smooth margins. On MEA surface greenish olivaceous to umber, reverse olivaceous
to umber; on OA surface brown vinaceous.
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Typus: New Zealand, Browns Bay, on unidentified grass (Poaceae), Nov. 2015, R. Thangavel (holotype
CBS H-23903, culture ex-type T15–06960B = CPC 29613= CBS 145416).
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Notes: Parastagonospora novozelandica is related to P. allouniseptata. Both species produce 1-septate
conidia, but these can be easily distinguished based on their conidial dimensions [(9–)11–13(–16) × (2–
)2.5(–3) µm in P. novozelandica vs. 16–22 × 2.5–3.5 m in P. allouniseptata].
Parastagonospora phragmitis Crous & Y. Marín, sp. nov. MycoBank MB829667. Fig. 41.
Etymology: Name reflects the genus Phragmites from which this fungus was isolated.
Conidiomata solitary, pycnidial, brown, globose, 250–300 µm diam, with central ostiole; conidiomatal
wall with 3–6 layers of pale brown cells of textura angularis. Conidiophores reduced to conidiogenous
cells lining the inner cavity. Conidiogenous cells hyaline, smooth, ampulliform to doliiform, 7–10 × 8–9
µm, proliferating percurrently at apex. Conidia solitary, hyaline to pale olivaceous, smooth, guttulate,
subcylindrical-fusoid, straight to slightly curved, with prominent taper in upper third to subobtuse apex,
widest in middle to lower third, base truncate, 3-septate, (18–)23–25(–27) × (3–)4 µm.
Culture characteristics: Colonies flat, spreading, covering dish after 2 wk, with moderate aerial
mycelium, and smooth, lobate margins. On MEA surface saffron, reverse sienna; on PDA surface saffron
to sienna, reverse sienna; on OA surface pale luteous to saffron.
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Typus: Australia, New South Wales, Sussex Inlet, on Phragmites sp. (Poaceae), 27 Nov. 2015, P.W.
Crous, HPC 1785 (holotype CBS H-23902, culture ex-type CPC 32075 = CBS 143446).
Notes: Parastagonospora phragmitis is related to P. fusiformis. However, P. phragmitis produces an
asexual morph, while in P. fusiformis only the sexual morph has been observed. Moreover, P. phragmitis
is the first species of the genus reported on Phragmites.
Description and illustration: Crous et al. (2016a).
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Phaeosphaeria phoenicicola (Crous & Thangavel) Y. Marín & Crous, comb. nov. MycoBank
MB829700.
Basionym: Parastagonospora phoenicicola Crous & Thangavel, Persoonia 37: 349. 2016.
Typus: New Zealand, Auckland, Botany road, on leaves of Phoenix canariensis (Arecaceae), 2015,
R. Thangavel (holotype CBS H-22892, culture ex-type CPC 28711 = CBS 142107).
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Notes: In the phylogenetic analysis based on ITS and LSU sequences (Fig. 28), the ex-type strain of
this species was located in the well-supported clade (84 % BS) representing the genus Phaeosphaeria.
Morphologically, this species produces conidia more similar to Phaeosphaeria than to
Parastagonospora, since these are subcylindrical, mostly straight, while in Parastagonospora the
conidia tend to be sigmoid and longer than in P. phoenicicola. Based on morphology and molecular
data, the new combination, Phaeosphaeria phoenicicola, is herewith proposed.
Authors: Y. Marin-Felix, R. Thangavel & P.W. Crous
Phaeosphaeriopsis M.P.S. Câmara et al., Mycol. Res. 107: 519. 2003. Fig. 42.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
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DNA barcode (genus): LSU. Fig. 28.
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Type species: Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S. Câmara et al., basionym:
Cryptosphaeria glaucopunctata Grev. [as “glauco-punctata”]. Epitype and ex-epitype strain
designated by Thambugala et al. (2014): MFLU 14-0029, MFLUCC 13-0265 = ICMP 20199.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 10. Fig. 43.
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Ascomata solitary or aggregated, immersed, subepidermal to erumpent, pushing up flaps of the
epidermis, globose to pyriform, often papillate, solitary or gregarious in a stroma of
scleroplectenchyma or dark brown cells of textura angularis, often surrounded by septate, brown
hyphae extending into the host tissues. Asci 8-spored, bitunicate, cylindrical to broadly fusoid, short
stipitate, with visible apical chamber. Ascospores uni- to triseriate, cylindrical, broadly rounded at
apex, tapering to narrowly rounded base, 4–5-septate, first septum submedian, often constricted,
medium brown, echinulate, punctate or verrucose. Asexual morph coniothyrium-like or
phaeostagonospora-like. Conidiomata pseudoparenchymatous, sometimes of scleroplectenchyma.
Conidiogenous cells lining locule, ampulliform, hyaline, proliferating percurrently, resulting in
inconspicuous annellations. Conidia cylindrical, with bluntly rounded ends, 0–3-septate, yellowish
brown, punctate (Quaedvlieg et al. 2013).
Culture characteristics: Colonies flat or rarely slightly raised, spreading, feathery, velvety or floccose,
with sparse to moderate aerial mycelium, circular or lobate, margins smooth or rarely slightly radiating.
On PDA, surface white, dirty white, pinkish white, primrose, pale grey or pale luteous; reverse dirty
white, light to dark grey, luteous or olivaceous buff. On MEA, surface dirty white, pale luteous, or white
to cream at the margins, pale yellowish to yellowish brown in the middle and pale brown to brown or
orange-brown at the centre, with small white to grey droplets; reverse luteous, umber with patches of
dirty white, isabelline in the middle and cinnamon in outer region, or white to cream at the margins,
brown to orange-brown in the middle and pale yellowish at the centre.
Optimal media and cultivation conditions: On MEA, PDA or OA at 25 °C.
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Distribution: Worldwide.
Hosts: Pathogens or saprophytes on Agapanthus praecox (Alliaceae), Agave spp. and Yucca spp.
(Agavaceae), Aloe sp. (Aloaceae), Dracaena lourieri, Dracaena sp. and Nolina erumpens
(Dracaenaceae), Grevillea sp. (Proteaceae), Phormium spp. (Phormiaceae) and Ruscus spp.
(Ruscaceae).
Disease symptoms: Leaf spots and leaf blight.
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Notes: The genus Phaeosphaeriopsis was introduced by Câmara et al. (2003) to accommodate some
species of Paraphaeosphaeria that were not congeneric based on phylogenetic data.
Phaeosphaeriopsis is characterised by having uni- or multi-loculate stromata and 4–5-septate
ascospores, and coniothyrium-like and phaeostagonospora-like asexual morphs, while
Paraphaeosphaeria produces 2-septate ascospores and has a microsphaeropsis-like asexual morph
(Câmara et al. 2003, Quaedvlieg et al. 2013). Phaeosphaeriopsis is related to Acericola, which is a
genus recently introduced to accommodate a saprobic fungus found on dead twigs of Acer campestre
(Hyde et al. 2017). Unfortunately, it appears that the LSU sequence of Acericola is incorrect (Crous et
al. 2019a).
In our phylogenetic analysis based on ITS and LSU sequences, 11 species are accepted in the genus
Phaeosphaeriopsis, and four strains located in independent branches are introduced as new species.
Species included in Phaeosphaeriopsis are saprobes or presumed pathogens. The type species,
Phaeosphaeriopsis glaucopunctata, is associated with leaf spot and necrosis of Ruscus aculeatus
(Câmara et al. 2003, Golzar & Wang 2012). Phaeosphaeriopsis agapanthi and Phs. dracaenicola are
also associated with necrotic leaf spots of Agapanthus precox and Dracaena lourieri, respectively
(Phookamsak et al. 2014b, Crous et al. 2016b).
References: Câmara et al. 2003, Quaedvlieg et al. 2013, Thambugala et al. 2014 (morphology and
phylogeny).
Phaeosphaeriopsis aloes Crous & Y. Marín, sp. nov. MycoBank MB829642. Fig. 44.
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Etymology: Name refers to Aloe, the host from which this fungus was collected.
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Conidiomata solitary, brown, pycnidial, globose, 150–180 µm diam, with central ostiole, 30–40 µm diam;
conidiomatal wall of 3–4 layers of brown cells of textura angularis. Conidiophores reduced to
conidiogenous cells lining the inner cavity, hyaline, smooth-walled, ellipsoid, phialidic, 4–6 × 3–4 µm.
Conidia solitary, aseptate, straight, verruculose, golden-brown, subcylindrical, apex obtuse, base bluntly
rounded, (4–)5(–6) × 3(–3.5) µm.
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Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium, covering dish
in 2 wk. On MEA surface honey, reverse cinnamon; on PDA surface and reverse sepia; on OA surface
saffron.
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Typus: USA, California, on leaves of Aloe sp. (Aloaceae), 6 Aug. 2016, P.W. Crous, HPC 1326 (holotype
CBS H-23870, culture ex-type CBS 145367 = CPC 31480).
Notes: Phaeosphaeriopsis aloes is related to Phs. obtusispora. This latter species only produces the
sexual morph, while only the asexual morph has been observed in our new species.
Phaesphaeriopsis aloes and Phs. aloicola, both described here, are the first species of the genus found
on a member of the family Aloaceae, an Aloe sp. However, the two species are not related.
Phaeosphaeriopsis aloes produces an asexual morph, while in Phs. aloicola only the sexual morph has
been observed.
Phaeosphaeriopsis aloicola Crous & Y. Marín, sp. nov. MycoBank MB829643. Fig. 45.
Etymology: Name refers to the host genus Aloe from which this fungus was collected.
Ascomata solitary, aggregated, erumpent, brown, globose, 150–200 µm diam, with papillate neck and
central ostiole, 40–50 µm diam; ascomatal wall of 4–6 layers of brown cells of textura angularis.
Pseudoparaphyses hyaline, smooth-walled, hyphae-like, 2–3 µm diam, anastomosing, branched,
ACCEPTED MANUSCRIPT
intermingled among asci. Asci bitunicate, subcylindrical, apex obtuse with well-defined apical chamber, 2
µm diam, fasciculate, short stipitate, 70–100 × 10–12 µm. Ascospores bi- to triseriate, subcylindrical,
straight to slightly curved, (2–)3-septate, at times slightly swollen in second cell from apex, medium
brown, verruculose, ends obtuse, (19–)22–25(–26) × (4–)5(–6) µm.
Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and even,
lobate margins, reaching 30 mm diam on PDA, covering dish on MEA and OA. On MEA surface buff,
outer region sienna, reverse sienna; on PDA surface and reverse buff to sienna; on OA surface scarlet.
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Typus: USA, California, on leaves of Aloe sp. (Aloaceae), 6 Aug. 2016, P.W. Crous, HPC 1306 (holotype
CBS H-23871, culture ex-type CBS 145368 = CPC 31454).
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Notes: Phaeosphaeriopsis aloicola is related to Phs. agapanthi and Phs. triseptata. Morphologically, Phs.
aloicola is similar to Phs. triseptata since both produce verruculose, 3-septate ascospores. However, both
species differ in the size of ascomata (up to 200 m in Phs. aloicola vs. up to 110 m in Phs. triseptata),
asci (70–100 × 10–12 m in Phs. aloicola vs. 56−70 × 7.5−9 m in Phs. triseptata) and ascospores [(19–
)22–25(–26) × (4–)5(–6) m in Phs. aloicola vs. 14.5−18× 3−4 m in Phs. triseptata]. Moreover, Phs.
triseptata also produces an asexual morph, which has not been observed in Phs. aloicola.
Phaeosphaeriopsis agapanthi only produces an asexual morph. The three species were isolated from
different hosts in different families. Phaeosphaeriopsis agapanthi was isolated from Agapanthus precox
(Amaryllidaceae), Phs. aloicola from Aloe sp. (Aloaceae), and Phs triseptata from Ruscus aculeatus
(Asparagaceae). Moreover, Phs. aloicola was found in the USA while the other species have been
reported from Europe. Phaeosphaeriopsis aloes was also isolated from Aloe in California. For
comparison see notes of Phaeosphaeriopsis aloes.
Phaeosphaeriopsis grevilleae Crous & Y. Marín, sp. nov. MycoBank MB829644. Fig. 46.
Etymology: Name refers to the host genus Grevillea from which this fungus was collected.
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Conidiomata solitary, pycnidial, scattered, globose, 180–250 µm diam, with central ostiole, exuding black
mucoid conidial mass; conidiomatal wall of 2–3 layers of brown cells of textura angularis.
Conidiophores reduced to conidiogenous cells lining inner cavity, hyaline, smooth-walled, ampulliform,
phialidic, 5–7 × 3–5 µm. Conidia solitary, aseptate, medium brown, verruculose, ellipsoid to ovoid, (4–
)5(–6) × (3–)3.5(–4) µm.
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Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and feathery margins,
reaching 35 mm diam on MEA, covering dish on PDA and OA. On MEA surface and reverse scarlet; on
PDA surface ochreous, reverse sienna; on OA surface scarlet with patches of ochreous.
Typus: Australia, Queensland, leaves of Grevillea sp. (Proteaceae), 14 Jul. 2009, P.W. Crous, (holotype
CBS H-23872, culture ex-type CBS 145369 = CPC 17003).
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Notes: In the phylogenetic analysis based on ITS, LSU, rpb2, tef1 and tub2, this species was located in an
independent branch. This is the first species reported on Grevillea, which is a member of the Proteaceae.
Phaeosphaeriopsis pseudoagavacearum Crous & Y. Marín, sp. nov. MycoBank MB829645. Fig. 47.
Etymology: Named after its similarity to Phaeosphaeriopsis agavacearum.
Conidiomata solitary, globose, brown, pycnidial, 200–250 µm diam, with central ostiole, 15–20 µm diam;
conidiomatal wall of 2–3 layers of brown cells of textura angularis. Conidiophores reduced to
conidiogenous cells lining the inner cavity, hyaline, smooth-walled, ampulliform, phialidic, 5–7 × 3–4
µm. Conidia solitary, golden-brown, verruculose, thick-walled, straight to slightly curved, 1-septate,
subcylindrical, apex obtuse, base bluntly rounded, (6–)8–9(–10) × 4(–4.5) µm.
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and even, smooth
margins, covering dish in 2 wk. On MEA surface ochreous, reverse bay; on PDA surface sienna with bay
outer region, reverse bay; on OA surface sienna with scarlet margins.
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Typus: France, Domaine la Fraysse, Valgorge, on leaves of Agave sp. (Asparagaceae), 15 Jul. 2010,
P.W. Crous (holotype CBS H-23873, culture ex-type CBS 145370 = CPC 18383).
Notes: Phaeosphaeriopsis pseudoagavacearum is closely related to Phs. agavacearum. Morphologically
these species are also similar in producing verruculose, aseptate conidia. However, they differ in the size
of their conidiomata (up to 180 m diam in Phs. agavacearum vs. up to 250 m diam in Phs.
pseudoagavacearum) and conidia [(5–)6–7(–9) × 3(–4) m in Phs. agavacearum vs. (6–)8–9(–10) × 4(–
4.5) µm in Phs. pseudoagavacearum].
Pleiocarpon L. Lombard & D. Aiello, IMA Fungus 8: 73. 2017. Fig. 48.
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Authors: Y. Marin-Felix & P.W. Crous
Classification: Sordariomycetes, Hypocreomycetidae, Hypocreales, Nectriaceae.
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Type species: Pleiocarpon strelitziae L. Lombard & D. Aiello. Holotype and ex-type strain: CBS H22967, CBS 142251.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, his3, rpb2, tef1, tub2. Table 11.
Conidiophores simple or aggregated, forming sporodochia; simple conidiophores arising laterally or
terminally from aerial mycelium, solitary to loosely aggregated, unbranched or sparsely branched,
septate, bearing up to two conidiogenous cells. Conidiogenous cells monophialidic, cylindrical,
tapering slightly towards the apex. Macroconidia cylindrical to subcylindrical, hyaline, straight to
curved, 1−6-septate, apex or apical cell typically slightly bent to one side and minutely beaked, base
with sometimes visible, centrally located or laterally displaced hilum. Microconidia absent or
abundant, aseptate, hyaline, ellipsoid to ovoid or subcylindrical, straight to slightly curved, with
clearly laterally displaced hilum. Chlamydospores absent or solitary, globose, brown, thich-walled,
guttulate. Sexual morph not observed (adapted from Aiello et al. 2017).
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Culture characteristics: Colonies on PDA with sparse to moderate aerial mycelium, even, smooth, with
lobate margins; surface and reverse umber or cinnamon to honey.
Optimal media and cultivation conditions: On PDA, MEA, OA or SNA with sterile filter paper and
carnation leaf pieces at 25 °C.
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Distribution: Italy and Sri Lanka.
Host: Livistona rotundifolia (Arecaceae) and Strelitzia reginae (Strelitziaceae).
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Disease symptoms: Basal rot and wilt.
Notes: Pleiocarpon was recently introduced by Aiello et al. (2017) to accommodate a new species
isolated from potted plants of Strelitzia reginae in an ornamental nursery located in eastern Sicily that
had a new basal rot disease. This basal stem rot disease resulted in the detachment of the roots from
the stem. Moreover, the diseased plants displayed symptoms of general wilting and rot of the internal
foliage. Pathogenicity tests indicated that Pl. strelitziae, was highly aggressive, killing all inoculated
test plants within 2 mo (Aiello et al. 2017).
The phylogenetic analysis based on ITS, LSU, tef1 and tub2 demonstrated that Pleiocarpon is
closely related to the genus Thelonectria, with both genera being characterised by cylindrocarpon-like
asexual morphs (Aiello et al. 2017). Recently, Thelonectria was segregated by introducing three new
genera, Cinnamomeonectria, Macronectria and Tumenectria (Salgado-Salazar et al. 2016). These four
related genera are mostly found on bark of exposed wood of dead, dying or diseased trees, and are
rarely associated with small cankers and root rots (Chaverri et al. 2011, Salgado-Salazar et al. 2016).
Moreover, Pleiocarpon can be distinguished from Thelonectria and these three new genera by the
absence of a sexual morph.
Hitherto, Pleiocarpon was monospecific. Here we introduce a new species isolated from Livistona
rotundifolia (Arecaceae) in Sri Lanka, causing root and corm rot.
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References: Aiello et al. 2017 (morphology, pathogenicity and phylogeny).
Pleiocarpon livistonae Crous & Quaedvl., sp. nov. MycoBank MB829613. Fig. 49.
Etymology: Name refers to Livistona, the host from which this fungus was collected.
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Conidiophores simple, solitary or aggregated, forming sporodochia, arising from superficial hyphae,
branched, 2–4-septate, 50–120 × 6–8 µm. Conidiogenous cells monophialidic, cylindrical, tapering
slightly towards apex, 10–30 × 4–5 µm, forming conidia in false chains that eventually aggregate in a
mucoid mass. Macroconidia hyaline, smooth-walled, subcylindrical, 1–6-septate, curved, apex
subobtuse, with base sometimes visible as lateral hilum; 1–3 septate conidia (21–)28–34(–45) × (5–)6
µm; 4–6-septate conidia (45–)55–65(–70) × (5–)6 µm. Chlamydospores solitary, globose, brown,
thick-walled, guttulate, 15–20 µm diam.
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Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, lobate
margins, reaching 60 mm diam after 2 wk at 25 °C. On MEA surface ochreous, reverse umber; on
PDA surface and reverse umber; on OA surface umber.
Typus: Sri Lanka, on Livistona rotundifolia (Arecaceae), W. Quaedvlieg, NAK Tuinbouw, INS-1720656D (holotype CBS H-23849, culture ex-type CBS 145030 = CPC 34576).
Notes: Pleiocarpon livistonae is distinguished from Pl. strelitziae by the absence of microconidia, the
production of chlamydospores, and the septation and size of its macroconidia (1–6-septate, up to 70
µm in Pl. livistonae vs. 1–5-septate, up to 50 µm in Pl. strelitziae). Pleiocarpon livistonae is
phylogenetically close but clearly differentiated from Pl. strelitziae based on ITS, his3, rpb2, tef1 and
tub2 sequence similarity (96 %, 85 %, 92 %, 94 %, and 91 %, respectively). Moreover, Pl. livistonae
was isolated from Livistona rotundifolia (Arecaceae) in Sri Lanka, while Pl. strelitziae was found on
Strelitzia reginae (Strelitziaceae) in Italy.
Authors: Y. Marin-Felix, W. Quaedvlieg & P.W. Crous
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Pyrenophora Fr., Summa veg. Scand. 2: 397. 1849. Fig. 50.
Synonyms: Polytrichia Sacc., Syll. fung. (Abellini) 1: 451. 1882.
Pleospora subgen. Scleroplea Sacc., Syll. fung. (Abellini) 2: 277. 1883.
Neilreichina Kuntze, Revis. gen. pl. (Leipzig) 2: 862. 1891.
Scleroplea (Sacc.) Oudem., Verslag. Meded. K. Akad. Wetensch., Afd. Natuurk., ser. 3 9: 152. 1900.
Drechslera S. Ito, Proc. Imp. Acad. Japan 6: 355. 1930.
Marielliottia Shoemaker, Canad. J. Bot. 76: 1559. 1999.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae.
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Type species: Pyrenophora phaeocomes (Rebent.) Fr., basionym: Sphaeria phaeocomes Rebent.
Neotype specimen designated by Shoemaker (1961): UPS 170980. Representative strain: DAOM
222769.
DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, gapdh, tef1. Table 12. Fig. 51.
Ascomata perithecial, immersed, becoming erumpent to near superficial, solitary or scattered, globose
to subglobose, broadly or narrowly conical, smooth-walled, with central ostiole; necks papillate,
covered with brown to reddish brown setae, which are darkened at the base; ascomatal wall
comprising 2–4 layers of brown, thick-walled cells of textura angularis. Pseudoparaphyses not
observed. Asci 8-spored, bitunicate, fissitunicate, clavate to subcylindrical, with a short, broad pedicel,
with a distinct ocular chamber surrounded by a large apical ring. Ascospores 2–3-seriate, muriform,
constricted at the septum, smooth-walled, surrounded by a mucilaginous sheath. Conidiophores semito macronematous, mononematous, sometimes caespitose, straight or flexuous, often geniculate,
usually unbranched, sometimes branched, pale brown to brown, rarely subhyaline to pale brown.
Conidiogenous cells polytretic, integrated, terminal, frequently becoming intercalary, sympodial,
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cylindrical, smooth-walled, or less frequently verruculose, cicatrized. Conidia solitary, in certain
species also sometimes catenate or forming secondary conidiophores which bear conidia,
acropleurogenous, simple, straight or curved, cylindrical, ellipsoidal or obclavate, less frequently
subglobose, obpyriform or fusiform, tapering towards apex, straw-coloured or pale to dark brown or
olivaceous brown, sometimes the end cells are paler than the intermediate ones, smooth-walled or
verruculose, pseudoseptate; hila protuberant or flat, darkened, thickened (adapted from Ellis 1971,
Ariyawansa et. al. 2014).
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Culture characteristics: Colonies flat or umbonate, cottony, sometimes granular or powdery, with
moderate to abundant aerial mycelium, sometimes with sparse aerial mycelium, margins fringed,
sometimes arachnoid. On PDA smoke-grey to olivaceous or olivaceous grey, primrose to greyish
yellow-green, greenish grey to olivaceous black, olivaceous black with patches white for the aerial
mycelium, honey to isabelline, orange to umber, cinnamon with centre white due to the aerial mycelium,
greyish sepia to fuscous black, or fuscous grey with margins buff; reverse olivaceous, olivaceous black
with or without margins, primrose or luteous, fuscous black with or without margins transparent or buff,
honey to isabelline, sienna to umber with margins luteous, or cinnamon with centre brick to dark brick.
On MEA white to pale greenish glaucous or buff, greyish sepia to pale mouse grey or mouse grey,
smoke-grey to pale olivaceous grey, glaucous grey to greenish grey, pale vinaceous to vinaceous buff,
purplish grey with margins vinaceous buff, luteous with margins white to pale smoke-grey, vinaceous
buff to hazel with margins white and saffron, pale greenish grey to greenish grey, or fuscous black with
margins luteous; reverse olivaceous black, smoke-grey to olivaceous grey with middle white due to the
aerial mycelium, fuscous black with or without margins luteous or olivaceous or buff to cinnamon or
honey to isabelline, chestnut with margins luteous to rust, blood colour with margins luteous or scarlet,
chestnut with margins luteous, or orange to sienna. On OA smoke-grey, olivaceous, olivaceous black,
hazel, buff, cinnamon, olivaceous grey with margins luteous, greyish sepia to fuscous black with
margins brick, or orange to umber with margins transparent; reverse olivaceous to olivaceous black,
smoke-grey to olivaceous grey, olivaceous black with margins transparent or luteous, leaden grey to
leaden black, buff with centre fuscous black and margins olivaceous, isabelline with centre olivaceous,
orange to umber or greyish sepia to fuscous black with margins transparent, olivaceous grey to
olivaceous black with margins transparent, or fuscous black with margins brick, or transparent with
centre brick to dark brick.
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Optimal media and cultivation conditions: On PDA, PNA, OA and MEA to induce sporulation of the
asexual morph, while for the sexual morph Sach's agar with sterilised rice or wheat straw at 25 °C is
used.
Distribution: Worldwide, mainly in Australia, Europe, New Zealand and North America.
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Hosts: Wide host range, occurring as pathogens, saprobes or endophytes. Mainly found in members of
Poaceae, being pathogens of cereals and grasses, including barley, oats and wheat. The most common
genera belonging to Poaceae which this genus is associated to are Agropyron, Agrostis, Avena, Bromus,
Dactylis, Festuca, Hordeum, Lolium, Poa and Triticum, among others. Pyrenophora species are also
reported from other genera outside this family, such as Protea and Leucospermum in the Proteaceae.
Disease symptoms: Leaf spots, leaf blight, leaf blotch, net blotch, melting out, head rot, foot rot, seedborne diseases, among others.
Notes: Pyrenophora is characterised by immersed to semi-immersed ascomata with necks covered
with brown to reddish brown setae, lack of pseudoparaphyses, clavate to saccate asci, usually with a
large apical ring, and muriform terete (cylindrical, frequently circular in section but narrowing to one
end) ascospores (Ariyawansa et al. 2014). The asexual morph was known as Drechslera and it is
characterised by brown, transversely septate conidia similar to those found in Bipolaris and
Curvularia. In order to properly delineate these three genera, phylogenetic analyses using sequence
data of different loci (i.e. LSU, SSU, ITS, gapdh and rpb2) were performed (Zang & Berbee 2001,
Ariyawansa et al. 2014, Manamgoda et al. 2014). The synonymy of Drechslera with Pyrenophora
was recently discussed by Ariyawansa et al. (2014). However, there are still a large number of species
which await treatment. Three new combinations are introduced here, i.e. Py. nisikadoi, Py. poae and
Py. wirreganensis. The main problem encountered is the lack of type material of the already known
species, and this resulted in few molecular studies being performed in the past.
Species delimitation in Pyrenophora based on morphology alone is complicated since many
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species have overlapping characters, similar to what is observed in Bipolaris and Curvularia (MarinFelix et al. 2017). Therefore, molecular data (ITS, gapdh and rpb2) are essential for an accurate
identification of species of Pyrenophora. In our phylogenetic analysis, 21 species are accepted and an
additional six are newly described.
Pyrenophora includes saprobic and plant pathogenic species with a worldwide distribution,
commonly associated with members of the family Poaceae. Some species are serious plants pathogens,
e.g. Py. teres, which is a necrotrophic pathogen causing net blotch in barley (Crous et al. 1995, Louw et
al. 1995, Campbell et al. 1999, 2002), and Py. tritici-repentis, which causes tan spot of wheat (Lamari &
Bernier 1989, Balance et al. 1996, Abdullah et al. 2017) in all the major wheat growing areas of the
world resulting in 3–50 % yield losses (Lamari & Bernier 1989).
References: Sivanesan 1987 (morphology and pathogenicity), Zhang & Berbee 2001, Ariyawansa et
al. 2014 (morphology and phylogeny).
Pyrenophora avenicola Y. Marín & Crous, sp. nov. MycoBank MB829614. Fig. 52.
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Etymology: Name refers to the host genus Avena from which it was isolated.
Hyphae hyaline, branched, septate, verrucose, 2–6(–7.5) m. Conidiophores arising in groups, septate,
straight or flexuous, sometimes geniculate at upper part, usually cells decrease in size towards apex,
sometimes branched, cells walls thicker than those of vegetative hyphae, semi- to macronematous,
subhyaline to pale brown, usually paler towards apex, not swollen at the base, up to 350 m long, 4.5–
7 m wide. Conidiogenous cells verruculose, terminal or intercalary, proliferating sympodially,
subhyaline to pale brown, subcylindrical to swollen, 14–33.5(–43.5) × 6.5–9.5 m. Conidia
verruculose, straight, middle cells enlarged, cylindrical to obclavate, tapering towards apex, pale
brown to brown, end cells rarely slightly paler, 1–4(–5)-distoseptate, 21.5–71.5 × 9.5–15 m, forming
secondary conidiophores or conidia; hila protuberant, darkened, thickened, 3–4.5
m.
Chlamydospores, microconidiation and sexual morph not observed.
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Culture characteristics: Colonies on MEA covering dish after 1 wk at 25 °C, with abundant aerial
mycelium, umbonate; surface fuscous black, margins luteous; reverse smoke-grey to olivaceous grey,
centre white due to the aerial mycelium. Colonies on PDA covering dish, with abundant aerial
mycelium; surface smoke-grey to olivaceous grey; reverse fuscous black to black, margins
transparent. Colonies on OA covering all dish, with moderate to abundant aerial mycelium; surface
smoke-grey to olivaceous grey; reverse olivaceous grey to olivaceous black, margins transparent.
Typus: Sweden, Uppsala, on seed of Avena sp. (Poaceae), unknown date, C. Svensson (holotype CBS
H-23840, culture ex-type CBS 307.84).
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Notes: Pyrenophora avenicola is closely related to Py. chaetomioides. Moreover, both species have
been found on the same host, Avena. However, Py. avenicola can be easily distinguished by its shorter
conidiophores (up to 350 m in Py. avenicola vs. 1 mm in Py. chaetomioides) and smaller conidia
(21.5–71.5 × 9.5–15 m in Py. avenicola vs. 25–140 × 12–22 m in Py. chaetomioides) with less
septa [1–4(–5) in Py. avenicola vs. 2–9 in Py. chaetomioides]. The sexual morph has only been
observed in Py. chaetomioides.
Pyrenophora cynosuri Y. Marín & Crous, sp. nov. MycoBank MB829615. Fig. 53.
Etymology: Name refers to the host genus Cynosurus from which it was isolated.
Hyphae hyaline to pale brown, branched, septate, smooth-walled or verrucose, 1.5–5 m.
Conidiophores arising in groups, septate, straight or flexuous, usually geniculate in upper part, size of
cells rarely decrease towards apex, rarely branched, cells walls thicker than those of vegetative
hyphae, macronematous, rarely micronematous, pale brown to brown, slightly paler towards apex, not
swollen at the base, (70–)95–700 × 4.5–8 m. Conidiogenous cells smooth-walled to slightly
verruculose, terminal or intercalary, proliferating sympodially, pale brown to brown, subcylindrical to
swollen, 10–30(–37) × 5.5–10 m. Conidia verruculose, mostly curved, middle cells sometimes
enlarged, cylindrical to obclavate, tapering towards apex, subhyaline to pale brown, end cells rarely
paler, 2–5-distoseptate, (25–)28–80(–83) × 9–16.5 m, forming secondary conidiophores or conidia;
hila protuberant, darkened, thickened, (3–)3.5–6 m. Chlamydospores, microconidiation and sexual
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morph not observed.
Culture characteristics: Colonies on MEA reaching 55–60 mm diam after 1 wk at 25 °C, with
moderate aerial mycelium, flat to umbonate, margins arachnoid; surface greyish sepia to mouse grey;
reverse blood colour, margins luteous. Colonies on PDA reaching 60–65 mm diam, with sparse aerial
mycelium, flat, margins arachnoid; surface greyish sepia to fuscous black; reverse fuscous black.
Colonies on OA covering dish, with moderate aerial mycelium; surface greyish sepia to fuscous black,
margins brick; reverse fuscous black, margins brick.
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Typus: New Zealand, on seeds of Cynosurus cristatus (Poaceae), 1975, E.H.C. McKenzie (holotype
CBS H-23841, culture ex-type CBS 127918 = BRIP 12355 a = NZ 14880).
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Notes: Pyrenophora cynosuri was isolated from seeds of Cynosurus cristatum, a member of the
Poaceae, which includes host genera commonly infected by species of Pyrenophora. However,
Cynosurus represents a new host genus for Pyrenophora.
Pyrenophora cynosuri is closely related and morphologically similar to Py. dictyoides. However,
both species differ in the size of their conidiophore length (up to 700 m in Py. cynosuri vs. up to 250
m in Py. dictyoides) and conidial length (up to 83 m in Py. cynosuri vs. up to 250 m in Py.
dictyoides), as well as conidial septation (up to 5 in Py. cynosuri vs. up to 15 in Py. dictyoides).
Pyrenophora dictyoides A.R. Paul & Parbery, Trans. Brit. Mycol. Soc. 51: 708. 1968.
Synonyms: Helminthosporium dictyoides Drechsler, J. Agric. Res. 24: 679. 1923.
Helminthosporium dictyoides var. dictyoides Drechsler, J. Agric. Res. 24: 679. 1923.
Helminthosporium dictyoides f. dictyoides Drechsler, J. Agric. Res. 24: 679. 1923.
Drechslera dictyoides (Drechsler) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Helminthosporium dictyoides f. perenne Braverman & J.H. Graham, Phytopathology 50: 695. 1960.
Drechslera andersenii Scharif, Studies on Graminicolous Species of Helminthosporium (Tehran): 29.
1963. (nom. inval., Art. 36.1).
Drechslera andersenii A. Lam, Trans. Brit. Mycol. Soc. 85: 601. 1986.
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Notes: Isolates belonging to Py. dictyoides (CBS 127933 and DAOM 63666) and Drechslera andersenii
(CBS 258.80 and CBS 967.87) clustered together in a well-supported clade (100 % BS / 1 PP). Moreover,
the morphology of the asexual morph is similar in both species, differing only in the production of
conidia with much less tapered apices in D. andersenii (Sivanesan 1987). Therefore, we reduce these
species to synonymy.
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Pyrenophora nisikadoi Y. Marín & Crous, nom. nov. MycoBank MB829616. Fig. 54.
Replaced synonym: Helminthosporium brizae Y. Nisik., Ber. Ohara Inst. Landw. Biol.: 121, 133.
1928, non Pyrenophora brizae C. Massal. ex Sacc. 1911.
Additional synonyms: Bipolaris brizae (Y. Nisik.) Shoemaker, Canad. J. Bot. 37: 882. 1959.
Drechslera brizae (Y. Nisik.) Subram. & B.L. Jain, Curr. Sci. 35: 354. 1966.
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Etymology: Named after the Japanese plant pathologist and mycologist, Y. Nisikado, who first
described and named this fungus.
Hyphae hyaline to pale brown, branched, septate, verrucose, 2–5.5 m. Conidiophores arising in
groups, septate, mostly flexuous, rarely straight, geniculate at upper part, sometimes size of cells
decrease towards apex, frequently branched, cells walls thicker than those of vegetative hyphae,
macronematous, pale brown to brown, sometimes paler towards apex, rarely swollen at the base, 50–
330 × 3.5–6.5(–8.5) m. Conidiogenous cells smooth-walled to verruculose, terminal or intercalary,
proliferating sympodially, pale brown to brown, subcylindrical to swollen, 7.5–20.5(–25) × 5–7.5 m.
Conidia verruculose, straight or curved, middle cells enlarged, cylindrical to obclavate, tapering
towards apex, pale brown to brown, basal cell sometimes paler, less frequently apical cell also paler,
(1–)2–4(–5)-distoseptate, (15–)17.5–42.5 × 8.5–12 m, not forming secondary conidiophores or
conidia; hila flat, darkened, thickened, 2–4 m. Chlamydospores immersed in all media tested (MEA,
OA and PDA), brown to dark brown, lineally or irregularly disposed, verrucose, globose to
subglobose, up to 30 m. Microconidiation and sexual morph not observed.
Culture characteristics: Colonies on MEA reaching 80–90 mm diam after 1 wk at 25 °C, with
abundant aerial mycelium, cottony, lobate; surface pale greenish grey to greenish grey; reverse
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olivaceous black. Colonies on PDA covering dish, with moderate aerial mycelium, cottony, powdery
at margins, flat; surface greenish grey to olivaceous black; reverse olivaceous black. Colonies on OA
covering the dish, with sparse aerial mycelium, powdery to granular, flat; surface grey olivaceous to
olivaceous black; reverse leaden grey to leaden black.
Typus: Japan, from Briza minor (Poaceae), Y. Nisikado [epitype designated by Manamgoda et al.
(2014) CBS H-7218, culture ex-epitype CBS 190.29 = MUCL 9613].
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Additional materials examined: South Africa, Western Cape Province, J.S. Marais Nature Reserve, from Protea burchellii
senescent flowerheads (Proteaceae), 6 Jun. 2000, S. Lee, CBS 119213. New Zealand, Auckland, Waitakere Ranges, from
Briza minor (Poaceae), 1 Nov. 1975, E.H.C. McKenzie, CBS 127912 = ICMP 6183.
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Notes: This species was originally described in Helminthosporium as Hel. brizae (Nisikado 1928),
then transferred to Bipolaris (Shoemaker 1959), and finally placed in Drechslera (Subramanian &
Jain 1966). Type material was not available, thus Manamgoda et al. (2014) designated CBS 190.29 as
ex-epitype since this strain was isolated by the original author from the same host and location. In our
phylogenetic analysis based on ITS, gapdh and rpb2, CBS 190.29 together with other two strains were
located in a well-supported clade (100 % BS / 1 PP) within the main clade representing the genus
Pyrenophora. Therefore, we propose to transfer this species to Pyrenophora, changing the epithet to
Py. nisikadoi since Py. brizae already exists.
Isolate CBS 119213 sporulated, enabling us to conduct a morphological comparison and provide a
modern description of this species in PDA. Moreover, Pyrenophora nisikadoi was formerly only
recorded from Briza minor, a member of the Poaceae, while it is here also recorded from Protea
birchellii, which belongs to the Proteaceae.
Pyrenophora novozelandica Y. Marín & Crous, sp. nov. MycoBank MB829620. Fig. 55.
Etymology: Name refers to New Zealand, the country from where it was isolated.
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Hyphae hyaline to pale brown, branched, septate, verrucose, 1–6(–8.5) m. Conidiophores arising in
groups, septate, straight or flexuous, sometimes geniculate at upper part, size of cells not decreasing
towards apex, rarely branched, cells walls thicker than those of vegetative hyphae, macronematous,
pale brown to brown, paler towards apex, not swollen at the base, 35–700 × 4.5–7.5 m.
Conidiogenous cells smooth-walled, terminal or intercalary, proliferating sympodially, brown,
terminal conidiogenous cells hyaline, cylindrical to subcylindrical, 11–22 × 5–8.5 m. Conidia
smooth-walled, straight, rarely slightly curved, sometimes middle cells slightly enlarged, cylindrical
to obclavate, tapering towards apex, pale brown to brown, sometimes basal cell slightly paler, (2–)3–
5(–6)-distoseptate, 20.5–58 × 9.5–14 m, not forming secondary conidiophores or conidia; hila
usually inconspicuous, flat, slightly darkened, slightly thickened, 2–4 m. Chlamydospores,
microconidiation and sexual morph not observed.
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Culture characteristics: Colonies on MEA reaching 27–30 mm diam after 1 wk at 25 °C, with sparse
aerial mycelium, raised, margins fringed; surface grey olivaceous; reverse dark mouse grey, margins
buff. Colonies on PDA reaching 38–40 mm diam, with sparse aerial mycelium, flat, margins fringed;
surface smoke grey to olivaceous grey; reverse olivaceous black. Colonies on OA reaching 37–40 mm
diam, with moderate aerial mycelium, flat, margins fringed; surface grey olivaceous; reverse
olivaceous black.
Typus: New Zealand, Wanganui, Palmerston North, Seed Testing Station, on seed of Triticum sp.
(Poaceae), 5 Oct. 1976, G.F. Laundon (holotype CBS H-23843, culture ex-type CBS 127934 = LEV
11079b = PDD 50697).
Notes: Pyrenophora novozelandica is similar and closely related to Py. fugax. However, both species
can be easily distinguished based on the size of their conidiophores (up to 250 m in Py. fugax vs. up
to 700 m in Py. novozelandica) and conidia (50–170 × 14–24 m in Py. fugax vs. 20.5–58 × 9.5–14
m in Py. novozelandica), as well as conidial septation [4–8(–10) in Py. fugax vs. (2–)3–5(–6) in Py.
novozelandica]. Pyrenophora novozelandica is known to occur on Triticum in New Zealand, which is
a common host of species belonging to Pyrenophora, including Py. fugax.
Pyrenophora poae (Baudyš) Y. Marín & Crous, comb. nov. MycoBank MB829617.
Basionym: Helminthosporium poae Baudyš, Lotos 63: 104. 1916.
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Synonyms: Helminthosporium vagans Drechsler, J. Agric. Res., Washington 24: 688. 1923.
Drechslera vagans (Drechsler) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Drechslera poae (Baudyš) Shoemaker, Canad. J. Bot. 40: 827. 1962.
Description and illustration: Sivanesan (1987).
Materials examined: Canada, Saskatchewan, Saskatoon, from Poa pratensis (Poaceae), Oct. 1973, J.D. Smith, DAOMC
145373. Germany, Husum, from P. pratensis, Aug. 1966, U.G. Schlösser, CBS 319.68. USA, Maryland, Beltsville, from P.
pratensis, Apr. 1979, A. Hagan, CBS 128045 = BRIP 12969a.
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Notes: Pyrenophora poae was introduced as Helminthosporium poae by Baudys (1916), then
transferred to Drechslera (Shoemaker 1959). The original description was based on a specimen
isolated from Poa trivialis in the Czech Republic. Type material is not available, but CBS 319.68 is
considered here as an authentic strain since was isolated from the same host genus and continent.
Unfortunately, it did not sporulate and thus we chose to not designate it as epitype. All the strains
identified as D. poae were located in the main clade belonging to Pyrenophora, and a new
combination is proposed here.
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Pyrenophora pseudoerythrospila Y. Marín & Crous, sp. nov. MycoBank MB829675. Fig. 56.
Etymology: Named after its close phylogenetic relation to Py. erythrospila.
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Only forming protoascomata in OA. Protoascomata composed of pale brown to brown cells of up to
100 m diam, of textura angularis to textura globulosa. Pyrenophora pseudoerythrospila differs from
its closest phylogenetic neighbour, Py. erythrospila by unique fixed alleles in three loci based on
alignments of the separate loci deposited in TreeBASE (S23834): LSU positions 70 (G), 395 (C), 396
(T), 397 (T), 500 (G), 536 (C), 537 (T); ITS positions 141 (A), 146 (A), 147 (T), 148 (A), 149 (G),
152 (G), 153 (A), 154 (G), 155 (T), 164 (T), 168 (T), 169 (G), 173 (G), 174 (C), 176 (A), 178 (T), 179
(G), 184 (T), 187 (T), 189 (T), 197 (C), 198 (C), 199 (C), 205 (C), 206 (T), 208 (C), 223 (T), 224 (T),
225 (T), 226 (T), 234 (A), 252 (T), 264 (T), 268 (C), 279 (G), 288 (C), 289 (A), 293 (A), 302–304
(indels), 327 (T), 330–332 and 341 (indels),511 (T), 536–540 (indels), 558 (C), 591 (A), 594 (G), 601
(T), 621 (A), 630 (C), 641 (G), 643 (A), 644 (G), 645 and 646 (indels), 659 (T), 660 (G), 664 (T), 666
(A), 667 (T); gapdh positions 30 (C), 49 A 54 and 55 (indels), 59 (G), 60 (A), 69 (C), 71 (C), 74 (A),
85 (T), 87 (A), 177 (G), 178 (G), 179 (C), 180 (C), 182 (C), 187 (G), 189 (C), 190 (T), 191 (A), 192
(T), 193 (C), 194 (A), 195 (G), 196 (A), 197 (C), 201 (G), 203 (A), 204 (G), 205 (A), 256 (G), 319
(G), 373 (T), 406 (C), 487 (T), 493 (T), 496 (T), 523 (G), 526 (C), 532 (C), 538 (A).
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Culture characteristics: Colonies on MEA reaching 55–60 mm diam after 1 wk at 25 °C, with sparse
aerial mycelium, flat, margins fringed; surface pale vinaceous to vinaceous buff; reverse orange to
sienna, margins transparent. Colonies on PDA reaching 56–59 mm diam, with sparse aerial mycelium,
flat, margins fringed; surface cinnamon, white mycelium in the centre; reverse cinnamon, centre brick
to dark brick. Colonies on OA reaching 50–55 mm diam, without aerial mycelium, flat; surface
cinnamon; reverse transparent, centre brick to dark brick.
Typus: Germany, West Germany, on Lolium sp. (Poaceae), 9 Sep. 1968, U.G. Schlosser (holotype
CBS H-23844, culture ex-type CBS 127931 = DAOMC 126772).
Notes: The ex-type strain of Pyrenophora pseudoerythrospila did not sporulate on any of the media
tested, producing only few protoascomata in OA. However, these remained sterile after several
months of incubation. Pyrenophora pseudoerythrospila is closely related to Py. erythrospila, which
produces both sexual and asexual morphs. The protoascomata were also reported in Py. erythrospila,
but these finally developed mature ascospores after 25 wk. Pyrenophora erythrospila is commonly
found on Agrostis spp. in Australia and North America, but has also been reported on Lolium in
Germany (Farr & Rossman 2019), having the same host and distribution as Py. pseudoerythrospila.
Pyrenophora sieglingiae Y. Marín & Crous, sp. nov. MycoBank MB829618. Fig. 57.
Etymology: Name refers to Sieglingia, the host genus from which this fungus was collected.
Hyphae hyaline to pale brown, branched, septate, verrucose, (1–)1.5–5.5 m. Sterile ascomata solitary
or arising in groups, brown to dark brown or black, sometimes apical part of neck yellowish brown,
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composed of cells of textura intricata, up to 1200 m long, up to 300 m wide, conidiophores arising
from the body and neck; inside consisting of angular to globose, hyaline cells. Conidiophores arising
in groups, septate, straight or flexuous, rarely geniculate in the upper part, cell size rarely decreases
towards the apex, unbranched, cell walls thicker than those of vegetative hyphae, macronematous,
brown, mostly paler towards apex, not swollen at the base, 100–700 × (5–)7–9(–11) m.
Conidiogenous cells verruculose, terminal or intercalary, proliferating sympodially, pale brown to
brown, subcylindrical to slightly swollen, (12–)15–33.5(–36.5) × 9–12(–13.5) m. Conidia
verruculose, straight or curved, sometimes with middle cells enlarged, cylindrical to obclavate,
tapering towards apex, pale brown to brown, end cells usually paler, 4–8-distoseptate, 56–108(–120) ×
15–23(–25.5) m, forming secondary conidiophores or conidia; hila not protuberant or flat, darkened,
thickened, (4–)4.5–6.5(–7) m. Chlamydospores, microconidiation and sexual morph not observed.
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Culture characteristics: Colonies on MEA reaching 26–29 diam after 1 wk at 25 °C, with abundant
aerial mycelium, raised, slightly lobate; surface white to buff; reverse fuscous black. Colonies on PDA
reaching 27–30 diam, with abundant aerial mycelium, lobate; surface olivaceous black with patches of
white due to aerial mycelium; reverse olivaceous black, margins luteous. Colonies on OA reaching
54–57 diam, with moderate to abundant aerial mycelium, flat; surface smoke-grey to olivaceous grey;
reverse smoke-grey to olivaceous grey.
Typus: New Zealand, Auckland, Waikumete, from leaf of Sieglingia decumbens (Poaceae), E.H.C.
McKenzie (holotype CBS H-23842, culture ex-type CBS 127930 = ICMP 6170 = PDDCC 6170).
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Notes: Pyrenophora sieglingiae is closely related to Py. semeniperda and Py. wirreganensis.
Morphologically, these species are similar, producing sterile ascomata with long necks. However, they
can be distinguished by the size of their conidiophores (up to 700 m in Py. sieglingiae vs. up to 180
m in Py. semeniperda vs. up to 1000 m in Py. wirreganensis) and conidia [56–108(–120) × 15–23(–
25.5) m in Py. sieglingiae vs. 70–160 × 13–17 m in Py. semeniperda vs. (30–)40–80(–100) × (10–
)12–19(–22) m in Py. wirreganensis). Moreover, Py. semeniperda produces conidia with more septa
(up to 12 in Py. semeniperda vs. up to 8 in Py. sieglingiae vs. up to 9 in Py. wirreganensis), and it is
the only one that produces a sexual morph. Pyrenophora sieglingiae has been isolated from Sieglingia
from New Zealand while Py. wirreganensis occurs on Hordeum in Australia. Pyrenophora
semeniperda has been isolated from both hosts in both locations, apart from other hosts that are
widely distributed, i.e. Agropyron, Avena, Bromus, Cortaderia, Ehrhartia, Pennisetum and Triticum.
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Pyrenophora teres Drechsler, J. Agric. Res., Washington 24: 656. 1923.
Synonyms: Helminthosporium secalis Fée, Mém. Soc. Mus. Hist. Nat. Strassbourg 3: 36. 1843.
Alternaria secalis (Fée) Sacc. & Traverso, Syll. fung. (Abellini) 20: 1184. 1911.
Helminthosporium gramineum Rabenh., Klotzschii Herb. Viv. Mycol., Edn Nov, Ser. Sec., Cent. 4:
no. 332. 1857.
Brachysporium gracile var. gramineum (Rabenh. ex Schltdl.) Sacc., Syll. fung. (Abellini) 4: 430.
1886.
Drechslera graminea (Rabenh. ex Schltdl.) S. Ito, Proc. Imper. Acad. Tokyo 6: 355. 1930.
Drechslera teres subsp. graminea (Rabenh. ex Schltdl.) Simay, Barley Newsletter 36: 174. 1992.
Helminthosporium teres Sacc., Syll. fung. (Abellini) 4: 412. 1886.
Drechslera teres (Sacc.) Shoemaker, Can. J. Bot. 37: 881. 1959.
Drechslera teres f. teres (Sacc.) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Drechslera teres subsp. teres (Sacc.) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Helminthosporium hordei Eidam, Der Landw. (Schles. Landw. Ztg), Breslau 27: 509. 1891.
Helminthosporium tuberosum G.F. Atk., Bulletin of Cornell University 3: 47. 1897.
Drechslera tuberosa (G.F. Atk.) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Helminthosporium fragosoi Bubák, Hedwigia 57: 13. 1915.
Pyrenophora teres f. teres Drechsler, J. Agric. Res., Washington 24: 656. 1923.
Pyrenophora teres subsp. teres Drechsler, J. Agric. Res., Washington 24: 656. 1923.
Helminthosporium japonicum S. Ito & Kurib., Proc. Imper. Acad. Tokyo 6: 353. 1930.
Pyrenophora japonica S. Ito & Kurib., Proc. Imper. Acad. Tokyo 6: 353. 1930.
Drechslera japonica (S. Ito & Kurib.) Shoemaker, Canad. J. Bot. 37: 881. 1959.
Pyrenophora graminea S. Ito & Kurib., Proc. Imp. Acad. Japan 6: 353. 1930.
Pyrenophora teres subsp. graminea (S. Ito & Kurib.) Simay, Barley Newsletter 36: 174. 1992.
Pyrenophora secalis M.D. Whitehead & J. Dicks., Mycologia 44: 752. 1952.
Drechslera teres f. maculata Smed.-Pet., Arb. Tiflis Bot. Gard.: 139. 1971.
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Pyrenophora teres f. maculata Smed.-Pet., The Royal Veterinary and Agricultural University
Yearbook: 139. 1971.
Description and illustrations: Sivanesan (1987).
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Materials examined: Denmark, from Hordeum vulgare (Poaceae), unknown date, V. Smedegaard-Petersen (ex-type culture of
Py. teres f. maculata CBS 228.76). Japan, from H. vulgare (Poaceae), unknown date, S. Ito, CBS 281.31; unkown substrate
and date, S. Ito, CBS 282.31; from H. vulgare (Poaceae), unknown date, Y. Nisikado, CBS 336.29 = MUCL 9687. Germany,
Niedersachsen, Rotenburg, from H. vulgare (Poaceae), Jul. 1968, U.G. Schlösser, CBS 314.69. Hungary, Eszteragpuszta, from
Hordeum vulgare leaf (Poaceae), unknown date, col. M. Csosz, dep. J. Bakonyi, CBS 123929; Taplanszentkereszt, from
Hordeum vulgare leaf (Poaceae), unknown date, col. A. Tomcsanyi, dep. J. Bakonyi, CBS 123932.
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Notes: In our phylogenetic analysis, isolates identified as Py. teres and Py. graminea were located in
the same well-supported clade (100 % BS / 1 PP), suggesting that these represent the same species. In
fact, Py. graminea has been recently considered as a subspecies of Py. teres (Simay 1992) since its
morphology is similar and both species share the same host, Hordeum. Therefore, we propose the
synonymy of both species under the name of Py. teres, which is well established and the most
commonly used name for this taxon. Moreover, an authentic strain of Py. japonica was also located in
this clade, supporting the synonymy of Py. japonica with Py. teres proposed by Crous et al. (1995),
which was based on their morphological, molecular and pathological similarity.
Pyrenophora teres produces net blotch on barley worldwide, causing cell death and feeding off the
nutrients released (Sivanesan 1987, Louw et al. 1995, Campbell et al. 1999, Ellwood et al. 2012).
Two different forms of Py. teres were recognised depending on the disease symptoms produced, i.e.
Py. teres f. teres producing the net form of net blotch, characterised by elongated lesions where
necrosis develops along leaf veins with occasional transverse striations, while Py. teres f. maculata
produces the spot form of net blotch, typified by more ovoid lesions, often surrounded by a chlorotic
zone (Campbell et al. 1999, Ellwood et al. 2012). However, both forms are considered the same
species, Py. teres. This disease becomes systemic in plants infected from seed (Sivanesan 1987).
Pyrenophora variabilis Hern.-Restr. & Y. Marín, sp. nov. MycoBank MB829619. Fig. 58.
Etymology: Name refers to the highly variable conidial morphology.
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Hyphae hyaline to brown, branched, septate, smooth-walled to verrucose, 2.5–7 m. Conidiophores
arising in groups, septate, straight or flexuous, sometimes geniculate in upper part, simple, cells walls
thicker than those of vegetative hyphae, semi- to macronematous, brown, not swollen at the base, up
to 321 m long, 5–10 m wide. Conidiogenous cells smooth-walled, terminal or intercalary,
proliferating sympodially, brown, subcylindrical, 18–27 × 8–10 m. Conidia smooth-walled, straight
to curved, cylindrical, subcylindrical, obclavate, obpyriform to subglobose, pale brown to brown, 1–3distoseptate, 20–75 × 13–19.5 m; hila flat, darkened, thickened, 4–7 m. Chlamydospores and sexual
morph not observed.
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Typus: Canada, British Columbia, Agassiz Research Station, from leaves of Poa trivialis (Poaceae),
Jul. 1972, J.D. Smith (holotype CBS H-23843, culture ex-type CBS 127920 = DAOMC 139513).
Notes: Pyrenophora variabilis was located on an independent branch far removed from the other
species in the genus. It can be easily distinguished from all the species of the genus by its highly
variable conidial morphology in size and shape, from cylindrical, subcylindrical or obclavate to
subglobose or obpyriform. It was isolated from Poa trivialis (Poaceae) leaves in Canada, a common
host of Pyrenophora spp.
Pyrenophora wirreganensis (Wallwork et al.) Y. Marín & Crous, comb. nov. MycoBank MB829621.
Basionym: Drechslera wirreganensis Wallwork et al., Mycol. Res. 96. 888. 1992.
Description and illustration: Wallwork et al. (1992).
Material examined: Australia, South Australia, from Hordeum sp. (Poaceae), unknown date, J. Bakonyi, CBS 109896.
Notes: Pyrenophora wirreganensis was introduced as Drechslera wirreganensis by Wallwork et al.
(1992) to accommodate a specimen isolated from Hordeum in Australia. In the phylogenetic analysis,
the strain CBS 109896 identified as Py. wirreganensis and isolated from the same host and location
than the ex-type strain (IMI 348323), was located in an independent branch within the main clade
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representing the genus Pyrenophora. Therefore, this species is here transferred to the latter genus. For
comparison with close species see notes of Py. sieglingiae.
Authors: Y. Marin-Felix, M. Hernández-Restrepo, P.W. Crous
Ramichloridium Stahel ex de Hoog, Stud. Mycol. 15: 59. 1977. Fig. 59.
Classification: Dothideomycetes, Dothideomycetidae, Capnodiales, Dissoconiaceae.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, rpb2, tef1. Table 13. Fig. 60.
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Type species: Ramichloridium apiculatum (J.H. Mill. et al.) de Hoog, basionym: Chloridium apiculatum
J.H. Mill. et al. Ex-type strain: CBS 156.59 = ATCC 13211 = IMI 100716 = JCM 6972 = MUCL
15753 = MUCL 7991 = QM 7716.
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Mycelium consisting of hyaline, subhyaline, pale brown, or brown, smooth-walled, septate, branched,
submerged and aerial hyphae. Conidiophores straight to gently curved, subcylindrical, unbranched,
smooth- and thick-walled, brown to dark brown, 0–3-septate, or sometimes reduced to intercalary
conidiogenous cells. Conidiogenous cells integrated, terminal, pale to medium brown, or goldenbrown, apical part subhyaline to pale brown, subcylindrical, rarely somewhat clavate, sometimes with
irregular, nodulose swellings along the length of the conidiogenous cells, tapering towards apex, with
sympodial proliferation, forming a rachis with slightly thickened and darkened, circular, somewhat
protruding scars. Conidia solitary, aseptate, pale brown, smooth-walled to finely verrucose, clavate or
oblong to ellipsoid, or obovate to obconical, apex obtuse or subobtuse, base truncate, with a
conspicuous, slightly darkened and thickened, not refractive hilum.
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Culture characteristics: Colonies after 1 mo at 25 °C in the dark flat, spreading, with sparse to moderate
aerial mycelium, margins smooth and even, or lobate and feathery. On SNA grey olivaceous, pale mouse
grey, or smoke-grey. On PDA olivaceous grey or smoke-grey; reverse olivaceous grey or iron-grey. On
OA olivaceous grey or iron-grey; reverse iron-grey, with pale luteous pigment diffusing into agar. On
MEA olivaceous green; reverse olivaceous black, often with a diffusing citron-yellow pigment.
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Optimal media and cultivation conditions: MEA, OA, PDA and SNA at 25 °C under near-ultraviolet
light.
Distribution: Africa, America and Asia.
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Hosts: Aloe sp. (Aloaceae), Cucumis sativus, Cucurbita maxima (Cucurbitaceae), Malus domestica,
Malus pumila and Pyrus pyrifolia (Rosaceae). Also isolated from soil.
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Disease symptoms: Sooty blotch and flyspeck diseases.
Notes: Ramichloridium was initially introduced by Stahel (1937), designating R. musae as type
species. However, the name was invalid due to the lack of Latin description or diagnosis.
Subsequently, de Hoog (1977) validated this genus to include species with erect, dark conidiophores
and predominantly aseptate conidia, designating as new type species R. apiculatum. In that study, 13
species were recognised in Ramichloridium, and subsequently more species were included. However,
several molecular studies demonstrated that some of them belonged to different genera, i.e.
Myrmecridium, Pachyramichloridium, Pleurothecium, Radulidium, Rhinocladiella and Zasmidium,
and were subsequently transferred (Arzanlou et al. 2007, Cheewangkoon et al. 2009, Videira et al.
2017). Some species still need to be molecularly studied to confirm their phylogenetic position. In the
present study, only five species have been demonstrated to belong to Ramichloridium, which is
characterised by aseptate, pale brown, smooth-walled to finely verrucose, clavate or oblong to
ellipsoid, or obovate to obconical conidia. Based on molecular data, R. indicum is proposed here as a
new genus, Globoramichloridium indicum, and R. ducassei as a new combination in Zasmidium (see
Zasmidium below). Moreover, R. apiculatum, R. cucurbitae and R. mali were located in a wellsupported clade (100 % BS / 1 PP) without phylogenetic distance. Unfortunately, the only loci
available and common in the three species are the ITS and LSU. Therefore, other loci should be
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sequenced to verify their status as separate species.
Species of Ramichloridium cause sooty blotch and flyspeck disease (SBFS) on members of the
family Rosaceae, which produces blemishes on the epicuticular wax layer and is regarded as an
economically serious disease (Wang et al. 2014). Ramichloridium cucurbitae and R. punctatum have
been found as SBFS pathogens only in the USA (Li et al. 2012), while R. apiculatum, R. luteum and
R. mali have been reported as causal agents of SBFS in China (Zhang et al. 2007, Li et al. 2012,
Wang et al. 2014).
Globoramichloridium Y. Marín & Crous, gen. nov. MycoBank MB829622.
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References: Arzanlou et al. 2007, Li et al. 2012 (morphology and phylogeny).
Etymology: Name reflects the characteristic globose conidia produced by this genus.
Illustration: Arzanlou et al. (2007).
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Mycelium consisting of submerged and aerial hyphae; submerged hyphae, smooth- and thin-walled,
hyaline, with thin septa; aerial hyphae coarsely verrucose, olivaceous green, rather thick-walled, with
thin septa. Conidiophores arising vertically from creeping hyphae at right angles, straight,
unbranched, thick-walled, smooth-walled, dark brown, with up to 10 thin septa, often with inflated
basal cells. Conidiogenous cells terminally integrated, smooth-walled, dark brown, sympodially
proliferating, rachis straight or flexuose, geniculate or nodose, subhyaline; scars thickened and
darkened, clustered at nodes. Microcyclic conidiation observed in culture. Conidia solitary, (0–)1septate, not constricted at the septum, subhyaline to pale brown, smooth-walled or coarsely verrucose,
rather thin-walled, broadly ellipsoidal to globose, with truncate base; hila conspicuous, slightly
darkened, not thickened.
Culture characteristics: Colonies on MEA reaching 35 mm diam after 2 wks at 24 °C. Colonies
velvety, rather compact, slightly elevated, with entire, smooth, whitish margins, dark olivaceous green
in the central part.
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Type species: Globoramichloridium indicum (Subram.) Y. Marín & Crous. Holotype: IMI 114625.
Representative strain: CBS 171.96.
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Notes: This genus is introduced to accommodate R. indicum, which differs from Ramichloridium spp.
by its broadly ellipsoidal to globose, mostly 1-septate, smooth-walled or coarsely verrucose conidia,
being clavate or oblong to ellipsoid, or obovate to obconical, aseptate, smooth-walled to finely
verrucose in Ramichloridium. This genus is related to Dissoconium, but the latter can be easily
distinguished by its percurrent and sympodial proliferation, and the ellipsoid to obclavate, smoothwalled conidia.
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Globoramichloridium indicum (Subram.) Y. Marín & Crous, comb. nov. MycoBank MB829623.
Basionym: Chloridium indicum Subram., Proc. Indian Acad. Sci., Sect. B 42: 286. 1955.
Synonyms: Veronaea verrucosa Geeson, Trans. Brit. Mycol. Soc. 64: 349. 1975.
Veronaea indica (Subram.) M.B. Ellis, in Ellis, More Dematiaceous Hyphomycetes: 209. 1976.
Ramichloridium indicum (Subram.) de Hoog, Stud. Mycol. 15: 70. 1977.
Description and illustration: Arzanlou et al. (2007).
Material examined: Unknown collection details, Feb. 1996, L. Marvanová, CBS 171.96.
Notes: The strain examined and included in the phylogenetic analysis, CBS 171.96, was not derived
from type material. However, the morphology of this strain fits perfectly with the morphology of the
holotype IMI 114625 (de Hoog 1977). Therefore, CBS 171.96 is considered here as a representative
strain, and we propose the new genus and combination based on the phylogenetic data derived from
this isolate, as well as on the morphological differences observed.
Authors: Y. Marin-Felix, J.Z. Groenewald & P.W. Crous
Seifertia Partr. & Morgan-Jones, Mycotaxon 83: 348. 2002. Fig. 61.
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Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Melanommataceae.
Type species: Seifertia azaleae (Peck) Partr. & Morgan-Jones, basionym: Periconia azaleae Peck.
Representative strain: DAOM 239136.
DNA barcode (genus): LSU.
DNA barcodes (species): LSU, tef1. Table 14.
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Mycelium superficial or immersed, composed of branched, septate, smooth- and thin-walled, pale
white to white or brown hyphae. Synnemata erect, simple, unbranched or very rarely branched, greenish
when young, becoming dark brown to black at maturity, capitate at the apex. Conidiophores
macronematous, synnematous, straight or slightly flexuous, unbranched or branched toward the upper
region, cylindrical, septate, smooth- and thin-walled, hyaline to pale brown or olivaceous brown to
brown. Conidiogenous cells mono- or polyblastic, integrated, terminal, determinate, cylindrical to
subclavate or doliiform, smooth-walled. Conidia acropleurogenous, holoblastic, in simple or branched
acropetal chains, dry, sometimes aggregated into slimy masses at the apex of the synnema, aseptate or
very rarely 1-septate, smooth- and thin-walled, oblong, ellipsoidal, subglobose or fusiform, hyaline to
subhyaline, pale brown or olivaceous (adapted from Partridge & Morgan-Jones 2002).
Culture characteristics: Colonies effuse, powdery or cottony to fairy fluffy, grey to dark brown.
Optimal media and cultivation conditions: MEA, OA and PDA at 25 °C.
Distribution: North America, Europe and Asia.
Hosts: Species of Rhododendron.
Disease symptoms: Bud and twig blight; Rhododendron bud blight disease.
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Notes: Seifertia was introduced by Partridge & Morgan-Jones (2002) to accommodate Pycnostysanus
azaleae based on morphological differences. Seifertia azaleae is morphologically similar to Sorocybe
resinae. However, Partridge & Morgan-Jones (2002) decided to erect the new genus Seifertia since
Sei. azaleae produces much narrower conidia and has minute denticles on the conidiogenous cells.
Seifertia is characterised by erect, simple, and dark synnemata, macronematous conidiophores,
holoblastic, integrated, terminal and determinate conidiogenous cells, and unicellular or very rarely 1septate, pale brown or olivaceous conidia. This cosmopolitan genus occurs on azaleas and
rhododendrons causing a disease known as Rhododendron bud blight disease, in which the flower buds
die, and twig blight occurs. Infected buds are easily recognisable by the blackening of the bud and the
development of numerous synnemata which appear as tiny black spines over the entire surface
(Partridge & Morgan-Jones 2002, Glawe & Hummel 2006).
This genus, which is relatively poorly studied, was recently placed in Melanommataceae by Li et al.
(2016b), when they introduced the second species belonging to Seifertia, Sei. shangrilaensis. However,
the relation of Seifertia with Mycopappus and its synasexual morph Xenostigmina, which are foliar
pathogens belonging to Melanommataceae, was demonstrated previously by Crous et al. (2009a).
References: Partridge & Jones 2002 (morphology), Glawe & Hummel 2006 (pathogenicity), Seifert et al.
2007, Li et al. 2016b (morphology and phylogeny).
Authors: Y. Marin-Felix & P.W. Crous
Seiridium Nees, Das System der Pilze und Schwämme: 22. 1817. Fig. 62.
Synonym: Pestalotia De Not., Mém. Reale Accad. Sci. Torino 3: 80. 1841.
Additional synonyms in Bonthond et al. (2018).
Classification: Sordariomycetes, Xylariomycetidae, Xylariales, Sporocadaceae.
Type species: Seiridium marginatum Nees, Syst. Pilze (Würzburg): 23. 1817. Neotype designated by
Shoemaker et al. (1966): K 200376. Epitype and ex-epitype culture designated by Jaklitsch et al. (2016):
WU 33575, CBS 140403.
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DNA barcode (genus): ITS.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 15. Fig. 63.
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Ascomata perithecial, immersed to semi-erumpent, depressed, globose to pyriform, scattered or confluent,
with central ostiole; necks slightly papillate, black, periphysate; ascomatal wall dark brown,
pseudoparenchymatous. Paraphyses hyaline, smooth-walled, filiform. Asci cylindrical, 8-spored,
unitunicate, thin-walled, stipitate, with an apical amyloid ring. Ascospores cylindrical-oblong, euseptate,
septa often thicker than the wall, yellow- to dark brown, guttulate. Conidiomata acervuloid to pycnidioid,
semi-immersed to erumpent, uni- to plurilocular, brown or black, glabrous, dehiscing by irregular splits in
the upper wall. Conidiophores lining the cavity of the conidioma, septate and sparsely branched at the
base, or reduced to conidiogenous cells, hyaline, smooth-walled. Conidiogenous cells discrete, integrated,
ampulliform to lageniform or subcylindrical, hyaline, smooth-walled, proliferating percurrently at the
apex. Conidia fusiform, euseptate (septal pores present or not), end cells hyaline, median cells dark brown
to brown, wall thick, smooth or with striations, constricted at septa or not; apical cell with single or
multiple, unbranched or branched appendages; basal cell with or without a centric, unbranched or
sometimes branched appendage (adapted from Bonthond et al. 2018).
Culture characteristics: Colonies on PDA circular to irregular, reaching 12–68 mm diam after 14 d at 22
°C, mostly flat, in some cultures elevated at margins, often white coloured or with other colours,
sporulation rare, with pycnidioid conidiomata. On SNA circular to irregular to rhizoid, reaching 20–54
mm diam after 14 d at 22 °C, mostly flat, white coloured, with moderate to abundant aerial mycelium,
sporulation for some species within and others after 2 wk, sporodochia often compact and scattered.
Optimal media and cultivation conditions: Colonies grow well on CMA, MEA, PDA and SNA at 22 °C.
Most species sporulate on SNA and some species on CMA, MEA or PDA as well.
Distribution: Worldwide.
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Hosts: A diversity of gymnosperms and angiosperms. The genus is most well-known from members of
Cupressaceae.
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Disease symptoms: Cankers on stems and twigs.
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Notes: Pestalotia was introduced in 1841 and is similar to Seiridium (1817), one of the older names in the
Sporocadaceae. The genus has been subjected to many rearrangements (reviewed in Sutton 1969) which
eventually resulted in the genus accommodating only the type species: P. pezizoides. One of the important
characters used to separate Pestalotia from the related genera Pestalotiopsis and Truncatella was the
production of 5-septate (or 6-celled) conidia. However, this morphology is typical for Seiridium as well,
and for this reason it was speculated that Pestalotia and Seiridium could be congeneric
(Maharachchikumbura et al. 2014). A fresh collection of P. pezizoides from Vitis vinifera collected in the
USA was recently obtained (Table 1), which matches in all respects with the type of P. pezizoides. DNA
sequence data generated here confirm that P. pezizoides clusters within Seiridium (Fig. 63). Therefore,
Pestalotia is reduced to synonymy with Seiridium. For Seiridium, the here synonymized Pestalotia and
related genera, not only the number but also the type of conidial septation has been a commonly reported
descriptor. Different authors have interpreted the appearance of the cell walls as either distoseptate (e.g.
Nag Raj 1993) or euseptate (e.g. Sutton 1980). Although when examined by light microscopy, conidia
can appear as distoseptate, electron microscopic studies on S. cupressi (Roberts & Swart 1980) and S.
pezizoides (Griffiths & Swart 1974) have shown that the conidial cell walls are differentiated into
multiple zones but arise from a single layered cell wall and are thus euseptate. Since the sexual morph is
known for only a few species, the taxonomy in Seiridium has been based mainly on asexual morphology.
However, the generic type (S. marginatum) is one of the exceptions where both morphs have been
characterised. This species was re-described and epitypified by Jaklitsch et al. (2016), who also provided
detailed illustrations of sexual and asexual morphology.
References: Nag Raj 1993 (morphology), Danti & Della Roca 2017 (pathogenicity), Bonthond et al.
2018 (morphology and phylogeny),
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Seiridium cupressi (Nattrass et al.) Bonthond, Sandoval-Denis & Crous, comb. nov. MycoBank
MB830554.
Basionym: Rhynchosphaeria cupressi Nattrass et al., Trans. Brit. Mycol. Soc. 46: 103. 1963.
Synonyms: Cryptostictis cupressi Guba, Monograph of Monochaetia and Pestalotia: 47. 1961. Nom.
inval. Art. 40.3 (Shenzhen).
Lepteutypa cupressi (Nattrass et al.) H.J. Swart, Trans. Brit. Mycol. Soc. 61: 79. 1973.
Seiridium cupressi (Guba) Boesew, Trans. Brit. Mycol. Soc. 80: 545. 1983. Nom. inval. Art. 40.3
(Shenzhen).
Known distribution: Africa (Kenya, Uganda) and Europe (Greece).
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Description: Sexual morph adapted from Nattrass et al. (1963). Asexual morph adapted from Bonthond et
al. (2018).
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Typus: Africa, Kenya, on Cupressus macrocarpa, July 1954, R.M. Nattrass (holotype of
Rhynchosphaeria cupressi IMI 56917); from cankers in branches of Cupressus macrocarpa, 1949, D.R.
Jones [epitype of Rhynchosphaeria cupressi designated here IMI 52254, MBT386544 (dried culture),
culture ex-epitype CBS 224.55].
Additional materials examined: Europe, Greece, from Cupressus sp., A. Graniti, CBS 122616 = CMW 1646. Africa, Kenya, nonpathogenic isolate from Cupressus sp., collection data unknown (CBS 320.51); on Cupressus macrocarpa, July 1948, R.M.
Nattrass, IMI 37158; on Cupressus macrocarpa, Dec. 1949, R.M. Nattrass, IMI 40096; from cankers in branches of Cupressus
forbesii, 1949, D.R. Jones, IMI 52255 (dried culture); CBS 225.55.
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Notes: This species has been a source of confusion since its introduction (Guba 1961). Bonthond et al.
(2018) showed that Guba’s diagnosis included three different species (S. cancrinum, S. cupressi and S.
kenyanum), and selected an epi- and lectotype for Cryptosticis cupressi; the latter name, however, was
invalidly published (article 40.1). Nattrass et al. (1963) re-examined the material from Guba (1961) and
synonomized Cryptostictis cupressi with Monochaetia unicornis. Despite noting small morphological
differences between the asexual stage of R. cupressi and the type of M. unicornis (i.e. smaller and slender
conidia), they were not able to confirm the exogenous origin of the basal appendage, which was the main
argument for Guba (1961) to place the species in Cryptostictis instead of Monochaetia. In the same study,
Nattrass et al. (1963) described the sexual morph of M. unicornis as Rhynchosphaeria cupressi based on
three specimens: IMI 37158, IMI 40096 and the holotype IMI 56917. Bonthond et al. (2018) examined
each of these specimens but incorrectly cited the holotype as IMI 37158. While only the sexual-morph
was found in these materials, the original description from Nattrass et al. (1963) includes drawings,
photographs and measurements of the conidia. These measurements (22–32 × 6–9.5 m) fall perfectly
within the range documented for the lineage currently assigned to S. cupressi (Guba) Boesew. (18–36 ×
5–11.5 m) (Bonthond et al. 2018). Consequently, being the oldest valid name for this lineage and in
accordance with the rule of priority Rynchosphaeria cupressi is recombined in Seiridium, as Seiridium
cupressi (Nattrass et al.) Bonthond, Sandoval-Denis & Crous, and an epitype is designated (CBS 224.55).
The similar species S. cancrinum and S. unicorne show smaller conidia (20–30.5 m and 19–28 m,
respectively), whereas conidia of S. kenyanum are considerably larger (24–39 m).
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Seiridium pezizoides (De Not.) Crous, comb. nov. MycoBank MB828021. Fig. 64.
Basionym: Pestalotia pezizoides De Not., Mém. Reale Accad. Sci. Torino 2, 3: 80. 1839.
Caulicolous. Isolated from branches of Vitis vinifera. On the host (described in more detail by Nag Raj
1993): Conidiomata irregularly scattered over the surface, gregarious to confluent, discoid to cupulate and
occasionally globose, erumpent from tissue, acervular to sporodochial, occasionally with aerial mycelium,
black to brown, (300–)350–500(–650) m. On SNA: Conidiophores tightly aggregated in the conidioma,
cylindrical, irregularly branched, hyaline or pale brown at the base, smooth- and thin-walled.
Conidiogenous cells discrete, hyaline, cylindrical, smooth- and thin-walled. Conidia lunate to falcate,
often curved, 5-septate, not striate, bearing a basal and two or more apical appendages, euseptate with
pores sometimes visible, (24–)28–33.5(–38.5) × (6–)7–8(–9) m, mean ± SD = 30.7 ± 2.8 × 7.5 ± 0.4 m;
basal cell obconic with truncate base, hyaline, smooth-walled, bearing marginal frills, 4–7 m; four
median cells pale brown, smooth-walled, cylindrical to doliiform; second cell from base 3.5–8 m; third
cell 3.5–7 m; fourth cell 3.5–6.5 m; fifth cell 3.5–7.5 m; apical cell conical, hyaline, smooth-walled,
4.5–8.5 m long; apical appendages single or multiple, centric, branched or unbranched, 8.5–27 m;
basal appendage single, cylindrical, centric, occasionally branched, 5.5–14 m.
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Culture characteristics: Colonies on PDA circular, reaching 34–37 mm diam after 10 d at 22 °C, flat,
olivaceous to luteous in the centre, white to brown at the margins, with abundant aerial mycelium at the
margins, not sporulating within 10 d. On CMA circular, reaching 39–41 mm diam after 10 d at 22 °C, flat
at centre and margins, dark brown to black, without aerial mycelium, not sporulating within 10 d. On
MEA circular to slightly irregular, reaching 25–27 mm diam after 10 d at 22 °C, flat, olivaceous to pale
green, with a white outer ring, with moderate aerial mycelium, massive spore production in the centre. On
SNA circular, reaching 31–33 mm diam after 10 d at 22 °C, raised in the centre, flat at the margins, with
moderate aerial mycelium, no sporulation within 10 d.
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Distribution: Europe (France, Italy), USA.
Typus: Italy, near Mailand, twig of Vitis vinifera (Vitaceae), leg. Oct. 1838, De Notaris, holotype, RO.
Additional material examined: USA, Virginia, Charlottesville, from a complex hybrid of Vitis æstivalis × Vitis cinerea × Vitis
vinifera, 2018, L. Morton, CBS 145115 = CPC 35011.
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Notes: After several rearrangements and the introduction of the genera Pestalotiopsis and Truncatella (see
Sutton 1969 and references therein), the generic type P. pezizoides was the only remaining species in
Pestalotia. A specimen of P. pezizoides (IMI 83642, from branches of Vitis vinifera, Italy, non-type) was
examined and redescribed by Sutton (1980). A more detailed description was provided by Nag Raj
(1993), based on several materials, including this specimen. The present study is the first to provide DNA
sequence data on this species. We sequenced four loci (ITS, rpb2, tef1 and tub2) and included P.
pezizoides in an updated phylogeny of Seiridium (Fig. 63) which supports the conclusion that Pestalotia
and Seiridium are congeneric. Consequently, the species is transferred to Seiridium as S. pezizoides. The
S. pezizoides strain (CBS 145115 = CPC 35011) that was included in this analysis morphologically
conforms with the description and was isolated from the same host, Vitis. However, since the specimen
was collected in Virginia (USA) and the holotype is from Italy, it is not suitable for epitypification. The
phylogeny generated here suggests that S. pezizoides is most closely related to S. rosarum (Rosa canina,
Italy).
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Seiridium unicorne (Cooke & Ellis) B. Sutton, Mycol. Pap. 138: 74. 1975. Fig. 65.
Basionym: Pestalotia unicornis Cooke & Ellis, Grevillea 7: 6. 1878, as "Pestalozzia".
Synonym: Monochaetia unicornis (Cooke & Ellis) Sacc. & D. Sacc., Syll. Fung. 18: 485. 1906.
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Caulicolous. Most commonly isolated from cankers on branches of species from Cupressaceae.
Conidiomata on PDA numerous, sporodochial, globose or clavate, mostly solitary, erumpent from agar,
partially immersed in mycelium, producing large black spore masses; on SNA, sporodochial, mostly
aggregated, erumpent from agar, producing large black spore masses. On SNA: Conidiophores septate,
cylindrical, irregularly branched, hyaline or brown, thin-walled, 22–68 m long, ex-epitype: 22–50 m
long. Conidiogenous cells discrete, hyaline, cylindrical, smooth- and thin-walled, 3.5–29.5 × 1.5–3.5 m,
ex-epitype: 16.2–28.9 × 1.7–3.5 m, proliferating percurrently, with visible collarettes and minute
periclinal thickenings. Conidia lunate to falcate, curved, 5-septate, rarely 4- or 6-septate, not striate,
bearing two appendages, euseptate with no visible pores, (19–)22.5–26.5(–28) × (6.5–)7.5–8.5(–9.5) m,
mean ± SD = 24.5 ± 1.8 × 7.9 ± 0.5 m, ex-epitype: (19–)22.5–26.5(–28) × (6.5–)7.5–8.5(–9) m, mean
± SD = 24.4 ± 1.9 × 7.8 ± 0.4 m; basal cell obconic with a truncate base, hyaline, walls smooth, bearing
minute marginal frills, 2.5–9.5 m, ex-epitype: 2.5–6 m (n = 119); four median cells colour varying
from pale to dark brown, smooth-walled, cylindrical to doliiform; second cell from base 3.5–6 m (n =
152); third cell 3–5.5 m; fourth cell 3–5.5 m; fifth cell 3–5.5 m, ex-epitype: second cell from base
3.5–6 m; third cell 3–5.5 m; fourth cell 3–5 m; fifth cell 3–5.5 m; apical cell conical, hyaline,
smooth-walled, 2–5.5 m long, ex-epitype: 2–5.5 m long; apical appendage single, mostly centric, 5–10
m, ex-epitype: 5–9.5 m; basal appendage single, cylindrical, mostly excentric, 2.7–7.1 m, ex-epitype:
4–6.5 m (adapted from Bonthond et al. 2018).
Culture characteristics: Colonies on PDA irregular, reaching 65–68 mm diam after 14 d at 22 °C, slightly
umbonate, colour citrine, olivaceous buff to olivaceous, with compact aerial mycelium on the surface,
abundant sporulation surrounding centre and at the margins of the colony. On CMA circular or irregular,
reaching 58–59 mm diam after 14 d at 22 °C, flat at the centre and margins, citrine to olivaceous
coloured, with moderate aerial mycelium on the surface, sporulating abundantly. On MEA irregular,
reaching 35–40 mm diam after 14 d at 22 °C, flat to crateriform, slightly sunk into the agar, buff to
olivaceous coloured at the centre becoming white at the margins, with dense mycelium on the surface,
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sporulating near the centre. On SNA circular to slightly irregular, reaching 20–21 mm diam after 14 d at
22 °C, umbonate, with moderate aerial mycelium, sporulation abundant between centre and margins.
Distribution: New Zealand, South Africa and USA.
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Typus: USA, Maryland, Pocomoke City, 38.072952N, 75.555852W, from branch canker of Cupressus
sp., 2017, S.A. Krueger-Hadfield (epitype of Pestalotia unicornis designated here CBS H-23739,
MBT383596, cultures ex-epitype CBS 143871 = CPC 34650, CBS 143872 = CPC 34649, CBS 143873 =
CPC 34651); New Jersey, from Chamaecyparis thyoides, 1878, J.B. Ellis (holotype of Pestalotia
unicornis IMI 5816).
Additional materials examined: New Zealand, from Cryptomeria japonica, 1981, H.J. Boesewinkel (CBS H-23151 reference
specimen, culture CBS 538.82 = CPC 23783 = NBRC 32684). South Africa, from Cupressus sempervirens, 1999, I. Barnes
(culture CBS 120306 = CMW 5596).
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Notes: Seiridium unicorne (basionym: Pestalotia unicornis) is the earliest described cypress pathogen
(Cooke & Ellis 1878) of Seiridium and was isolated from “cedar wood” collected in New Jersey (USA).
The host was later identified by W.W. Wagener as Chamaecyparis thyoides, as indicated on the holotype
label (see Bonthond et al. 2018). The genus Seiridium accommodates multiple species infecting
Cupressaceae and S. unicorne has traditionally been regarded as a mild pathogen but capable of infecting
a broad range of hosts, including plant species beyond the Cupressaceae (Guba 1961). The holotype (IMI
5816) was obtained and examined during a preceding study (Bonthond et al. 2018) and found to be
limited to two microscope slides. Therefore, the selection of an epitype for S. unicorne is important to
consolidate a stable taxonomic concept for this taxon. Given the occurrence of related species which are
pathogenic on the same hosts (i.e. S. cancrinum, S. cardinale, S. cupressi and S. neocupressi) the
availability of ex-epitype DNA sequence data provides a valuable reference for the identification of future
collections. The specimen we introduce here as epitype (CBS H-23739) was collected from necrotic
lesions of a Cupressus sp. in Maryland, USA and matches morphologically in all respects with the
holotype (IMI 5816) and the reference strain (CBS 538.82) of S. unicorne. Furthermore, in the four-locus
phylogeny (Fig. 63E) the ex-epitype strain clusters under a fully supported node in the clade that was
assigned to S. unicorne based on morphology (Bonthond et al. 2018). Conidial measurements strongly
overlap between the selected epitype, holotype and reference strain (Fig. 63A–D), although the median
width of the epitype being slightly narrower in comparison to the holotype and reference strain.
Measurements of basal and apical appendages and distributions of those measurements are highly similar
between epitype and reference strain. For both appendages, the variation in measurements is higher for
the holotype compared to the reference strain and the epitype, which, however, likely results from the age
and condition of the material as we observed that conidial appendages from the holotype were often
damaged.
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Authors: G. Bonthond, M. Sandoval-Denis, S.A. Krueger-Hadfield, L. Morton, C. Ambers & P.W. Crous
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Septoriella Oudem., Ned. kruidk. Archf, ser. 2, 5: 504. 1889. Fig. 66.
Synonyms: Allophaeosphaeria Ariyaw. et al., Fungal Diversity 72: 137. 2015.
Poaceicola W.J. Li et al., Mycosphere 6: 696. 2015.
Vagicola Chethana & K.D. Hyde, Fungal Diversity 75: 113. 2015.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
Type species: Septoriella phragmitis Oudem. Epitype and ex-epitype strain designated by Crous et al.
(2015a): CBS H-22281, CBS 140065.
DNA barcode (genus): LSU. Fig. 28.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 16. Fig. 67.
Ascomata ostiolate, solitary to gregarious, immersed to semi-immersed or superficial, broadly
ellipsoidal to globose, subglobose, or obpyriform, brown to dark brown or black, smooth-walled,
coriaceous, uni- to biloculate; necks central, flush to papillate, brown to dark brown or black, with or
without periphyses, rarely comprising short, hyaline setae; ascomatal wall thin-walled, outer layers
composed of brown to dark brown or blackish cells of textura angularis, inner layers composed of
brown cells of textura prismatica, or of hyaline or brown cells of textura angularis, rarely composed
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of hyaline gelatinous cells. Hamathecium composed of numerous, 1–3 m wide, filiform to broadly
cylindrical, septate, cellular pseudoparaphyses, or lacking pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, cylindrical, broadly cylindrical, cylindrical-fusiform, cylindrical-clavate,
clavate, or broadly clavate, pedicellate, apically rounded with an ocular chamber. Ascospores
overlapping, 1–3-seriate, hyaline to yellowish brown or pale brown, brown, golden brown, or reddish
brown, narrowly or broadly fusiform or oblong to narrowly oblong, straight or slightly curved,
smooth-walled or echinulate, multi-septate, with transverse septa, with or without longitudinal septa,
sometimes with enlarged medium cells, constricted or not at septa, conical or obtuse, rounded ends,
with or without sheath. Conidiomata pycnidial, solitary or aggregated, immersed to semi-immersed,
globose to subglobose, unilocular, pale brown to brown or dark brown, with central, papillate, circular
ostiole; conidiomatal wall of brown cells of textura angularis, rarely of textura oblita, inner layers
becoming hyaline. Conidiophores lining the inner cavity, reduced to conidiogenous cells, invested in
mucus. Conidiogenous cells ampulliform to lageniform, hyaline, smooth-walled, proliferating via
inconspicuous percurrent proliferations near apex. Conidia cylindrical to subcylindrical, fusiform, or
subfusiform, apex obtuse to subobtuse, base truncate, straight or curved, euseptate, pale brown to
brown, thin-walled, smooth-walled or minutely verruculose, bearing mucoid appendages at both ends
(type H sensu Nag Raj 1993) (description of asexual morph adapted from Crous et al. 2015a).
Culture characteristics: Colonies with moderate to abundant aerial mycelium, variable in colour,
circulate or lobate. On PDA surface white to iron-grey or grey olivaceous, greyish white, pale olivaceous
grey, or dull green margins, mouse grey in the middle and pale mouse grey at the center; reverse grey,
olivaceous grey, dull green, buff to dark brown, olivaceous to pale brown or black, brown or olivaceous
brown to black. On MEA surface white, dirty white, or white to pale yellow or iron-grey; reverse yellow,
umber, buff, or dark grey to black.
Optimal media and cultivation conditions: CMA, MEA, OA and PDA at 25 °C.
Distribution: Mostly Europe, but also reported in Asia and North America.
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Hosts: Mostly saprophytes of grasses (Poaceae), including Arundo spp., Agrostis stolonifera, Bromus
sterilis, Calamagrostis spp., Dactylis glomerata, Elymus glaucus, E. repens, Poa sp., Phragmites spp.,
Setaria verticillata. Also found on Rosa canina (Rosaceae) and Juncus sp. (Juncaceae), and others hosts
not molecularly corroborated. Septoriella hirta is considered and important secondary pathogen of
grasses, including Agropyron spp., Bromus spp., Dactylis glomerata, Festuca spp., Poa spp., Stipa spp.,
and Triticum spp., among others.
Disease symptoms: Secondary foot rot and rot of mature straw. Discoloured culms and predisposition of
the plant to premature collapse.
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Notes: Septoriella was considered an asexual genus characterised by pycnidial, unilocular conidiomata,
and cylindrical to fusoid, euseptate conidia bearing mucoid appendages at both ends (Crous et al. 2015a).
However, in our phylogenetic studies based on ITS and LSU, and on the combined dataset, the ex-type
strains of the sexual genera Allophaeosphaeria, Poaceicola and Vagicola were located in the clade
representing the genus Septoriella. Therefore, these genera are synonymised with Septoriella in the
present study.
Allosphaeosphaeria was recently introduced by Liu et al. (2015) to incorporate two new
saprophytic species found on Dactylis glomerata from Italy, i.e. Al. dactylidis and Al. muriformia, the
latter designated as type species. These species only produce the sexual morph characterised by
ascospores with transverse and longitudinal septa, and a gelatinous sheath. Subsequently, three other
new species were introduced in the genus, i.e. Al. clematidis, Al. cytisi and Al. subcylindrospora.
Allosphaeosphaeria clematidis and Al. cytisi only produce the sexual morph, while Al.
subcylindrospora only produces an asexual morph. The morphology of this asexual morph fits
perfectly in the description of Septoriella, corroborating the synonymy proposed in the present study
based on the phylogenetic data. Allosphaeosphaeria clematidis was recently excluded from the genus
and transferred to the new genus Embarria (Wanasinghe et al. 2018). Moreover, in our phylogenetic
studies, Al. cytisi formed an independent lineage in Phaeosphaeriaceae far from the clade
representing Septoriella. Therefore, a new genus is proposed to accommodate this species.
The genus Poaceicola was introduced by Li et al. (2015) to accommodate Phaeosphaeria elongata
and two new species, i.e. Po. arundinis and Po. bromi. The two latter species are characterised by the
production of an asexual morph similar to Septoriella. Poaceicola elongata produces a sexual morph
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characterised by ascospores with transverse septa. The presence of only transverse septa could be a
morphologic difference from Allophaeosphaeria. However, seven more new species have been
included in the genus, including one species presenting ascospores with transverse and longitudinal
septa, Po. arundinis, demonstrating that the longitudinal septation of the ascospores is not
phylogenetically informative in these genera.
Vagicola was recently introduced by Ariyawansa et al. (2015a), in the same year as the other two
genera. Ariyawansa et al. (2015a) raised the subgenus Vagicola (Shoemaker & Babcock 1989) to
generic rank to accommodate Phaeosphaeria vagans, a species characterised by a sexual morph
similar to the species of Poaceicola, having ascospores with transverse septa only. Subsequently,
Jayasiri et al. (2015) introduced two new species: V. chlamydospora, which presents both morphs, and
V. dactylidis, which produces only the sexual morph. The sexual morph of V. chlamydospora is
similar to the two former species of the genus, while V. dactylidis produces ascospores with transverse
and longitudinal septa as seen in species of Allophaeosphaeria, indicating again that the longitudinal
septation of ascospores is not phylogenetically informative. Vagicola chlamydospora was recently
transferred to Septoriella based on phylogenetic data (Jayasiri et al. 2015). Surprisingly, the asexual
morph reported in that species does not fit with the morphology of Septoriella, since it produces
micro- to macronematous conidiophores and chlamydospore-like conidia. Recently, Thambugala et al.
(2017) introduced the last species of the genus, V. arundinis, which produces both morphs and is
characterised by ascospores with transverse septa and an asexual morph similar to Septoriella, which
demonstrates the link of Vagicola with Septoriella. This last species was invalid because two
holotypes were designated. Therefore, this taxon is validated in the present study.
Moreover, in our phylogenetic analyses, the ex-type strain of the most recently described species
of Neostagonospora was located in the clade representing Septoriella. This species is characterised by
the production of conidia that are subcylindrical or fusiform, euseptate, with a subobtuse apex and
truncate base. However, the presence of mucoid appendages at both ends, as the other species of
Septoriella, has not been reported. Septoriella artemisiae is saprobic or weakly necrotrophic on dead
and dying stems of Artemisia austriaca.
Most of the species now included in Septoriella are saprophytes, except for Sep. hirta, which is an
important secondary pathogen of grasses (Sprague 1950). This species is often found in association
with other fungi such as Gaeumannomyces graminis (Johnston et al. 2014) and Oculimacula
yallundae causing foot rot of wheat (Crous et al. 2003, 2015a). Other disease symptoms observed in
plants affected by Sep. hirta are discoloured culms and predisposition to premature collapse,
especially in rainy and windy seasons, since this species produces a weakness in the culms of plants
with ripe grains. The result of all these symptoms resulted in the increasing of the cost of harvesting
and decreasing of the grain quality (Sprague 1950).
References: Sprague 1950 (pathogenicity), Crous et al. 2014b, Ariyawansa et al. 2015a, Li et al. 2015,
Liu et al. 2015, Thambugala et al. 2017 (morphology and phylogeny).
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Septoriella agrostina (Mapook et al.) Y. Marín & Crous, comb. nov. MycoBank MB829676.
Basionym: Poaceicola agrostina Mapook et al., Fungal Diversity 10.1007/s13225-018-0395-7: 132.
2018.
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Description and illustration: Wanasinghe et al. (2018).
Septoriella artemisiae (Wanas. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829677.
Basionym: Neostagonospora artemisiae Wanas. et al, Fungal Diversity 89: 130. 2018.
Description and illustration: Wanasinghe et al. (2018).
Septoriella arundinicola (Wanas. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829678.
Basionym: Poaceicola arundinicola Wanas. et al., Fungal Diversity 89: 135. 2018.
Description and illustration: Wanasinghe et al. (2018).
Septoriella arundinis (W.J. Li et al.) Y. Marín & Crous, comb. nov. MycoBank MB829679.
Basionym: Poaceicola arundinis W.J. Li et al., Mycosphere 6: 698. 2015.
Description: Li et al. (2015).
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Septoriella bromi (Wijayaw. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829680.
Basionym: Poaceicola bromi Wijayaw. et al., Mycosphere 6: 698. 2015.
Description and illustration: Li et al. (2015).
Septoriella dactylidicola Y. Marín & Crous, nom. nov. MycoBank MB829681.
Replaced synonym: Poaceicola dactylidis Tibpromma et al., Mycosphere 8: 755. 2017, non
Septoriella dactylidis (Wanas. et al.) Y. Marín & Crous (2019).
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Description and illustration: Thambugala et al. (2017).
Septoriella dactylidis (Wanas. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829682.
Basionym: Vagicola dactylidis Wanas. et al., Phytotaxa 6: 725. 2015.
Description and illustration: Jayasiri et al. (2015).
Description and illustration: Wehmeyer (1952).
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Septoriella elongata (Wehm.) Y. Marín & Crous, comb. nov. MycoBank MB829683.
Basionym: Leptosphaeria elongata Wehm., Mycologia 44: 633. 1952.
Synonym: Poaceicola elongata (Wehm.) W.J. Li et al., Mycosphere 6: 701. 2015.
Septoriella forlicesenica (Thambug. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829684.
Basionym: Poaceicola forlicesenica Thambug et al., Mycosphere 8: 756. 2017.
Description and illustration: Thambugala et al. (2017).
Septoriella garethjonesii (Thambug. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829685.
Basionym: Poaceicola garethjonesii Thambug et al., Mycosphere 8: 756. 2017.
Description and illustration: Thambugala et al. (2017).
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Septoriella germanica Crous, R.K. Schumach. & Y. Marín, sp. nov. MycoBank MB829701. Fig. 68.
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Etymology: Name refers to Germany, from where this fungus was isolated.
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Conidiomata solitary, pycnidial, erumpent, globose, brown, 180–220 µm diam, in vivo gregarious,
caespitose or in rows, but also pseudostromatic, up to 300 µm diam, with central ostiole, 30–40 µm diam;
conidiomatal wall of 3–4 layers of brown cells of textura angularis. Conidiophores reduced to
conidiogenous cells lining the inner cavity, hyaline, smooth-walled, globose to ampulliform, phialidic, 4–
6 × 4–5 µm. Conidia solitary, scolecosporous, fusoid to subcylindrical, apex subobtuse, base truncate,
straight to sligthly curved, 3–6-septate, golden-brown, smooth-walled, granular with mucoid caps at each
end, (35–)37–42(–46) × 3(–3.5) µm, in vivo 29–46 × 3–4.5 µm.
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium, covering dish in 2 wk.
On MEA, PDA and OA surface olivaceous grey, reverse iron-grey.
Typus: Germany, near Berlin, on dead culm of Phragmites australis (Poaceae), 16 Apr. 2016, R.K.
Schumacher, HPC 1168 (holotype CBS H-23875, culture ex-type CBS 145372 = CPC 30511).
Notes: Septoriella germanica is related to Sep. artemisiae. However, both species differ in the size of the
conidia [15–25 × 2–2.5 µm in Sep. artemisiae vs. (35–)37–42(–46) × 3(–3.5) µm in Sep. germanica], as
well as in the conidial septation (2–4 in Sep. artemisiae vs. 3–6 µm in Sep. germanica). Septoriella
germanica was isolated from Phragmites australis (Poaceae), while Sep. artemisiae was found on
Artemisia austriaca (Asteraceae).
Septoriella hibernica Crous, Quaedvl. & Y. Marín, sp. nov. MycoBank MB829703.
Etymology: Name refers to Ireland, where this fungus was collected.
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Culture sterile. Septoriella hibernica differs from its closest phylogenetic neighbour, Septoriella
subcylindrispora by unique fixed alleles in the ITS locus based on the alignment deposited in
TreeBASE (S23834): positions 5 (T), 33 (T), 34 (A), 46 (T), 61 (A), 89 (T), 477 (A), 479 (T), 480
(A), 512 (T), 528 (G), 534 (G).
Culture characteristics: Colonies erumpent, spreading, covering dish in 2 wk, with fluffy aerial mycelium
and even margins. On MEA, PDA and OA surface and reverse olivaceous grey.
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Typus: Ireland, on unidentified grass species (Poaceae), Mar. 2014, W. Quaedvlieg (holotype CBS H23874, culture ex-type CBS 145371 = CPC 24290).
Notes: Septoriella hibernica remained sterile on all media tested. It is related to Sep. subcylindrispora, but
the ITS sequences of both species showed only 97.75 % of nucleotide similarity (Identities = 522/538, 3
gaps).
Septoriella hollandica Crous, Quaedvl. & Y. Marín, sp. nov. MycoBank MB829702. Fig. 69.
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Etymology: Name refers to the Netherlands, where this fungus was collected.
Ascomata solitary, erumpent, globose, brown, 150–180 µm diam with central ostiole; conidiomatal wall
of 3–4 layers of brown cells of textura angularis. Pseudoparaphyses hyphae-like, hyaline, smoothwalled, branched, septate, 1.5–2 µm diam. Asci subcylindrical, flexuous, bitunicate, with well-defined
apical chamber, 1–1.5 µm diam, fasciculate, short stipitate, 70–90 × 8–10 µm. Ascospores bi- to triseriate,
fusoid-ellipsoid, 5-septate, constricted at median septum, medium brown, smooth-walled, guttulate,
widest above median septum, (27–)28–30(–32) × (4–)4.5(–5) µm.
Culture characteristics: Colonies flat, spreading, covering dish in 2 wk with moderate aerial mycelium.
On MEA surface vinaceous buff, reverse sienna; on PDA surface isabelline, reverse hazel; on OA surface
saffron.
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Typus: The Netherlands, Oosterbeek, on leaves of Phragmites australis (Poaceae), 24 Jan. 2014, W.
Quaedvlieg (holotype CBS H-23877, culture ex-type CBS 145374 = CPC 24109).
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Notes: Septoriella hollandica is related to Sep. chlamydospora and Sep. tridentina. Septoriella hollandica,
as well as Sep. chlamydospora and Sep. tridentina, produce sexual morphs in culture. Septoriella
hollandica can be easily distinguished from Sep. chlamydospora by its 5-septate ascospores, being 9septate in Sep. chlamydospora. Septoriella tridentina is the only species of this complex that produces
ascospores surrounded by a mucilaginous sheath. The asexual morph was reported only for Sep.
chlamydospora. However, as it was mentioned above, the asexual morph described in Sep.
chlamydospora (Jayasiri et al. 2015) corresponds to chains of chlamydospores instead of scolecosporous
conidia typical of Septoriella. Septoriella hollandica was isolated from Phragmites australis, while the
other two species were found on Dactylidis spp.
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Septoriella italica (Thambug. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829704.
Basionym: Poaceicola italica Thambug et al., Mycosphere 8: 759. 2017.
Description: Thambugala et al. (2017).
Septoriella muriformis (Ariyaw., Camporesi & K.D. Hyde) Y. Marín & Crous, comb. nov.
MycoBank MB829705.
Basionym: Allophaeosphaeria muriformis Ariyaw., et al., Fungal Diversity 72: 137.
Descriptions and illustrations: Liu et al. (2015), Thambugala et al. (2017).
Septoriella neoarundinis Y. Marín & Crous, nom. nov. MycoBank MB829706.
Replaced synonym: Vagicola arundinis Phukhams., Camporesi & K.D. Hyde, sp. nov. MycoBank
MB831056, non Septoriella arundinis (W.J. Li et al.) Y. Marín & Crous
Synonym: Vagicola arundinis Phukhams. et al. Mycosphere 8: 763. 2017. (nom. inval., Art. 40).
Etymology: Name reflects the host genus Arundo from which it was isolated.
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Description and illustration: Thambugala et al. (2017)
Typus: Italy, Province of Marsignano, Predappio, on a dead stem of Arundo plinii (Poaceae), 10 Nov.
2014, E. Camporesi IT 2223A (holotype MFLU 17-0016, ex-type living culture MFLUCC 15-0027).
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Notes: This species was initially introduced by Thambugala et al. (2017) as Vagicola arundinis.
However, it was invalid since two different holotype numbers were cited. Therefore, this species is
validated here and a new name in Septoriella is proposed, using a new epithet since Sep. arundinis is
occupied.
Septoriella neodactylidis Y. Marín & Crous, nom. nov. MycoBank MB829707.
Replaced synonym: Allophaeosphaeria dactylidis Wanas. et al., Fungal Diversity 72: 137. 2015, non
Septoriella dactylidis (Wanas. et al.) Y. Marín & Crous (2019).
Description and illustration: Liu et al. (2015).
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Septoriella pseudophragmitis Crous, Quaedvl. & Y. Marín, sp. nov. MycoBank MB829708. Fig. 70.
Etymology: Name refers to its morphological similarity with Sep. phragmitis, which occurs on the same
host.
Conidiomata solitary, pycnidial, erumpent, globose, brown-black, 200–250 µm diam with central ostiole;
conidiomatal wall of 6–8 layers of brown textura angularis. Conidiophores reduced to conidiogenous
cells lining the inner cavity, hyaline, smooth-walled, globose to ampulliform, phialidic, 4–10 × 4 µm.
Conidia solitary, subcylindrical, golden-brown, guttulate, smooth-walled, apex obtuse, base truncate, 3(–
6)-septate, with mucoid caps at each end, (20–)24–28(–32) × (3–)3.5(–4) µm.
Culture characteristics: Colonies erumpent, spreading, covering dish after 2 wk at 25 °C, with moderate
aerial mycelium and feathery margins. On MEA surface olivaceous grey, reverse iron-grey.
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Typus: The Netherlands, on leaves of Phragmites sp. (Poaceae), 5 Mar. 2014, W. Quaedvlieg (holotype
CBS-H 23904, culture ex-type CBS 145417 = CPC 24166).
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Notes: Septoriella pseudophragmitis is similar to Sep. phragmitis, which is reported from the same host,
Phragmites (Poaceae). These species differ in the size of their conidiomata (up to 250 µm diam in Sep.
pseudophragmitis vs. 350 m diam in Sep. phragmitis) and conidia [(20–)24–28(–32) × (3–)3.5(–4) µm in
Sep. pseudophragmitis vs. (29–)32–40(–46) × 3(–3.5) m diam in Sep. phragmitis], as well as in the
conidial septation, being mostly 3-septate in Sep. pseudophragmitis and 5-septate in Sep. phramitis.
Based on our phylogenetic analysis, Septoriella pseudophragmitis is related to Sep. allojunci. However,
Sep. allojunci produces smaller conidiomata (up to 150 µm) and larger conidia (48–70 × 3–6.6 m).
Moreover, Sep. allojunci was isolated from Juncus (Juncaceae).
Septoriella rosae (Mapook et al.) Y. Marín & Crous, comb. nov. MycoBank MB829713.
Basionym: Poaceicola rosae Mapook et al., Fungal Diversity 89: 136. 2018.
Description and illustration: Wanasinghe et al. (2018).
Septoriella subcylindrospora (W.J. Li et al.) Y. Marín & Crous, comb. nov. MycoBank MB829709.
Basionym: Allophaeosphaeria subcylindrospora W.J. Li et al., Fungal Diversity 75: 100. 2015.
Description and illustration: Ariyawansa et al. (2015a).
Septoriella vagans (Niessl) Y. Marín & Crous, comb. nov. MycoBank MB829710.
Basionym: Pleospora vagans Niessl, Verh. nat. Ver. Brünn 14: 174. 1876.
Synonym: Vagicola vagans (Niessl) O.E. Erikss. et al., Fungal Diversity 75: 115. 2015.
Description and illustration: Jayasiri et al. (2015).
Arezzomyces Y. Marín & Crous, gen. nov. MycoBank MB829711.
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Etymology: Name reflects the Italian province Arezzo where it was collected.
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Ascomata solitary, scattered, immersed to erumpent, obpyriform, dark brown to black, coriaceous, with
ostiole filled with hyaline cells, appearing as a white ring around ostiole; necks papillate, black, smooth;
ascomatal wall comprising 6–8 layers, outer layer heavily pigmented, comprising blackish to dark
brown, thick-walled cells of textura angularis, inner layer composed of brown, thin-walled cells of
textura angularis. Hamathecium comprising numerous, filamentous, branched, septate
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical, pedicel furcate, rounded and
thick-walled at the apex, with an ocular chamber. Ascospores mostly uniseriate, initially hyaline,
becoming yellowish brown at maturity, ellipsoidal, muriform, with 6–8 transverse septa, 3–7 vertical
septa, strongly constricted at the central septa, weakly constricted at the other septa, with conical and
narrowly rounded ends, lacking a mucilaginous sheath. Asexual morph not observed.
Culture characteristics: Colonies spreading, surface erumpent, with moderate aerial mycelium, and
feathery margins. On MEA, PDA and OA surface dirty white; reverse dirty white to luteous.
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Type species: Arezzomyces cytisi (Wanas. et al.) Y. Marín & Crous. Holotype and ex-type cultures:
MFLU 15-1502, MFLUCC 15-0649.
Notes: Arezzomyces is introduced to accommodate Allophaeosphaeria cytisi since, based on
phylogenetic data, it is located in an independent lineage distant to the clade representing the genus
Septoriella. Moreover, based on a megablast search using the ITS sequence, the closest matches in
NCBIs GenBank nucleotide database were Ophiosimulans tanaceti [GenBank KU738890; Identities =
534/586 (91 %), 11 gaps (1 %)], Ophiobolus cirsii [GenBank KM014664; Identities = 514/566 (91 %),
22 gaps (1 %)], and Chaetosphaeronema hispidulum [GenBank KX096655; Identities 535/588 (91 %),
22 gaps (3 %)]. Arezzomyces cytisi is a saprobe found on dead herbaceous branches of Cytisus.
Arezzomyces cytisi (Wanas. et al.) Y. Marín & Crous, comb. nov. MycoBank MB829712.
Basionym: Allophaeosphaeria cytisi Wanas. et al., Fungal Diversity 75: 97. 2015.
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Description and illustration: Ariyawansa et al. (2015a).
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Typus: Italy, Arezzo Province, Casuccia di Micheli in Quota, dead and hanging branches of Cytisus sp.
(Fabaceae), 20 Jun. 2012, E. Camporesi (holotype MFLU 15-1502, culture ex-type MFLUCC 150649).
Authors: Y. Marin-Felix, W. Quaedvlieg, R.K. Schumacher & P.W. Crous
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Setophoma Gruyter et al., Mycologia 10: 1077. 2010. Fig. 71.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
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Type species: Setophoma terrestris (H.N. Hansen) Gruyter et al., basionym: Phoma terrestris H.N.
Hansen. Lectotype and ex-lectotype strain designated by de Gruyter et al. (2010): CBS H-20311, CBS
335.29.
DNA barcode (genus): LSU. Fig. 28.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 17. Fig. 72.
Ascomata scattered, gregarious, immersed, visible as raised, minute black dots on host surface,
uniloculate, globose to subglobose, glabrous, brown to dark brown, ostiole central, with minute
papilla; ascomatal wall thin, of equal thickness, composed of pseudoparenchymatous cells, arranged
in flattened textura angularis to textura prismatica. Hamathecium composed of numerous, filiform,
broad cellular pseudoparaphyses, with distinct septa, anastomosing at apex. Asci 8-spored, bitunicate,
fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with well-developed
narrowly ocular chamber. Ascospores overlapping, 2-seriate, phragmosporous, cylindrical to
cylindrical-clavate, hyaline, 3-septate, usually enlarged at the second cell from apex, smooth-walled
with large guttules. Conidiomata pycnidial, solitary to confluent, superficial or submerged in agar,
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globose to subglobose, setose, with papillate necks, honey to olivaceous or olivaceous black, with 2–
7(–11) layers of pseudoparenchymatal cells. Conidiogenous cells hyaline, monophialidic. Conidia
aseptate, ellipsoidal to subcylindrical or subfusoid, guttulate (adapted from Quaedvlieg et al. 2013,
Phookamsak et al. 2014a).
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Culture characteristics: Colonies spreading with sparse to moderate aerial mycelium, smooth or folded
surface, even or lobate margins. On PDA surface iron-grey or grey olivaceous with outer region irongrey; reverse olivaceous grey or iron-grey. On MEA surface olivaceous grey or umber with patches of
apricot and dirty white; reverse ochreous or cinnamon with patches of olivaceous grey. On OA surface
isabelline or iron-grey surrounded by orange to apricot diffuse pigment layer in agar.
Optimal media and cultivation conditions: OA and SNA at 25 °C under continuous near-ultraviolet light
to promote sporulation. Sterile bamboo pieces on WA to induce sporulation of the sexual morph.
Distribution: Worldwide.
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Hosts: Mainly on members of Poaceae, but also of Amaryllidaceae and Asteraceae, among others.
Disease symptoms: Leaf spots and necrosis, leaf dieback, and pink root.
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Notes: Setophoma was introduced by de Gruyter et al. (2010) to accommodate two species previously
placed in Pyrenochaeta, i.e. Pyr. sacchari and Pyr. terrestris. Setophoma is characterised by pycnidial
conidiomata covered by setae, phialidic conidiogenous cells, and hyaline, ellipsoidal to subcylindrical,
aseptate, guttulate conidia (de Gruyter et al. 2010, Quaedvlieg et al. 2013). When Setophoma was
introduced, the sexual morph had not been observed. Subsequently, Phookamsak et al. (2014a) reported
the sexual morph of this genus. It was found causing leaf spots of sugarcane (Saccharum officinarum),
and based on the phylogenetic data it was shown to be the sexual morph of Set. sacchari. This sexual
morph is similar to Phaeosphaeria species, producing ascospores with three septa with the second cell
from the apex being swollen; these cells differ in colour (hyaline in Setophoma vs. yellowish to brown
in Phaeosphaeria).
Setophoma encompasses pathogenic or saprobic species associated with monocotyledonous plants (de
Gruyter et al. 2010). The type species, Set. terrestris, causes pink root on Allium spp., and also on Zea
mays and Oryza sativa, but it is asymptomatic on other hosts (Farr & Rossman 2019). Setophoma
vernoniae produces leaf spots on Vernonia polyanthes (Crous et al. 2014b), while Set. sacchari is
considered a weak pathogen of members of the Poaceae that is only noticeable when conditions are
favourable for disease spread, and causes leaf spots and necrosis and leaf dieback (Farr & Rossman
2019).
In our phylogenetic analysis based on ITS and LSU (Fig. 28), the clade representing the genus
Setophoma is well-supported (95 % BS / 1 PP). However, the most recently described species, Set.
cyperi, is not located in that clade, representing a new genus in the family Phaeosphaeriaceae.
At the proof stage of this paper, a new publication appeared on Setophoma (Liu et al. 2019), which
contains four new species.
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References: de Gruyter et al. 2010, Quaedvlieg et al. 2013, Phookamsak et al. 2014a (morphology and
phylogeny).
Setophoma brachypodii Crous, R.K. Schumach. & Y. Marín, sp. nov. MycoBank MB829669.
Etymology: Name reflects the host genus Brachypodium from which it was isolated.
Culture sterile. Setophoma brachypodii differs from its closest phylogenetic neighbour, Setophoma
terrestris by unique fixed alleles in two loci based on alignments of the separate loci deposited in
TreeBASE (S23834): LSU positions 42 (G), 67 (C), 75 (C), 77 (T), 79 (G), 81 (C), 82 (A), 89 (C),
133 (C), 144 (G), 145 (C), 146 (C), 147 (T), 150 (G), 302 (C), 348 (T), 380 (C), 392 (A), 437 (T), 445
(T), 446 (C), 473 (A), 477 (G), 601 (G), 636 (T), 637 (T), 638 (A); ITS positions 32 (C), 34 and 35
(indels), 36 (T), 37 (T), 38 (T), 42 (G), 43 (T), 44 (A), 54 (C), 56 (G), 57 (T), 58 (T), 59 (C), 60 (G),
61 (C), 62 (T), 63 (G), 64 (T), 66 (G), 67 (T), 72 (T), 77 (G), 78 (T), 80 (T), 99 (T), 100 (G), 101 (A),
103 (C), 114 (C), 117 (G), 118 (T), 119 (A), 121 (C), 122 (T), 124 (C), 130 (A) 138 (C), 140 (A), 143
(T), 146 (A), 148 (C), 172 (A), 176 (T), 178 (A), 180 (T), 182 (A), 186 (indel), 354 (T), 379 (T), 381
(indel), 388 (T), 389 (G), 390 (G), 391 (T), 392 (C), 393 (C), 394 (T), 395 (C), 396 (T), 399 (G), 400
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(A), 401 (C), 402 (C), 409 (A), 418 (A), 419 (T), 433 (G), 434 (T), 435 (A), 441 (G), 444 (T), 467
(A), 470 (indel), 473 (T), 475 (C), 477 (A), 478 (C), 479 (T), 482 (A), 485 (C), 486 (C), 490 (A), 495–
498 (indels), 499 (C), 500 (C), 502 (T), 504 (A), 506 (T), 507 (A), 511 (C).
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and even, lobate
margins, reaching 60 mm diam after 2 wk. On MEA, PDA and OA, surface and reverse olivaceous grey.
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Typus: Belgium, Dinant, 173 m a.s.l., on border of calcareous meadow, on a dead and attached leaf of
Brachypodium sylvaticum (Poaceae), 2 Nov. 2016, L. Bailly & R.K. Schumacher, HPC 1503, RKS 1
(holotype CBS H-23905, culture ex-type CBS 145418 = CPC 32492).
Notes: Setophoma brachypodii remained sterile on all media tested, and the original specimen was
depleted, hence we could not describe it based on morphology. This is the first species of Setophoma
reported on Brachypodium.
Setophoma pseudosacchari Crous & Y. Marín, sp. nov. MycoBank MB829670. Fig. 73.
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Etymology: Named after its closely phylogenetic relation to Setophoma sacchari.
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Ascomata developing on OA, solitary, erumpent, brown, 200–300 µm diam, globose, with large central
ostiole, 30–40 µm diam; ascomatal wall of 3–4 layers of brown cells of textura angularis, ascomata
setose; setae brown, flexuous, thick-walled, septate, base verruculose, with slight taper to obtuse apex, up
to 150 µm long. Pseudoparaphyses hyphae-like, anastomosing, branched, septate, hyaline, occurring
intermingled among asci. Asci bitunicate, ellipsoid to subcylindrical, hyaline, curved to straight,
fasciculate, apex obtuse, with well-defined ocular chamber, 2 µm diam, stipitate, 70–100 × 10–13 µm.
Ascospores bi- to triseriate, fusoid-ellipsoid with subobtuse ends, straight, 3-septate, widest in second cell
from apex, prominently guttulate, hyaline, smooth-walled, (22–)25–30 × (5.5–)6 µm. Conidiomata
developing on SNA, solitary to aggregated, erumpent, brown, globose, 200–300 µm diam, with 1–2
ostioles, lacking setae; conidiomatal wall of 2–3 layers of brown textura angularis. Conidiophores
reduced to conidiogenous cells lining inner cavity, dissolving at maturity, hyaline, smooth-walled,
globose to ampulliform, phialidic, 4–6 × 5–6 µm. Conidia solitary, aseptate, straight to slightly curved,
subcylindrical to fusoid-ellipsoid, apex obtuse, base truncate, hyaline, smooth-walled, guttulate, (8–)11–
12(–14) × (3–)4 µm.
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Culture characteristics: Colonies erumpent, spreading, surface folded, with moderate aerial mycelium
and even, lobate margins, reaching 55 mm diam. On MEA surface peach, outer region scarlet, reverse
sienna; on PDA surface umber, outer region saffron, reverse sienna with patches of saffron; on OA
surface sienna with patches of saffron.
Typus: France, La Réunion Island, leaf spots on Saccharum officinarum (Poaceae), May 2015, P.W.
Crous, HPC 296 (holotype CBS H-23876, CBS 145373 = CPC 26421).
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Notes: This species is closely related to Nph. sacchari, which is a species also isolated from sugarcane.
However, the ITS sequences of the type material of both species showed only a 97.68 % of nucleotide
similarity. Unfortunately, tef1 and tub2 sequences of Nph. sacchari are not available in order to compare
both species. These species produce both the sexual and asexual morphs, with morphological differences
most obvious in the sexual morph. Neosetophoma pseudosacchari can be easily distinguished by its
larger ascomata (up to 300 m diam in Nph. pseudosacchari vs. up to 180 m diam in Nph. sacchari), asci
[70–100 × 10–13 µm in Nph. pseudosacchari vs. 60–75(–85) × 12–15(–17) m in Nph. sacchari] and
ascospores [(22–)25–30 × (5.5–)6 µm in Nph. pseudosacchari vs. 20–23(–25) × 5–6 m in Nph. sacchari].
Wingfieldomyces Y. Marín & Crous, gen. nov. MycoBank MB829671. Table 18. Fig. 74.
Etymology: Named in honour of its collector, Prof. dr M.J. Wingfield, who contributed greatly to the
elucidation of African fungal biodiversity.
Ascomata immersed on host, erumpent in culture, black, globose, with central ostiole; ascomatal wall of
3–4 layers of dark brown cells of textura angularis. Pseudoparaphyses intermingled among asci,
hyaline, septate, branched prominently, constricted at septa. Asci bitunicate with apical chamber,
subcylindrical, hyaline, smooth, fasciculate, stipitate, 8-spored. Ascospores tri- to multiseriate, fusoid
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with subobtusely rounded ends, finely verruculose, red-brown, guttulate, 2-septate, slightly constricted at
septa, with central cell somewhat swollen.
Culture characteristics: Colonies spreading, erumpent surface, with moderate aerial mycelium, margins
feathery. On MEA, PDA and OA surface dirty white; reverse dirty white to luteous.
Type species: Wingfieldomyces cyperi (Crous & M.J. Wingf.) Y. Marín & Crous. Holotype and ex-type
cultures: CBS H-22622, CBS 141450 = CPC 25702.
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Notes: Wingfieldomyces is introduced to accommodate Set. cyperi since, based on phylogenetic data, it is
located in an independent lineage distant to the clade representing the genus Setophoma. Moreover,
based on a megablast search using the ITS sequence, the closest matches in NCBIs GenBank nucleotide
database were Pringsheimia euphorbiae [GenBank NR_145344; Identities = 456/500 (91 %), 8 gaps (1
%)] and Phaeosphaeria caricis [GenBank KY090633; Identities 439/485 (91 %), 12 gaps (2 %)]. It only
produces a sexual morph in culture, characterised by tri- to muliseriate, 2-septate, red-brown ascospores,
while Setophoma produces both morphs and 2-seriate, 3-septate, hyaline ascospores with the second cell
from the apex becoming swollen. Wingfieldomyces is associated with leaf scorch symptoms on Cyperus.
Wingfieldomyces cyperi (Crous & M.J. Wingf.) Y. Marín & Crous, comb. nov. MycoBank MB829672.
Fig. 74.
Basionym: Setophoma cyperi Crous & M.J. Wingf., Persoonia 36: 385. 2016.
Description: Crous et al. (2016b).
Typus: South Africa, Eastern Cape Province, Haga Haga, on leaves of Cyperus sphaerocephala
(Cyperaceae), Dec. 2014, M.J. Wingfield (holotype CBS H-22622, culture ex-type CPC 25702 = CBS
141450).
Authors: Y. Marin-Felix & P.W. Crous
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Stagonosporopsis Died., Ann. Mycol. 10: 142. 1912. Emend. Aveskamp et al., Stud. Mycol. 65: 44.
2010. Fig. 75.
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Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Didymellaceae.
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Type species: Stagonosporopsis boltshauseri (Sacc.) Died., designated as lectotype by Clements & Shear
(1931), basionym: Ascochyta boltshauseri Sacc. = Stagonosporopsis hortensis (Sacc. & Malbr.) Petr.,
basionym: Hendersonia hortensis Sacc. & Malbr. Representative strain of Sta. hortensis: CBS 572.85 =
PD 79/269.
DNA barcode (genus): LSU.
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DNA barcodes (species): ITS, rpb2, tub2. Table 19. Fig. 76.
Ascomata pseudothecial, globose to subglobose, sometimes with a somewhat conical neck. Asci
cylindrical or subclavate, 8-spored, biseriate. Ascospores ellipsoid, fusoid or obovoid, uniseptate,
guttulate, sometimes with a gelatinous sheath. Conidiomata pycnidial, globose to subglobose, glabrous
or with hyphal outgrowths, superficial on agar surface or immersed, solitary or confluent, ostiolate or
poroid, occasionally papillate; conidiomatal wall pseudoparenchymatous, 2–6 layered, with an outer
wall composed of 1–3 layers of brown to olivaceous cells. Conidiogenous cells phialidic, hyaline,
simple, smooth-walled, ampulliform or doliiform. Conidia often dimorphic: mainly aseptate, hyaline,
ellipsoid to subglobose, thin-walled, smooth-walled, eguttulate or with several polar or scattered
guttules; second type of conidia larger in size, can be produced both in vivo and in vitro in the same
pycnidia as the other type of conidia, 0–3-septate.
Culture characteristics: Colonies on OA regular to somewhat irregular, colourless, buff, luteous to
ochraceous or amber, or olivaceous grey to greenish grey, with scarce or abundant floccose white to
pale salmon, or olivaceous grey aerial mycelium.
Optimal media and cultivation conditions: On OA at 20–24 °C under near-ultraviolet light (13 h light, 11
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h dark) to induce sporulation of the asexual morph, while MEA stimulates pigmentation and crystal
formation. Changes in colour of the fungal cultures upon a sudden increase of pH (NaOH spot test),
which may be used for taxonomic characterisation, are best observed on OA.
Distribution: Worldwide.
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Hosts: Associated with at least 30 plant genera in Asteraceae, Brassicaceae, Campanulaceae,
Caricaceae, Cucurbitaceae, Fabaceae, Lamiaceae, Ranunculaceae, Solanaceae and Valerianaceae as
saprobes or pathogens. Stagonosporopsis oculo-hominis is the only species that is not associated with a
plant host and was isolated from human corneal ulcer in the USA.
Disease symptoms: Plant stunting, seedling damping-off, leaf spots and dieback, crown rot, stem canker,
flower blight, and fruit rot.
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Notes: Many species of Stagonosporopsis are phytopathogens, causing devastating diseases on plants
from various families. Some species have a worldwide distribution, e.g., Sta. cucurbitacearum on
Cucurbitaceae, and Sta. hortensis on Fabaceae, while others represent important quarantine plant
pathogens limited to certain geographical areas. For example, Sta. andigena and Sta. chrysanthemi are
classified as A1 and A2 quarantine pathogens, respectively, by the European and Mediterranean Plant
Protection Organisation (EPPO; 2016). Stagonosporopsis tanaceti is a destructive pathogen of
pyrethrum (Tanacetum cinerariifolium) in Australia but has not been reported elsewhere in the world
(Vaghefi et al. 2012). Some Stagonosporopsis species have been isolated from plants but their
pathogenicity has not been established. For example, Sta. dennisii has been reported from dead stems of
Solidago spp. but no data are available on its pathogenicity (Boerema et al. 2004).
Species identification based on only morphology is unreliable. Stagonosporopsis was originally
separated from Ascochyta based on occasional formation of multi-septate (stagonospora-like) conidia
(Diedicke 1912). However, later phylogenetic studies revealed that some Stagonosporopsis spp. lack
the stagonospora-like spores or any features except for globose pycnidial conidiomata, and aseptate,
hyaline conidia (Aveskamp et al. 2010). Thus, multilocus sequence typing is essential for
identification of Stagonosporopsis species. The emended description of the genus Stagonosporopsis by
Aveskamp et al. (2010) states that the sexual morph of Stagonosporopsis, if present, occurs only in vivo.
However, some strains of Sta. chrysanthemi, Sta. caricae and Sta. inoxydabilis have been shown to
produce pseudothecial ascomata intermingled with pycnidial conidiomata on agar media (Boerema et al.
2004, Vaghefi et al. 2012).
Currently, more than 40 species are linked to the genus Stagonosporopsis. However, only 27
species are recognised based on molecular data (Table 19). Previous phylogenetic studies have used
one locus (act in De Gruyter et al. 2012), three loci (LSU, ITS and tub2 in Aveskamp et al. 2010),
four loci (LSU, ITS, tub2 and act in Hyde et al. 2014; LSU, ITS, tub2 and rpb2 in Chen et al. 2015;
ITS, tub2, chs and cal in Stewart et al. 2015) and five loci (LSU, ITS, tub2, act and tef1 in Vaghefi et
al. 2012) for phylogenetic species recognition in Stagonosporopsis. However, in most cases, ITS and
tub2 sequences are sufficient for achieving resolution to species level. While ITS sequences alone
may be used to distinguish Stagonosporopsis as a monophyletic clade within Didymellaceae, tub2
fails to distinguish Stagonosporopsis and, thus, needs to be always combined with ITS (Chen et al.
2015). A phylogeny produced by rpb2 alone is highly similar to the combined four-locus phylogeny
based on LSU, ITS, tub2 and rpb2. However, amplification of rpb2 has not been successful for many
Stagonosporopsis spp. (Chen et al. 2015). Likewise, while partial cal sequences provide high
resolution for Stagonosporopsis species delineation, it has not been successfully amplified in some
strains (Aveskamp et al. 2010, Vaghefi et al. 2012). Thus, the use of ITS and tub2 is recommended as
they will provide sufficient resolution for almost all Stagonosporopsis species, are easier to amplify,
and are available for the majority of Stagonosporopsis spp. described to date (Table 19). The only two
species that cannot be separated based on the LSU-ITS-tub2 phylogeny are S. bomiensis and S.
papillata, for which sequencing of rpb2 was necessary (Chen et al. 2017).
References: Boerema et al. 2004 (morphology and distribution); Aveskamp et al. 2010, Chen et al.
2015 (morphology and phylogeny).
Stagonosporopsis chrysanthemi (F. Stevens) Crous et al., Australas. Pl. Pathol. 41: 681. 2012.
Basionym: Ascochyta chrysanthemi F. Stevens, Bot. Gaz. 44: 246. 1907.
Synonyms: Mycosphaerella ligulicola, K.F. Baker et al., Phytopathology 39: 799. 1949.
Didymella ligulicola (K.F. Baker et al.) Arx, Beitr. Kryptfl. Schweiz 11: 364. 1962.
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Didymella ligulicola var. ligulicola (K.F. Baker et al.) Arx, Stud. Mycol. 32: 199. 1990.
Phoma ligulicola var. ligulicola Boerema, Stud. Mycol. 32: 9. 1990.
Stagonosporopsis ligulicola var. ligulicola (K.F. Baker et al.) Aveskamp et al., Stud. Mycol. 65: 46.
2010.
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Typus: USA, North Carolina, West Raleigh, on Chrysanthemum indicum, Dec. 1906, F.L. Stevens
(Bartholomew, Fungi Columbiani no. 2502, Field Museum of Natural History, C0004169F;
designated here as lectotype, MBT385563); on Chrysanthemum morifolium, 1949, L.H. Davis
[epitype of Ascochyta chrysanthemi designated here ATCC 10748, MBT385567 (preserved in a
metabolically inactive state)].
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Notes: Stevens (1907) described Ascochyta chrysanthemi on Chrysanthemum indicum from North
Carolina; however, he did not refer to a holotype specimen in the original description. A specimen at
Field Museum of Natural History (C0004169F) is chosen as lectotype, among numerous other
duplicates deposited at BPI, CUP, NYBG, MSC, and various other herbaria that include collections
distributed as E. Bartholomew, Fungi Columbiani 2502. Since no living cultures derived from these
specimens are available, we designate ATCC 10748, isolated from Chrysanthemum morifolium from
North Carolina, as ex-epitype culture of Ascochyta chrysanthemi here.
Authors: N. Vaghefi, Y. Marin-Felix, P.W. Crous & P.W.J. Taylor
Stemphylium Wallr., Flora Cryptogamica Germaniae 2: 300. 1833. Fig. 77.
Synonyms: Scutisporium Preuss, Linnaea 24: 112. 1821.
Epochniella Sacc., Michelia 2: 127. 1880.
Soreymatosporium Sousa da Câmara, Proposta Stemphylium: 18. 1930.
Thyrodochium Werderm., Annls mycol. 22: 188. 1942.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae.
Type species: Stemphylium botryosum Wallr. = Pleospora tarda E.G. Simmons. Holotype of Ste.
botryosum: "Ad sparagam" in herb. Wallroth, STR. Ex-type strain of Ple. tarda: CBS 714.68.
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DNA barcode (genus): ITS.
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DNA barcodes (species): cmdA, gapdh. Table 20. Fig. 78.
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Ascomata pseudothecial, globose or ovoid, membranous, dark brown to black, sometimes with a slender
neck. Asci oblong to clavate, with distinct outer and inner walls. Ascospores elongate to oval, with 7
horizontal and 3–5 longitudinal septa, yellowish to brown, muriform on maturity. Conidiophores dark due
to percurrent proliferation forming phaeodictyospores, mostly solitary, straight or flexuous, short or long,
branched or unbranched, aseptate or septate. Conidiophores proliferate further after a conidium is
produced, producing new cells and new conidia. Conidiogenous cells swollen at apex, single or in group.
Conidia olive, dark or pale brown, verrucose, oblong or muriform, with 3 or more constricted transverse
and 1–2 longitudinal or oblique septa.
Culture characteristics: Stemphylium colonies grow rapidly on a variety of media. On most media, the
colonies are velvety to cottony in texture with a pale or dark olivaceous grey, brown or brownish black
colour and black pigmentation on the colony reverse. Conidial density is low in cultures produced under
laboratory conditions even when the isolate is grown under alternate cycle of 12 h light and 12 h darkness
on PDA. Aerial mycelia flat/effuse, woolly or compact. Margins smooth and sharp or crenate and lobate.
Optimal media and cultivation conditions: SNA for pigmentation and morphological identification, PCA for
morphological identification and alternatively PDA for pigmentation and morphological identification.
Incubation for 1–2 wk at moderate temperatures from 23–27 °C (depending on the species) under cool white
florescent light with an 8-hr or 12-hr photoperiod.
Distribution: Worldwide.
Hosts: Many Stemphylium species are saprophytes and grow on plant debris and cellulose material.
However, plant pathogenic species, such as Ste. beticola, Ste. botryosum, Ste. loti, Ste. solani and Ste.
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vesicarium, can cause devastating damage and significant loss of agriculturally important crops annually.
Stemphylium spp. are pathogenic to a wide range of hosts, such as tomato, garlic, asparagus, alfalfa, lupin,
lentil and cotton. The ability of pathogens to infect a wide range of crops reflects its adaptability to wide
range of climatic conditions and provides better survival chances.
Disease symptoms: Leaf spot, defoliation, curling and bending of the leaf margins and stems. Lesion size
differs in various hosts and can grow to encompass the entire leaf and reduce photosynthesis.
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Notes: Simmons (1967) established criteria for morphological identification of various Stemphylium spp.
and introduced Pleospora herbarum as the sexual morph of Ste. botryosum. However, Simmons (1985)
subsequently reclassified Stemphylium/Pleospora holomorphs and reported P. tarda as the sexual morph
of Ste. botryosum and P. herbarum the sexual morph of Ste. herbarum (Moslemi et al. 2017). The asexual
morph Stemphylium has been well studied, though the sexual morph Pleospora is poorly defined. The
number of Pleospora spp. identified may be as many as 1000 and they are reported to be polyphyletic.
Stemphylium is morphologically similar to the closely related genus Alternaria. However, unlike
Alternaria in which its conidia remain attached and form a chain, Stemphylium conidia are always
solitary, arising from a conidiogenous cell with a swollen apex (Inderbitzin et al. 2009). Morphological
features such as spore shape and size, conidiophores, ascospores and the size and time of pseudothecial
maturation are important characteristics in species identification. Other features such as variation in
conidial wall ornamentation and septum development are not considered as important parameters in
Stemphylium identification (Câmara et al. 2002).
For phylogenetic analyses of Stemphylium species, cmdA and gapdh were identified as the most
informative genes, and rpb2 and actA as the least informative (Woudenberg et al. 2017). Among ITS,
cmdA and gapdh, cmdA provides the highest resolution; however, more significant support is obtained
when all three loci are combined.
Environmental factors such as temperature and moisture are key in Stemphylium disease development.
Plant debris and seeds are primary sources of inoculum of Stemphylium in most host plant species. When
environmental conditions are favourable, the pathogen can cause significant loss to various agricultural
crops such as lupin and cotton (Boshuizen et al. 2004).
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References: Simmons (1967), Ellis (1971), Bayaa & Erskine (1998), Câmara et al. (2002) (morphology);
Bashi & Rotem (1975), Mwakutuya (2006) (culture characteristics); Boshuizen et al. (2004) (biology and
life cycle); Wang et al. (2010) (host range); Woudenberg et al. (2017), Crous et al. (2019b) (optimal
media and growing conditions).
Stemphylium rombundicum Moslemi, Y.P. Tan & P.W.J. Taylor, sp. nov. MycoBank MB829291. Fig.
79.
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Etymology: Named after the famous beverage, Bundaberg Rum (Bundy Rum), produced in Bundaberg,
Queensland, Australia, where the fungus was first isolated.
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Conidiophores long, solitary, straight, septate, verrucose, light or dark brown, (62–)111–258(–307) m,
bearing one thickened, darkened, percurrent rejuvenation site. Conidiogenous cells swollen at apex,
darkened, (5–)6–9(–10) m wide. Conidia solitary, conidium body light brown to golden, turning into
dark brown around longitudinal and transverse septa, verrucose, oblong or cylindrical, occasionally ovoid
with curved apex, (27.5–)35–55.5(–61) × (10–)12.5–24(–26.5) m, with 3–4 transverse septa, 1–2(–3)
longitudinal or oblique septa per transverse sector. Sexual morph not observed.
Culture characteristics: Colonies on SNA after 1 wk reaching 35 mm diam, effuse, hairy or velvety,
white, colourless, mycelia mostly immersed in the agar. On PDA reaching 35 mm diam, fast growing,
with compact, entire, aerial mycelium, fine, woolly on the surface; reverse dark orange to dark brown in
the centre with central pale brown rings growing towards the sides; thick yellow margins, and grey zones
can also be seen.
Typus: Australia, Queensland, Bundaberg (Burnett Heads), from fruit lesions of Solanum lycopersicum
(Solanaceae), 9 Aug. 2000, J. Maltby (holotype BRIP 27486, culture ex-type BRIP 27486).
Notes: Colonies of Ste. lycopersici produce a yellow dark red pigmentation diffusing out in PDA and
other media (Yamamoto 1960). As Ste. rombundicum is closely related to species in the Ste. lycopersici
complex, similar physiological characters can be observed on PDA. Colonies only sporulated at 23 °C
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under 12 h photoperiod on PDA. Against Ste. lycopersici in which the conidia are mostly ovoid with a
pointed apex, Ste. rombondicum mostly contains cylindrical or oblong conidia. It is difficult to observe
the longitudinal septa in conidia of Ste. rombundicum. Conidiophores are significantly long compared to
the type Ste. lycopersici in which the conidiophores length do not exceed 140 µm. Conidia of Ste.
lycopersici are significantly longer (50–74 µm × 16–23 µm) than those of Ste. rombundicum.
Stemphylium truncatulae Moslemi, Y.P. Tan & P.W.J. Taylor, sp. nov. MycoBank MB829282. Fig. 80.
Etymology: Named after the host species, Medicago truncatula, from which it was first collected.
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Immature ascomata observed on SNA and PDA embedded in the agar. Ascomata pseudothecial, dark
brown to black, globose or flask shaped, solitary or aggregated in groups of 3–5, (83.5–)185.5–186(–304)
× (351–)453–483(–603) µm, with outgrowing dark mycelia; ascomatal wall 27–32 µm thick.
Conidiophores solitary, straight to flexuous, mostly branched, septate or occasionally aseptate, smoothwalled, light brown, mostly 7–11 m length, some 17–39 m in length, bearing 2–6 thickened, pale,
percurrent rejuvenation sites. Conidiogenous cells slender or slightly swollen at apex, pale, (5.5–)6.5–7(–
9.5) m. Conidia solitary, conidium body pale brown to golden or olivaceous brown, mostly smoothwalled, sometimes minutely verrucose, usually ovoid, occasionally with pointed apex, (8–)9.5–18(–21) ×
(14–)15.5–29.5(–32) m, with 2–4 transverse septa, one longitudinal or oblique septa per transverse
sector.
Culture characteristics: Colonies on SNA reaching 21–25 mm diam after 1 wk, with white and fluffy
aerial mycelia in the centre; reverse colourless with pale olivaceous grey centre. Colonies on PDA
reaching 25 mm diam after 1 wk, with white fluffy mycelia in the centre; reverse dark green to grey with
thin white margins.
Typus: Australia, Victoria, from seeds of Medicago truncatula (Fabaceae), 10 Sep. 1982, M. Mebalds
(holotype BRIP 14850, culture ex-type BRIP 14850).
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Notes: Differs from the type species Ste. botryosum described by Simmons (1969) by producing
significantly smaller conidia. According to Simmons (1969), conidia of Ste. botryosum are 24–26 µm
wide and 33–35 µm long. Additionally, colonies of Ste. botryosum grow rapidly to 48 mm diam after 6 d
of incubation at 25 °C (Hosen et al. 2013), while Ste. truncatulae is slow growing. The morphological
identifications along with sequence analyses support Ste. truncatulae as a unique taxon closely related to
the type species Ste. botryosum (CBS 714.68).
Stemphylium waikerieanum Moslemi, Jacq. Edwards & P.W.J Taylor, sp. nov. MycoBank MB829283.
Fig. 81.
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Etymology: Named after the location, Waikerie in South Australia, from where it was collected.
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Immature ascomata observed on SNA after 2 wk embedded in the agar. Ascomata pseudothecial, dark
brown to black, mostly flask-shaped or occasionally globose, solitary or aggregated in groups of 4–6,
(267.5–)292–349.5(–374) × (191.5–)235.5–337(–381) µm, with outgrowing mycelia; ascomatal wall
thin, 10–15 µm thick. Conidiophores solitary, straight, simple or occasionally 1-branched, septate,
smooth-walled, pale brown, (29–)42–85(–98) m long, cylindrical, enlarging apically to the site of
conidium production. Conidiogenous cells swollen at apex, darkened, (6–)6.5–7(–7.5) m wide. Conidia
solitary, dark reddish brown, verrucose, ovoid, oblong or cylindrical, (18–)25–49(–52) × (10.5–)14–26(–
30) m, with 2–5 transverse septa, 1–2(–3) longitudinal or oblique septa per transverse sector, constricted
at multiple darkened transverse septa.
Culture characteristics: Colonies on SNA reaching 28 mm diam after 1 wk with flat, entire and fluffy
aerial mycelia in the centre, sub-hyaline. On PDA reaching 20 mm diam after 1 wk, compact, entire,
aerial mycelium white, woolly, with rings of light olivaceous grey in the centre and dark olivaceous grey
to black on the reverse side, margins regular, thick and white.
Typus: Australia, South Australia, Waikerie, from leaf spots on Allium sativum (Alliaceae), 14 Nov.
1997, H. Suheri (holotype VPRI 21969, culture ex-type VPRI 21969).
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Notes: Stemphylium waikerieanum is morphologically similar to species in the Ste. vesicarium species
complex. However, the conidium length in Ste. vesicarium complex does not exceed 45 µm (Simmons
1969), whereas conidia of Ste. waikerieanum were observed up to 49 µm long. The multigene
phylogenetic analysis of ITS, gapdh and cmdA also support (PP value =1) this species as a novel taxon.
Authors: A. Moslemi, J. Edwards, Y.P. Tan & P.W.J. Taylor
Tubakia B. Sutton, Trans. Brit. Mycol. Soc. 60: 164. 1973. Fig. 82.
Synonyms: Actinopelte Sacc., Annls mycol. 11: 315. 1913.
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Classification: Dothideomycetes, Diaporthomycetidae, Diaporthales, Tubakiaceae.
Type species: Tubakia japonica (Sacc.) B. Sutton, basionym: Actinopelte japonica Sacc. Epitype and exepitype strains designated by Braun et al. (2018): NBRC H-11611, NBRC 9268 = MUCC2296 = ATCC
22472.
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DNA barcodes (genus): ITS, LSU and rpb2.
DNA barcodes (species): ITS, tef1 and tub2. Table 21.
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Ascomata perithecial, pigmented, dark, on fallen overwintered leaves, rostrate, beak short, usually
lateral-eccentric, slightly protuberant, with periphysate ostiolate; ascomatal wall variable in thickness,
paler than stromatic layers, polyascal. Paraphyses lacking. Asci unitunicate, 8-spored, oblongellipsoid, stalk short to oblong, ascal apex with two refractive conoid structures, asci deliquescing at
maturity. Ascospores more or less uniseriate, becoming irregularly biseriate, one-celled, hyaline,
ellipsoid to fusiform, often inequilateral or slightly curved, wall finely ornamented, content granularguttulate. Conidiomata pycnothyrial, usually circular or subcircular when viewed from above,
superficial, easily removable, scutellate, fixed to the leaf by a central columella; scutellum composed
of loose to dense hyphal strands, mostly branched, thick-walled, pigmented, margin compact or outer
portions of the radiating hyphal strands looser to free, tips rounded, truncate or pointed, margin
usually not recurved. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic,
usually subcylindrical-conical, lageniform, hyaline to pale brown, arising from small, colourless
fertile cells around the upper part of the central pycnothyrial columella, percurrently proliferating,
sometimes forming indistinct periclinal thickenings or annellations (collarettes). Conidia formed
singly, globose to broad ellipsoid-obovoid, sometimes subcylindrical or somewhat irregular, wall thin
to somewhat thickened, smooth to faintly rough, hyaline to pigmented, apex rounded, base rounded to
attenuated, sometimes with distinct frill or peg-like basal hilum (adapted from Braun et al. 2018).
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Culture characteristics: Colonies on MEA flat, with dense or fluffy, sparse to moderate aerial mycelium,
margins regular or scalloped; surface ivory white, dingy white to pale yellow, straw, cream to light grey,
smoke grey, white with center green olive and brown hyphal stripes, creamy white with or without
concentric rings of olivaceous mycelium, or white to grey with wet conidial masses olive green to black;
reverse pale grey, greyish white with olivaceous margins, smoke grey with olivaceous grey margins,
golden yellow to slightly darker, yellowish grey with concentric rings, yellow with dark grey concentric
rings, straw with dark brown concentric rings, or middle dark grey and yellow to medium brown towards
the rim.
Optimal media and cultivation conditions: MEA, OA, PDA and PNA at 25 °C under near-ultraviolet
light.
Distribution: North America, Asia, Australia and Europe.
Hosts: Castanea spp., Chrysolepis chrysophylla, Lithocarpus densiflorus and Quercus spp (Fagaceae),
Liquidambar styraciflua (Hamamelidaceae), Lindera glauca (Lauraceae), and Pinus tabuliformis
(Pinaceae). Reported from other hosts not verified such as Acer spp. (Aceraceae), Carya spp.
(Juglandaceae) and Fraxinus spp. (Oleaceae).
Disease symptoms: Leaf spots, necrosis and death. Tubakia iowensis also causes petiole necrosis and
death of whole leaves on bur oak (bur oak blight).
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Notes: Tubakia was recently revised by Braun et al. (2018), resulting in the introduction of five new
genera to accommodate species previously placed in Tubakia, i.e. Apiognomonioides,
Involutiscutellula, Oblongisporothyrium, Paratubakia and Saprothyrium. All these genera together
with Racheliella and Sphaerosporithyrium, both genera also described during the revision of the genus
Tubakia, were accommodated in the new family Tubakiaceae (Braun et al. 2018). Presently, 16
species are accepted in the genus based on molecular data (ITS, tef1 and tub2 sequences). These
species may form different types of asexual morphs, being the punctiform conidiomata (pycnothyria)
composed of convex scutella with radiating threads of cells fixed to the substratum by a central
columella the most common and characteristic structure formed. Other asexual morphs include
sporodochial conidiomata and crustose or pustulate pycnidioid conidiomata. The conidia are globose
to broad ellipsoid-obovoid, sometimes subcylindrical or somewhat irregular, aseptate, hyaline,
subhyaline to pigmented. Tubakia suttoniana is the only species that produces sexual morph
characterised by ostiolate ascomata, unitunicate asci with two refractive conoid structures in the ascal
apex, and one-celled, hyaline ascospores.
Species of Tubakia are endophytes and/or pathogens in leaves and twigs of many tree species,
causing distinct leaf lesions in different hosts including oak (Quercus spp.), chestnut (Castanea spp.)
and other hardwood species. Moreover, T. iowensis is also capable of causing petiole necrosis and
death of whole leaves on bur oak (Q. macrocarpa), sometimes killing nearly every leaf on a
susceptible tree. This disease is known as bur oak blight and is most common in Iowa and Minnesota,
but it has been noted in western and southern Wisconsin, northern Illinois, northeast Kansas, eastern
Nebraska, and eastern South Dakota, with isolated groups of affected trees in counties of Illinois and
Missouri that border Iowa (Harrington & McNew 2016).
References: Harrington et al. 2012 (morphology and pathogenicity), Harrington & McNew 2016
(pathogenicity, bur oak blight), Harrington & McNew 2018 (morphology and phylogeny), Braun et al.
2018 (morphology, pathogenicity and phylogeny).
Authors: Y. Marin-Felix & P.W. Crous
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Zasmidium Fr., Summa veg. Scand., Sectio Post. (Stockholm): 407. 1849. Fig. 83.
Synonyms: Periconiella Sacc., Atti Ist. Veneto Sci. Lett. Arti 3: 727. 1885.
Biharia Thirum. & Mishra, Sydowia 7: 79. 1953.
Stenellopsis B. Huguenin, Bull. Trimestriel Soc. Mycol. France 81: 695. 1966.
Verrucispora D.E. Shaw & Alcorn, Proc. Linn. Soc. New South Wales 92: 171. 1967. (nom. illegit., Art.
53.1).
Verrucisporota D.E. Shaw & Alcorn, Austral. Syst. Bot. 6: 273. 1993.
Classification: Dothideomycetes, Dothideomycetidae, Capnodiales, Mycosphaerellaceae.
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Type species: Zasmidium cellare (Pers.) Fr., basionym: Racodium cellare Pers. Neotype designated by
Videira et al. (2017): CBS 146.36 (duplicate cultures ATCC 36951 = IFO 4862 = IMI 044943 = LCP
52.402 = LSHB BB274 = MUCL 10089).
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DNA barcode (genus): LSU.
DNA barcodes (species): ITS, rpb2, act, tef1 and tub2. Table 22. Fig. 84.
Ascomata pseudothecial, amphigenous or epiphyllous, dark brown or black, globose, single to
aggregated; necks rarely perceptible, usually a paler coloured circular area, composed of convergent
yellow hyphae; ascomatal wall composed of 2–3 layers of cells of textura angularis. Asci bitunicate,
fasciculate, subsessile, obpyriform, obovoid to ellipsoidal, obclavate to fusoid-ellipsoidal, saccate or
clavate to cylindrical, aparaphysate, 8-spored. Ascospores 2–3-seriate to multiseriate, hyaline, smoothwalled, guttulate, without sheath, fusoid to ellipsoidal with obtuse ends, straight to slightly curved,
uniseptate, constricted or not at the septum, widest in the middle of apical cell. In plant pathogenic
species, mycelium mostly immersed as well as superficial, rarely only immersed; hyphae branched,
septate, hyaline or almost so to pigmented, pale olivaceous to brown, wall thin to somewhat
thickened, immersed hyphae smooth-walled or almost so to faintly rough, external hyphae distinctly
verruculose to verrucose (in culture immersed hyphae usually smooth-walled or almost so, aerial
hyphae verruculose). Stromata lacking to well-developed, pigmented. Conidiophores solitary, arising
from superficial hyphae, lateral, occasionally terminal, in vivo (in plant pathogenic taxa) sometimes
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also fasciculate, arising from internal hyphae or stromata, semimacronematous to macronematous, in
culture occasionally micronematous, cylindrical, filiform, subuliform, straight to strongly geniculatesinuous, mostly unbranched, aseptate, i.e. reduced to conidiogenous cells, to pluriseptate, subhyaline
to pigmented, pale olivaceous to medium dark brown, wall thin to somewhat thickened, smooth to
verruculose. Conidiogenous cells integrated, terminal, occasionally intercalary, rarely pleurogenous,
or conidiophores reduced to conidiogenous cells, mostly polyblastic, sympodial, with conspicuous,
somewhat thickened and darkened-refractive, planate loci. Conidia solitary or catenate, in simple or
branched acropetal chains, shape and size variable, ranging from amero- to scolecosporous, aseptate to
transversely plurieuseptate, subhyaline to pigmented, pale olivaceous to brown, wall thin to somewhat
thickened, smooth or almost so to usually distinctly verruculose (in plant pathogenic species without
superficial mycelium always verruculose); hila somewhat thickened and darkened-refractive, planate,
conidial secession schizolytic (asexual morph description adapted from Braun et al. 2013).
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Culture characteristics: Colonies slow growing, with sparse to moderate aerial mycelium, rarely with
aerial mycelium absent, sometimes with mucoid exudate, margins smooth and regular, lobate or
feathery. On MEA olivaceous, olivaceous green with margins whitish, brown olivaceous, olivaceous
grey, iron mouse grey, grey, cream, yellowish brown, vinaceous buff to olivaceous buff, dark brown, or
dark brown with margins grey; reverse pale or dark grey olivaceous, olivaceous black, mouse grey, irongrey, greenish black, pale orange, isabelline, buff, dark brown, or brown vinaceous. On PDA pale white
with margins pale olivaceous grey, pale mouse grey, olivaceous grey, iron-grey, iron-grey with patches
orange, or brown olivaceous; reverse olivaceous grey, iron-grey, iron-grey with patches orange, or
isabelline. On OA mouse grey, olivaceous grey, smoke grey with margins olivaceous grey, iron-grey, or
iron-grey with broad margins of orange; reverse olivaceous, olivaceous grey, dark mouse grey, or irongrey.
Optimal media and cultivation conditions: MEA, OA, PDA and SNA at 25 °C under near-ultraviolet
light.
Distribution: Worldwide.
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Hosts: Wide range of hosts belonging to 25 different families, including Alocasia odora and Anthurium
sp. (Araceae), Aporosa villosa (Euphorbiaceae), Brabejum stellatifolium, Grevillea spp. and Hakea
undulata (Proteaceae), Citrus spp. (Rutaceae), Cyathea delgadii (Cyatheaceae), Dasypogon sp.
(Dasypogonaceae), Daviesia latifolia (Fabaceae), Elaeocarpus kirtonii (Elaeocarpaceae), Eucalyptus
spp. (Myrtaceae), Gahnia sieberiana (Cyperaceae), Geniostoma rupestre (Loganiaceae), Itea parvifolia
(Escalloniaceae), Lonicera japonica (Caprifoliaceae), Maclura cochinchinensis (Moraceae), Malus spp.
(Rosaceae), Pittosporum tenuifolium (Pittosporaceae), Podocarpus sp. (Podocarpaceae), Pseudotsuga
menziesii and Tsuga heterophylla (Pinaceae), Restio subverticillatus (Restionaceae), Rothmannia
engleriana (Rubiaceae), Sasa sp. (Poaceae), Scaevola taccada (Goodeniaceae), Schinus terebinthifolius
(Anacardiaceae), Strelitzia sp. (Strelitziaceae), and Syzygium cordatum (Myrtaceae).
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Disease symptoms: Causing various lesions, ranging from yellowish discolorations to distinct leaf spots.
Also associated with sooty blotch and flyspeck diseases. Also isolated from wall in wine cellar.
Notes: The genus Zasmidium is morphologically similar to Stenella, producing thickened and
darkened conidiogenous loci and hila (Braun et al. 2013). However, these genera differ in the conidial
hila and scars, being flat in Stenella and planate and somewhat thickened, darkened in Zasmidium.
Moreover, Stenella belongs to Teratosphaeriaceae while Zasmidium is located within
Mycosphaerellaceae (Arzanlou et al. 2007, Quaedvlieg et al. 2013, Videira et al. 2017).
A recent phylogenetic analysis based on LSU, ITS and rpb2 showed that species belonging to other
genera, i.e. Mycosphaerella, Parastenella, Periconiella, Ramichloridium, Rasutoria, Stenella and
Verrucisporota, were located in the monophyletic clade representing the genus Zasmidium (Videira et
al. 2017). Therefore, 12 new combinations were introduced, and the genera Periconiella and
Verrucisporota reduced to synonymy with Zasmidium. Based on our phylogenetic analysis, 48 species
are accepted in the genus together with one new species described here, and a new combination based
on Ramichloridium ducassei.
The type species of the genus, Z. cellare, has been isolated from wine cellars in Europe and
America, while the other species of the genus are associated to plants as saprobic or mostly biotrophic,
usually foliicolous, symptomless or causing various lesions, ranging from yellowish discolorations to
distinct leaf spots. Zasmidium spp. are pathogens of a wide range of hosts such as Z. biverticillatum
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and Z. musigenum, which cause tropical speckle disease on members of Musaceae (Stahel 1937, Jones
2000), and Z. fructicola and Z. fructigenum, both pathogens of Citrus causing a disease known as
citrus greasy spot (Huang et al. 2015).
References: Arzanlou et al. 2007, Videira et al. 2017 (morphology and phylogeny), Braun et al. 2013
(morphology).
Description and illustration: Shivas et al. (2011).
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Zasmidium ducassei (R.G. Shivas et al.) Y. Marín & Crous, comb. nov. MycoBank MB829646.
Basionym: Ramichloridium ducassei R.G. Shivas et al., Australas. Pl. Path. 40: 63. 2010.
Typus: Australia, Queensland, Daintree, on leaves of Musa acuminata × balbisiana (Musaceae), 14
Apr. 2010, M. Berridge & K.R.E. Grice (holotype BRIP 53367, culture ex-type BRIP 53367).
Additional material examined: Malaysia, on leaves of Musa sp., 2016, P.W. Crous, CPC 32929.
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Notes: This species was initially introduced as R. ducassei to accommodate some isolates associated
with a severe leaf speckle disease of Ducasse banana (Musa acuminata × babisiana cv. Pisang awak)
in northern Queensland (Shivas et al. 2011). The authors noticed that this species was similar to
Zasmidium in having pigmented conidiophores with integrated conidiogenous cells that sympodially
proliferate near the apex, with slightly thickened and refractive scars and aseptate, subhyaline conidia
also with slightly thickened and refractive hila. However, it was classified in Ramichloridium in
preference to Zasmidium because, at the time, Zasmidium was a paraphyletic genus in the
Mycosphaerellaceae. In our phylogenetic analysis based on the combined dataset, the ex-type strain of
this species was located in the well-supported clade (100 % BS / 1 PP) representing Zasmidium, and
therefore a new combination Z. ducassei is proposed. Moreover, additional isolates belonging to this
species were obtained from the same host genus, but from different locale, Malaysia.
Zasmidium thailandicum Crous, sp. nov. MycoBank MB829647. Fig. 85.
D
Etymology: Named reflects the country from where it was collected, Thailand.
EP
TE
On SNA. Conidiophores solitary, arising from superficial hyphae, subcylindrical, pale brown, 1–3septate, unbranched or branched below, 20–100 × (1.5–)2 m. Conidiogenous cells subhyaline,
smooth-walled, subcylindrical, apical and intercalary, apical part with well-defined rachis bearing
minute (0.5 m diam) slightly darkened scars, 10–30 × 1.5–2 m. Ramoconidia fusoid to obclavate,
hyaline, smooth-walled, aseptate, guttulate, 8–12(–17) × 2.5–3 m. Conidia solitary, hyaline, smoothwalled, guttulate, aseptate, ellipsoid, apex obtuse, base protruding, truncate, 0.5–1 m diam, (3–)4–
4.5(–5) × (2–)2.5 m.
AC
C
Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium and smooth,
lobate margins, reaching 20 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface pale mouse
grey, reverse mouse grey.
Typus: Thailand, Rachaburi province, Bangkok, on leaves of Musa sp. (Musaceae), 2010, P.W.
Crous, HPC 2158 (holotype CBS H-23850, culture ex-type CBS 145027 = CPC 33960).
Notes: Zasmidium thailandicum is closely related to Z. ducassei. Moreover, both species have been
reported from the same host genus, Musa, causing leaf spots on banana leaves. However, these species
can be easily distinguished by the length of their conidia [5–10 m in Z. ducassei vs. (3–)4–4.5(–5)
m in Z. thailandicum].
Authors: P.W. Crous, J.Z. Groenewald, J.J. Luangsa-ard & Y. Marin-Felix
ACKNOWLEDGEMENTS
Sincere thanks are due to the curators Tara Rintoul (DAOM) and Bevan Weir (ICMP and PDD). We also thank the MycoBank
curator Konstanze Bench, and the technical staff at the Westerdijk Institute, Arien van Iperen (cultures and deposit of herbarium
samples), Mieke Starink-Willemse (DNA isolation and sequencing) and Trix Merkx (deposit of isolates) for their invaluable
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assistance. Jacqueline Edwards thanks Robyn Brett, VPRI curatorial assistant, for maintaining and culturing the VPRI specimens
and Tonya Wiechel for extracting DNA and performing PCR on the VPRI specimens. Sampling in Maryland was supported by
start-up funds from the University of Alabama at Birmingham to S.A. Krueger-Hadfield. We are thankful to Dr. Paul Kirk for
helpful advice regarding the treatment of the invalid name Seiridium cupressi (Guba) Boesew. The study of the genus Alternaria
was supported by the Spanish Ministerio de Economía y Competitividad, Grant CGL2017-88094-P.
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Table 1. DNA barcodes of accepted Allophoma spp.
Isolates1
GenBank accession numbers2
LSU
rpb2
LN907376
LT593058
Allophoma cylindrispora
CBS 142453T
ITS
LT592920
Al. hayatii
CBS 142859T
KY684812
KY684814
CBS 142860
KY684813
Al. labilis
CBS 124.93
Al. minor
References
RI
PT
Species
Valenzuela-Lopez et al. (2018)
MF095108
KY684816
Babaahmadi et al. (2018)
KY684815
MF095109
KY684817
Babaahmadi et al. (2018)
GU237765
GU238091
KT389552
GU237619
Aveskamp et al. (2010), Chen et al. (2015)
CBS 325.82T
GU237831
GU238107
KT389553
GU237632
Aveskamp et al. (2010), Chen et al. (2015)
Al. nicaraguensis
CBS 506.91T
GU237876
GU238058
KT389551
GU237596
Aveskamp et al. (2010), Chen et al. (2015)
Al. oligotrophica
CGMCC 3.18114T
KY742040
KY742194
KY742128
KY742282
Chen et al. (2017)
CGMCC 3.18115
KY742041
KY742195
KY742129
KY742283
Chen et al. (2017)
GU237816
GU238129
KT389554
GU237644
Aveskamp et al. (2010), Chen et al. (2015)
CBS 108.93
GU237921
GU238130
KT389555
GU237645
Aveskamp et al. (2010), Chen et al. (2015)
CGMCC 3.19245T
MK088573
MK088580
MK088587
MK088594
Present study
LC12181
MK088569
MK088576
MK088583
MK088590
Present study
LC12182
MK088570
MK088577
MK088584
MK088591
Present study
LC12183
MK088571
MK088578
MK088585
MK088592
Present study
LC12184
MK088572
MK088579
MK088586
MK088593
Present study
GU237864
GU238149
KT389556
GU237663
Aveskamp et al. (2010), Chen et al. (2015)
FJ427084
GU238159
KT389557
FJ427188
Aveskamp et al. (2010), Chen et al. (2015)
KT389473
KT389690
KT389558
KT389767
Aveskamp et al. (2010), Chen et al. (2015)
Al. tropica
CBS 436.75IsoT
Al. zantedeschiae
CBS 131.93
CBS 229.32
1
M
AN
U
D
TE
EP
Al. pterospermicola
CBS 268.93
AC
C
Al. piperis
ET
SC
tub2
LT592989
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbiological Culture Collection Center, Beijing, China; LC:
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C
EP
TE
D
M
AN
U
SC
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PT
Dr Lei Cai's personal culture collection, housed at CAS, China. T, ET and IsoT indicate ex-type,ex-epitype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; rpb2: partial DNA-directed RNA polymerase II second
largest subunit gene; tub2: partial β-tubulin gene.
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Table 2. DNA barcodes of accepted Alternaria spp.
A. alternata
A. anagallidis
A. anigozanthi
A. anodae
A. aragakii
A. arborescens
A. arbusti
A. argyranthemi
A. argyroxiphii
A. armoraciae
A. arrhenatheri
A. aspera
A. atra
A. avenicola
A. axiaeriisporifera
A. azadirachtae
A. bataticola
A. betae-kenyensis
Alternaria
Porri
Eureka
Porri
Porri
Alternaria
Infectoriae
CBS 916.96T
CBS 117128
CBS 121920T
PPRI 12376
CBS 594.93T
CBS 102605T
CBS 596.93T
CBS 116530T
CBS 117222T
CBS 118702T
LEP 140372T
CBS 115269T
CBS 195.67T
CBS 121459T
CBS 118715T
CBS 116444T
CBS 531.63T
CBS 118810T
AF347031
KJ718106
KC584180
KJ718110
KJ718111
AF347033
JQ693644
KC584181
KJ718112
KC584182
JQ693677
KC584242
AF229486
KC584183
KC584184
KJ718115
KJ718117
KP124419
AC
C
Porri
Chalastospora
Pseudoalternaria
Pseudoulocladium
Ulocladioides
Panax
Gypsophilae
Porri
Porri
Alternaria
AY278808
JQ646338
KC584097
KJ717963
KJ717964
AY278810
JQ646365
KC584098
JQ646350
KC584099
JQ693635
KC584166
KC584167
KC584100
KC584101
KJ717967
JQ646349
KP124270
RI
PT
CBS 534.83T
CBS 541.94T
FMR 17111T
CBS 117221
CBS 577.94T
CBS 107.28T
CBS 118809T
FMR 16476T
CBS 124392
CBS 105.51T
CBS 126989T
CBS 119396T
M
AN
U
Chalastospora
Porri
Infectoriae
Porri
Porri
Porri
Alternaria
Pseudoalternaria
Althernantherae
Porri
Ulocladium
Infectoriae
gapdh
KC584154
KJ717952
LR133965
KJ717953
JQ646356
KJ717954
KP124154
LR133900
KC584096
KJ717959
AY278815
JQ646289
EP
Alternaria abundans
A. acalyphicola
A. aconidiophora
A. agerati
A. agripestis
A. allii
A. alstroemeriae
A. altcampina
A. alternantherae
A. alternariacida
A. alternariae
A. alternarina
ITS
JN383485
KJ718097
LR133931
KJ718098
KJ718099
KJ718100
KP124297
LR133895
KC584179
KJ718105
AF229485
JQ693648
GenBank accession numbers2
rpb2
tef1
ATPase
KC584448 KC584707 JQ671802
–
KJ718271
KJ718446
LR133967 LR133968 LR133969
–
KJ718272
KJ718447
–
KJ718273
KJ718448
–
KJ718274
KJ718449
KP124765 KP125072 –
–
–
LR133906
KC584374 KC584633 –
–
KJ718279
KJ718454
KC584470 KC584730 –
JQ905199
LR134367 JQ671817
SC
Isolates1
D
Section
TE
Species
KC584375
KJ718280
KC584376
KJ718284
KJ718285
KC584377
LR134184
KC584378
KJ718286
KC584379
KC584634
EU130544
KC584635
KJ718458
KJ718459
KC584636
–
–
–
–
–
–
–
JQ671940
KC584637
KJ718460
KC584638
–
–
–
–
LR134098
JQ693603
KC584474
KC584475
KC584380
KC584381
KJ718289
KJ718291
KP124888
KC584734
KC584735
KC584639
KC584640
KJ718463
KJ718465
KP125197
–
–
–
–
–
–
–
References
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2014)
Present study
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2015)
Present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Poursafar et al. (2018), Geng et al. (unpubl. data),
present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013), present study
Poursafar et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2015)
ACCEPTED MANUSCRIPT
–
–
–
–
–
–
–
–
–
KC584442
LR134194
KP124889
KC584492
KJ718301
LR134181
KC584384
KC584479
KC584385
KC584467
KC584386
KJ718305
KJ718308
KJ718309
KJ718313
KJ718314
LR135746
KC584700
LR134262
KP125198
KC584752
KJ718475
LR134245
KC584643
KC584739
KC584644
KC584726
KC584645
KJ718479
KJ718482
KJ718483
KJ718487
KJ718488
LR135745
KP711386
KC584699
KC584741
KC584646
KC584647
KC584710
LR133929
KJ718489
KC584648
KJ718491
KJ718492
KY412558
LR134099
KY412557
–
–
AY562398
KC584172
KC584107
KC584108
KC584156
LR133927
KJ717991
KC584109
KJ717993
KJ717994
KC584441
KC584481
KC584387
KC584388
KC584451
LR133926
KJ718315
KC584389
KJ718317
KJ718318
M
AN
U
SC
KJ718300
KC584491
KC584461
KC584476
KC584382
KC584478
KC584383
RI
PT
KJ718474
KC584751
KC584720
KC584736
KC584641
KC584738
KC584642
AY562405
FJ357305
AY278831
KC584168
KC584102
KC584170
KC584103
KM821537
KC584148
KM821538
JQ646305
KC584178
KJ717977
JQ646285
KC584104
KC584171
KC584105
AY278826
KC584106
KJ717981
KJ717984
KJ717985
KJ717989
KJ717990
LR135747
D
KJ718126
FJ357317
AY278844
KC584243
KC584185
KC584244
JX499031
JQ317188
FJ839608
KM821536
KP124420
KC584250
KJ718127
JQ693645
KC584186
KC584245
KC584187
AY278839
KC584188
KJ718131
KJ718134
KJ718135
KJ718139
KJ718140
LR135744
KP711383
JN383482
AF229488
AF229457
KC584189
KC584231
LR133924
KJ718141
KC584190
KJ718143
KJ718144
TE
Ulocladioides
Brassicicola
Pseudoalternaria
Chalastospora
Infectoriae
Alternaria
Infectoriae
Porri
Infectoriae
Panax
Ulocladioides
Ulocladium
Nimbya
Radicina
Porri
Porri
Porri
Porri
Porri
Infectoriae
Infectoriae
Chalastospora
Pseudoulocladium
Cheiranthus
Phragmosporae
Embellisia
Radicina
Porri
Sonchi
Porri
Porri
CBS 117364T
DAOM 231361T
CBS 478.90T
CBS 197.67T
CBS 116528
CBS 121493T
CBS 118699
CNU 111118T
CBS 121331T
CBS 118485T
CBS 107.38T
CBS 177.80T
CBS 224.76T
CBS 119409T
CBS 121545T
CBS 123007T
CBS 504.74
CBS 480.90T
CBS 109381T
CBS 635.80
CBS 117092T
CBS 478.81
CBS 137456T
CBS 116446T
CBS 121923T
MFLUCC 13-0450T
CBS 121340T
CBS 200.67T
CBS 109384
CBS 491.72T
CBS 341.71
FMR 17360T
CBS 102.33T
CBS 116495
CBS 113261T
CBS 103.32T
EP
Porri
Undifilum
Embellisioides
Ulocladium
AC
C
A. blumeae
A. bornmuelleri
A. botryospora
A. botrytis
A. brassicae
A. brassicaepekinensis
A. brassicicola
A. brassicifolii
A. breviramosa
A. broccoli-italicae
A. burnsii
A. caespitosa
A. calendulae
A. californica
A. calycipyricola
A. cantlous
A. capsici-annui
A. caricis
A. carotiincultae
A. carthami
A. carthamicola
A. cassiae
A. catananches
A. centaureae
A. cerasidanica
A. cesenica
A. cetera
A. chartarum
A. cheiranthi
A. chlamydospora
A. chlamydosporigena
A. chlamydosporifera
A. cichorii
A. cinerariae
A. cirsinoxia
A. citrullicola
–
LR134114
–
JQ671813
–
–
–
JQ671780
–
–
–
–
–
–
LR135748
–
LR134101
–
–
–
–
–
–
–
–
–
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Deng et al. (2018)
Woudenberg et al. (2013), present study
Deng et al. (2018), present study
Woudenberg et al. (2015)
Woudenberg et al. (2013), present study
Woudenberg et al. (2014)
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Present study
Liu et al. (2015)
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Present study
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
ACCEPTED MANUSCRIPT
–
–
KC584740
KJ718493
KC584649
KC584711
KC584742
KJ718495
KJ718504
KC584650
KJ718501
KC584743
LR133902
KJ718005
JQ646294
JN383469
KJ718077
KC584113
KJ718012
FJ357300
KC584480
KC584390
KC584452
KC584482
KJ718320
KJ718329
KC584391
KJ718326
KC584483
LR133901
KY750720
KJ718331
LR134177
KC584454
KJ718424
KC584394
KJ718340
KC584455
–
–
–
KJ718015
KP124276
KC584114
KC584115
KC584155
AY562416
AY278795
KJ718018
JN383471
LR133921
KJ718343
KP124895
KC584395
KC584396
KC584449
KC584397
KC584398
KJ718346
KC584456
LR133923
KJ718518
KP125204
KC584654
KC584655
KC584708
KC584656
KC584657
KJ718521
KC584715
LR133922
–
–
–
–
JQ646295
KC584116
KC584117
JQ646312
JQ646291
LR134172
KC584399
KC584400
KP124900
LR134180
LR134370
KC584658
KC584659
KP125209
LR134249
JQ671823
M
AN
U
–
JQ671824
–
–
–
–
–
–
–
RI
PT
SC
AY762950
KJ717995
AY562401
FJ348227
KC584173
KJ717997
KJ718003
KC584110
KJ718000
AY562418
LR133899
D
KC584246
KJ718145
FJ266475
AF348226
KC584247
KJ718147
KJ718156
KC584191
KJ718153
FJ266483
LR133898
KY703616
KJ718158
JQ693653
JN383488
KJ718249
KC584194
KJ718167
FJ357312
MG828864
KJ718170
KC146356
KC584195
KC584196
KC584230
JQ693661
AY278833
KJ718173
JN383490
LR133920
KY769657
JQ693654
KC584197
KC584198
KP124430
JQ693650
TE
Porri
Dianthicola
Porri
Phragmosporae
Alternaria
Porri
Alternaria
Dianthicola
Gypsophilae
Embellisia
Panax
Infectoriae
Euphorbiicola
Eureka
Infectoriae
Infectoriae
Infectoriae
Alternaria
Eureka
Alternaria
Infectoriae
CBS 120006T
CBS 137457T
CBS 196.86T
CBS 132.89
CBS 104.31T
CBS 110.38T
CBS 364.67
CBS 121329T
CBS 116114T
CBS 483.81
FMR 16901T
MFLUCC 15-0466T
CBS 111.38T
CBS 119398T
CBS 476.90T
CBS 110799T
CBS 116491
CBS 200.74T
CBS 766.79
KUMCC 17-0263T
CBS 116117T
CBS 489.92T
CBS 109159T
CBS 119674T
CBS 339.71
CBS 121339
CBS 197.86T
CBS 119410
CBS 193.86T
FMR 17110T
MFLUCC 13-0456T
CBS 119401T
CBS 632.93
CBS 118701T
CBS 104.32T
CBS 119400T
EP
Pseudoulocladium
Porri
Infectoriae
Brassicicola
Ulocladioides
Porri
Porri
Eureka
Porri
Ulocladioides
Infectoriae
Infectoriae
Porri
Infectoriae
AC
C
A. concatenata
A. conidiophora
A. conjuncta
A. conoidea
A. consortialis
A. crassa
A. cyamopsidis
A. cumini
A. cucumerina
A. cucurbitae
A. curvata
A. dactylidicola
A. dauci
A. daucicaulis
A. dennisii
A. deserticola
A. dianthicola
A. dichondrae
A. didymospora
A. doliconidium
A. echinaceae
A. eichhorniae
A. elegans
A. ellipsoidea
A. embellisia
A. eryngii
A. ethzedia
A. euphorbiicola
A. eureka
A. fimeti
A. forlicesenensis
A. frumenti
A. gaisen
A. geniostomatis
A. gossypina
A. graminicola
–
KJ718506
LR134241
KC584713
KJ718595
KC584653
KJ718515
KC584714
LR133905
–
–
JQ671822
–
–
–
–
JQ671796
–
–
–
–
–
–
–
JQ671805
–
–
LR133925
–
–
–
JQ671819
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013), Poursafar et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Present study
Thambugala et al. (2017)
Woudenberg et al. (2014)
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013), Deng et al. (2018)
Wanasinghe et al. (2018)
Woudenberg et al. (2014)
Woudenberg et al. (2015)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013), Poursafar et al. (2018)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Present study
Thambugala et al. (2017)
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2015)
Poursafar et al. (2018), present study
ACCEPTED MANUSCRIPT
KC584120
KC584176
JQ646283
JQ646284
JQ646293
KC584158
JQ646287
KC584159
AY278793
LR133938
JQ646297
KJ718020
KP124284
KC584121
KP124287
KJ718022
KC584122
MF033826
LR133998
LR133997
KJ649618
LR133908
KC584160
KC584164
KC584123
JQ646329
KJ718024
AY278811
JN383473
KC584124
JQ646278
LR134077
–
–
–
–
RI
PT
JX101649
KC584248
JQ693641
JQ693642
JQ693652
KC584233
JQ693658
KC584234
AF347034
LR133930
JQ693656
KJ718175
KP124435
KC584201
KP124438
KJ718177
KC584202
MF033843
LR133970
LR133995
KJ443262
LR133907
KC584235
KC584240
KC584203
KJ718178
KJ718180
AY278835
JN383492
KC584204
JQ693638
LR134074
EU130547
KC584660
EU130548
KC584748
LR134243
LR134371
LR134199
KC584716
LR134250
KC584717
KC584662
SC
Soda
Infectoriae
Eureka
Teretispora
Phragmosporae
Euphorbiicola
Porri
Alternaria
Embellisioides
Porri
Chalastospora
CBS 208.86T
CBS 123376T
CBS 119402T
CBS 121458T
CBS 119404T
CBS 416.71T
CBS 121330T
CBS 536.83T
CBS 210.86T
FMR 16477 T
CBS 119406T
CBS 219.79T
CBS 118486T
CBS 118390
CBS 133751T
CBS 133855T
CBS 119673T
IRAN 16888FT
FMR 17061
FMR 17372
CBS 137525T
FMR 17004T
CBS 477.90T
CBS 421.65T
CBS 481.81T
CBS 483.90T
CBS 105.41T
CBS 540.94
CBS 115266T
CBS 117228T
CBS 135.31
FMR 17369
–
KJ718414
KC584401
MG82924
7
KC584403
KC584488
LR134179
LR134175
LR134174
KC584457
LR134185
KC584458
KC584404
–
–
JQ671811
JQ671812
JQ671821
–
JQ671815
–
LR134170
KJ718348
KP124905
KC584405
KP124908
KJ718350
KC584406
–
–
JQ671804
LR133966
JQ671826
KJ718523
KP125214
KC584663
KP125217
KJ718525
KC584664
–
–
–
–
–
–
–
–
–
–
–
–
MF033860
LR134001
LR133999
KJ443176
LR133911
KC584459
KC584472
KC584407
KJ718351
KJ718353
KC584409
KC584460
KC584410
KJ443219
LR133912
KC584718
KC584732
KC584665
KJ718526
KJ718528
KC584667
KC584719
KC584668
–
–
–
–
M
AN
U
A. kulundii
A. lawrencei
A. leptinellae
A. leucanthemi
A. limaciformis
A. limicola
A. linariae
A. longipes
A. lolii
A. macrospora
A. malorum
JQ646341
KC584118
D
Ulocladioides
Infectoriae
Infectoriae
Infectoriae
Embellisioides
Infectoriae
Cheiranthus
Infectoriae
Pseudoalternaria
Infectoriae
Porri
Alternaria
Japonicae
Alternaria
Porri
Gypsophilae
Pseudoalternaria
KJ718239
KC584199
MG828866
TE
A. helianthiinficiens
A. heterospora
A. hordeiaustralica
A. hordeicola
A. humuli
A. hyacinthi
A. incomplexa
A. indefessa
A. infectoria
A. inflata
A. intercepta
A. ipomoeae
A. iridiaustralis
A. japonica
A. jacinthicola
A. jesenskae
A. juxtiseptata
A. kordkuyana
CBS 109158T
CBS 107.41T
MFLUCC 17-0783T
EP
Porri
Gypsophilae
Infectoriae
AC
C
A. grandis
A. gypsophilae
A. hampshirensis
–
–
LR133914
–
–
JQ671798
–
–
–
–
–
JQ671800
LR134029
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Wanasinghe et al., (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Poursafar et al. (2018), present study
Poursafar et al. (2018), present study
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Poursafar et al. (2018), present study
Woudenberg et al. (2013)
Woudenberg et al. (2013), Poursafar et al. (2018)
Present study
Poursafar et al. (2018), present study
Woudenberg et al. (2014)
Woudenberg et al. (2015)
Woudenberg et al. (2013)
Woudenberg et al. (2015)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Poursafar et al. (2018)
Present study
Present study
Grum-Grzhimaylo et al. (2014)
Present study
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Poursafar et al. (2018)
Present study
ACCEPTED MANUSCRIPT
A. merytae
A. metachromatica
Infectoriae
Infectoriae
CBS 119403T
CBS 553.94T
JQ693651
JQ693660
JQ646292
AY562404
LR134119
JQ905189
LR134198
FJ214931
A. mimicula
A. molesta
A. montanica
A. montsantina
A. mouchaccae
A. multiformis
A. multirostrata
A. murispora
A. neoipomoeae
A. nepalensis
A. nitrimali
A. nobilis
A. novae-guineensis
A. novae-zelandiae
A. obclavata
A. obovoidea
A. obtecta
A. oregonensis
A. oudemansii
A. panax
A. papavericola
A. paralinicola
A. passiflorae
A. parvicaespitosa
A. penicillata
A. perpunctulata
A. petroselini
A. petuchovskii
A. peucedani
A. photistica
A. phragmospora
A. pipionipisi
A. planifunda
Brassicicola
Phragmosporae
Porri
Infectoriae
Phragmosporae
Ulocladioides
Porri
Infectoriae
Porri
Japonicae
Porri
Gypsophilae
Porri
Infectoriae
Chalastospora
Ulocladioides
Porri
Infectoriae
Ulocladium
Panax
Crivellia
Porri
Porri
Pseudoalternaria
Crivellia
Althernantherae
Radicina
Soda
CBS 118696T
CBS 548.81T
CBS 121343T
FMR 17060T
CBS 119671T
CBS 102060T
CBS 712.68T
MFLU 14-0758T
PPRI 11845T
CBS 118700T
CBS 109163T
CBS 116490
CBS 116120T
CBS 119405T
CBS 124120T
CBS 101229
CBS 117367
CBS 542.94T
CBS 114.07T
CBS 482.81
CBS 116606T
CBS 116652T
CBS 113.38
LEP 014858T
CBS 116608T
CBS 115267T
CBS 112.41T
CBS 137517T
CNU 111485T
CBS 212.86T
CBS 274.70T
CBS 116115T
CBS 537.83T
FJ266477
KC584205
KJ718194
LR133913
KC584206
FJ266486
KJ718195
NR_137964
KJ718198
KC584207
KJ718201
KC584208
KJ718202
JQ693655
KC584225
FJ266487
KJ718204
FJ266478
FJ266488
KC584209
FJ357310
KJ718206
KJ718207
MF033859
FJ357311
KC584210
KC584211
KJ443254
KF728231
KC584212
JN383493
KJ718214
FJ357315
AY562415
KC584125
KJ718033
LR133915
AY562399
KC584174
JQ646362
KC584411
KC584412
KJ718367
LR133918
KC584413
KC584484
KJ718368
KC584669
KC584670
KJ718541
LR133919
KC584671
KC584744
EU130546
–
–
–
KJ718036
KC584126
JQ646358
KC584127
KJ718039
JQ646296
KC584149
FJ266498
KJ718041
FJ266491
KC584175
KC584128
FJ357298
KJ718043
JQ646353
MF033842
FJ357299
KC584129
KC584130
KJ649616
KF889361
KC584131
JN383474
KJ718049
FJ357303
KJ718371
KC584414
KJ718374
KC584415
KJ718375
LR134120
KC584443
KC584485
KJ718377
KC584416
KC584486
KC584417
KC584446
KJ718379
KJ718380
KJ718544
KC584672
KJ718547
KC584673
KJ718548
LR134197
KC584701
KC584745
KJ718550
KC584674
KC584746
KC584675
KC584705
KJ718552
KJ718553
–
–
KJ908217
KC584440
KC584418
KC584419
KJ443170
KC584698
KC584676
KC584677
KJ443211
–
–
–
–
–
–
–
–
RI
PT
SC
M
AN
U
D
TE
EP
AC
C
Panax
Phragmosporae
Porri
Embellisioides
JQ671820
JQ671809
–
–
KC584420
KC584462
KJ718387
KC584463
KC584678
KC584721
KJ718560
KC584722
LR133916
–
–
–
–
–
–
–
–
–
JQ671825
LR134100
–
–
JQ671827
–
–
–
–
–
JQ671807
–
–
–
Poursafar et al. (2018), present study
Andersen et al. (2009), Poursafar et al. (2018), Geng
et al. (unpubl. data)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Present study
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Ariyawansa and Hyde (unpubl. data)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Poursafar et al. (2018), present study
Woudenberg et al. (2013), present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013), Poursafar et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Poursafar et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Grum-Grzhimaylo et al. (2014)
Deng et al. (2014)
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
ACCEPTED MANUSCRIPT
Brassicicola
–
KY460971
–
–
–
–
LR133897
KJ718053
KC584161
JQ646335
AY562406
LR133935
KC584133
KJ718058
KJ718059
JQ646279
LR134070
LR13403
LR134072
LR134103
JQ646332
KC584135
KC584163
JN383480
JN383481
JQ646363
AY278800
JQ646344
FJ266500
KC584136
KJ718061
KJ649620
KJ718062
KC584137
KJ718063
KC584150
KC584138
KJ718066
KJ718071
KC584140
KC584141
KJ718391
KC584464
KJ718394
KC584422
LR133934
KC584423
KJ718398
KJ718399
KJ718564
KC584723
KJ718567
KC584680
LR133936
KC584681
KJ718571
KJ718572
–
–
–
–
–
–
–
–
–
–
–
–
RI
PT
LR133903
SC
M
AN
U
D
Infectoriae
Porri
Gypsophilae
Nimbya
Nimbya
Nimbya
Porri
Radicina
Porri
Pseudoulocladium
Brassicicola
Porri
Soda
Porri
Dianthicola
Porri
Infectoriae
Radicina
Porri
Porri
KY026587
LR133896
KJ718218
AY278842
KJ718221
JN383483
LR133928
KC584213
KJ718225
KJ718226
JQ646279
LR134076
LR134071
LR134073
LR134102
KJ718229
KC584215
KC584237
JN383499
JN383500
KJ718190
AF229455
KJ718230
FJ266489
KC584216
JF780939
KJ443257
KJ718232
JF780937
KJ718233
KC584226
AF229456
KJ718236
KJ718243
KC584218
KC584219
TE
A. roseogrisea
A. rostellata
A. saponariae
A. scirpicola
A. scirpinfestans
A. scirpivora
A. scorzonerae
A. selini
A. sennae
A. septospora
A. septorioides
A. sesami
A. shukurtuzii
A. sidae
A. simsimi
A. silybi
A. slovaca
A. smyrnii
A. solani
A. solani-nigri
A. soliaridae
A. solidaccana
MFLUCC 13-0346T
FMR 16448T
CBS 116699T
CBS 475.90T
CBS 116696
CBS 119411T
FMR 16900T
CBS 245.67T
CBS 116330T
CBS 215.31T
CBS 121341T
FMR 15720
FMR 17376
FMR 17377
CBS 121921T
CBS 117366T
CBS 116492
CBS 481.90
EGS 49-185T
EGS 50-021T
CBS 103.46
CBS 109382T
CBS 477.81T
CBS 109.38
CBS 106.41T
CBS 115264
CBS 137520T
CBS 117730T
CBS 115265T
CBS 134092T
CBS 567.66T
CBS 109380
CBS 106.21
CBS 113403
CBS 118387T
CBS 118698T
EP
Infectoriae
Chalastospora
Porri
Embellisioides
Porri
Porri
Infectoriae
Radicina
Porri
Porri
Pseudoalternaria
AC
C
A. poaceicola
A. pobletensis
A. porri
A. proteae
A. protenta
A. pseudorostrata
A. pseudoventricosa
A. radicina
A. ranunculi
A. ricini
A. rosae
LR134192
KJ718402
KC584425
KC584469
–
–
KJ718363
KC584426
KJ718403
KC584487
KC584427
KJ718405
KJ443172
KJ718406
KC584428
KJ718407
KC584444
KC584429
KJ718410
KJ718418
KC584431
KC584432
LR134260
KJ718575
KC584683
KC584728
JQ672404
JQ672405
KJ718537
KC584684
EU130543
KC584747
KC584685
KJ718577
KJ443214
KJ718578
KC584686
KJ718579
KC584702
KC584687
KJ718582
KJ718589
KC584689
KC584690
LR133937
–
–
–
JQ671803
LR134004
LR134003
LR134028
LR134104
–
–
JQ671781
JQ671783
JQ671782
–
–
–
–
–
–
–
–
–
–
LR134368
–
–
–
–
–
Thambugala et al. (2017)
Present study
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Poursafar et al. (2018)
Present study
Present study
Present study
Present study
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013), Deng et al. (2018)
Lawrence et al. (2012), Lawrence et al. (unpubl. data)
Lawrence et al. (2012), Lawrence et al. (unpubl. data)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Grum-Grzhimaylo et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013), present study
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
ACCEPTED MANUSCRIPT
KC584142
KJ718079
EU855803
KC584143
AY562419
KC584177
KJ718087
KC584144
KJ718084
KP124294
KC584145
JQ646280
JQ646281
KC584433
KJ718427
KC584489
KC584434
KC584465
KC584490
KJ718435
KC584436
KJ718432
KP124915
KC584437
LR134183
LR134186
KC584691
KJ718598
KC584749
KC584692
KC584724
KC584750
KJ718606
KC584694
KJ718603
KP125224
KC584695
–
JQ671806
JQ671808
A. tropica
A. tumida
A. quercicola
A. vaccariae
A. vaccariicola
A. venezuelensis
A. ventricosa
Porri
Embellisioides
Infectoriae
Gypsophilae
Gypsophilae
Porri
Infectoriae
CBS 631.93T
CBS 539.83T
CBS 141466T
CBS 116533
CBS 118714T
CBS 116121T
CBS 121546T
KJ718261
FJ266481
KX228295
KC584223
KC584224
KJ718263
JQ693649
KJ718088
FJ266493
KX228362
KC584146
KC584147
KJ718090
JQ646290
KJ718436
KC584466
LR134188
KC584438
KC584439
KJ718438
LR134134
KJ718607
KC584725
LR134259
KC584696
KC584697
KJ718609
KY352501
A. viburni
A. zinniae
Infectoriae
Porri
CBS 119407T
CBS 117223
JQ693647
KJ718270
LR134166
KJ718445
LR134200
KJ718616
1
SC
M
AN
U
D
TE
Sonchi
Porri
Ulocladioides
Porri
Embellisia
Ulocladioides
Porri
JQ646288
KJ718096
RI
PT
KC584220
KJ718252
KC584249
KC584221
FJ357316
FJ266490
KJ718260
EU040211
KJ718257
KP124445
KC584222
JQ693657
AY278834
–
–
–
–
Porri
Alternaria
Eureka
Infectoriae
Infectoriae
CBS 119675
CBS 117362T
CBS 121491T
CBS 479.81
CBS 538.83T
CBS 202.67T
CBS 116116T
CBS 121712T
CBS 116331T
CBS 103.30
CBS 119676T
CBS 578.94T
CBS 763.84T
A. sonchi
A. steviae
A. subcucurbitae
A. tagetica
A. tellustris
A. terricola
A. tillandsiae
A. thalictrigena
A. thunbergiae
A. tomato
A. triglochinicola
A. triticimaculans
A. triticina
FJ214942
JQ671794
–
–
–
–
–
–
–
–
LR134115
–
–
–
JQ671818
JQ671816
–
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2013), Deng et al. (2018)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Woudenberg et al. (2015)
Woudenberg et al. (2013)
Poursafar et al. (2018), present study
Andersen et al. (2009), Poursafar et al. (2018),
Present study
Woudenberg et al. (2014)
Woudenberg et al. (2013)
Woudenberg et al. (2016), present study
Woudenberg et al. (2013)
Woudenberg et al. (2013)
Woudenberg et al. (2014)
Poursafar et al. (2018), Fotedar et al. (unpubl. data),
present study
Poursafar et al. (2018), present study
Woudenberg et al. (2014)
AC
C
EP
CBS: Culture collection of the Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CNU: Culture Collection Center of the Chungnam National University; DAOM: Canadian Collection of Fungal Cultures, Ottawa, Canada;
EGS: Personal collection of Dr. E.G. Simmons; FMR: Facultat de Medicina, Universitat Rovira i Virgili, Reus, Spain; IRAN: Fungal Culture Collections of the Iranian Research Institute of Plant Protection; KUMCC, Culture collection of
Kunming Institute of Botany, Kunming, China; LEP: Mycological Herbarium of All-Russian Institute of Plant Protection, Saint Petersburg, Russia; MFLU: Herbarium of Mae Fah Luang University,Chiang Rai, Thailand; MFLUCC: Mae Fah
Luang University Culture Collection, Chiang Ria, Thailand; PPRI: ARC-Plant Protection Research Institute, Roodeplaat, South Africa. T indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial translation
elongation factor 1-alpha gene; ATPase: partial plasma membrane ATPase gene.
ACCEPTED MANUSCRIPT
Table 3. DNA barcodes of accepted Brunneosphaerella spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
GenBank accession numbers2
References
chs
rpb2
tef1
ITS
ET
Brunneosphaerella jonkershoekensis CPC 13902
JN712439 JN712609 MF951441 JN712571 Crous et al. (2011), Videira et al. (2017)
T
B. nitidae
CBS 130595
GU214625 JN712619 MF951442 JN712581 Crous et al. (2009b, 2011), Videira et al.
(2017)
ET
B. protearum
CBS 130597
GU214626 JN712620 MF951443 JN712582 Crous et al. (2009b, 2011), Videira et al.
(2017)
T
B. roupeliae
CBS 144602
MK539950 –
MK540080 –
Present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at the Westerdijk Fungal
Biodiversity Institute. T and ET indicate ex-type and ex-epitype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; chs: partial chitin synthase-1 gene; rpb2: partial DNA-directed RNA polymerase II second
largest subunit gene; tef1: partial translation elongation factor 1-alpha gene.
Species
ACCEPTED MANUSCRIPT
Table 4. DNA barcodes of accepted Elsinoe spp.
References
RI
PT
tef1
KX886831
KX886832
KX886835
KX886836
KX886837
KX886840
KX886841
KX886844
–
KX886845
KX886846
KX886847
KX886850
KX886851
KX886852
KX886853
KX886855
KX886856
KX398203
KX886857
KX886860
KX886861
KX886862
KX886863
KX886864
KX886865
KX886869
KX886870
SC
M
AN
U
D
CBS 510.50
CBS 208.25
CBS 470.62T
CBS 228.64
CBS 511.50T
CBS 220.50T
CBS 141937T
CBS 314.32ET
STE-U 2678T
CBS 113734T
CBS 471.62T
CBS 512.50T
CPC 18528ET
CBS 221.50T
CBS 222.50T
CPC 18535T
CBS 275.76T
CBS 223.50T
DAR 83016T
CBS 472.62T
CPC 18542ET
CBS 124765T
CBS 120084T
CBS 401.63T
CBS 514.50T
CBS 139.25T
CBS 515.50
CBS 473.62T
ITS
KX887185
KX887186
KX887189
KX887190
KX887191
KX887194
KX887195
KX887198
AF227197
KX887199
KX887200
KX887201
KX887204
KX887205
KX887206
KX887207
KX887209
KX887210
KX372292
KX887211
KX887214
KX887215
KX887216
KX887217
KX887218
KX887219
KX887223
KX887224
TE
T
GenBank accession numbers2
rpb2
LSU
KX886949
KX887068
KX886950
KX887069
KX886953
KX887072
KX886954
KX887073
KX886955
KX887074
KX886958
KX887077
KX886959
KX887078
KX886962
KX887081
–
–
KX886963
KX887082
KX886964
KX887083
KX886965
KX887084
–
KX887087
KX886968
KX887088
KX886969
KX887089
KX886970
KX887090
KX886972
KX887092
KX886973
KX887093
–
KX398204
KX886974
–
KX886977
KX887096
KX886978
KX887097
KX886979
KX887098
KX886980
KX887099
KX886981
KX887100
KX886982
KX887101
KX886986
KX887105
KX886987
KX887106
EP
Elsinoe abutilonis
E. ampelina
E. anacardii
E. annonae
E. arachidis
E. arrudai
E. asclepiadea
E. australis
E. banksiae
E. banksiicola
E. barleriicola
E. bidentis
E. brasiliensis
E. caleae
E. centrolobii
E. citricola
E. coryli
E. diospyri
E. eelemani
E. embeliae
E. erythrinae
E. eucalypticola
E. eucalyptorum
E. euphorbiae
E. fagarae
E. fawcettii
E. fici
E. fici-caricae
Isolates1
AC
C
Species
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Swart et al. (2001)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Crous et al. (2016a)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
ACCEPTED MANUSCRIPT
E. flacourtiae
E. freyliniae
E. genipae
E. genipae-americanae
E. glycines
E. hederae
E. ichnocarpi
E. jasminae
E. jasminicola
E. krugii
E. lagoa-santensis
E. ledi
E. lepagei
E. leucopogonis
E. leucospermi
E. lippiae
E. mangiferae
E. mattiroloanum
E. menthae
E. mimosae
E. oleae
CBS 474.62T
CBS 128204T
CBS 342.39T
CBS 516.50T
CBS 389.64ET
CBS 517.50T
CBS 475.62T
CBS 224.50T
CBS 212.63T
CPC 18531ET
CBS 518.50T
CBS 167.33ET
CBS 225.50T
CBS 144439T
CBS 111207T
CBS 166.40T
CBS 226.50T
CBS 287.64
CBS 322.37ET
CPC 19478ET
CBS 227.59T
KX887225
KX887226
KX887227
KX887228
KX887229
KX887231
KX887232
KX887233
KX887234
KX887235
KX887239
KX887240
KX887241
MH327822
KX887242
KX887248
KX887249
KX887250
KX887253
KX887255
KX887256
E. othonnae
CBS 139910T
KR476726
–
MK540083
–
E. perseae
E. phaseoli
E. picconiae
E. piri
E. pitangae
E. poinsettiae
E. pongamiae
E. populi
E. preissianae
E. proteae
CBS 406.34T
CBS 165.31T
CBS 145026T
CBS 163.29
CBS 227.50T
CBS 109333
CBS 402.63ET
CBS 289.64
CBS 142129T
STE-U 1349T
KX887258
KX887263
MK539951
KX887267
KX887269
KX887270
KX887272
KX887273
KY173406
AF097578
KX887021
KX887026
MK540022
KX887030
KX887032
KX887033
KX887035
KX887036
KY173498
–
KX887139
KX887144
MK540081
KX887148
KX887150
KX887151
KX887153
KX887154
–
–
KX886903
KX886908
MK540164
KX886912
KX886914
KX886915
KX886917
KX886918
–
–
D
TE
EP
AC
C
RI
PT
KX886871
KX886872
KX886873
KX886874
KX886875
KX886877
KX886878
KX886879
KX886880
KX886881
KX886885
KX886886
–
MH327897
KX886887
KX886893
KX886894
KX886895
KX886898
KX886900
KX886901
SC
KX887107
KX887108
KX887109
KX887110
KX887111
KX887113
KX887114
KX887115
–
KX887116
KX887120
KX887121
KX887122
MH327885
KX887123
KX887129
KX887130
KX887131
KX887134
KX887136
KX887137
M
AN
U
KX886988
KX886989
KX886990
KX886991
KX886992
KX886994
KX886995
KX886996
KX886997
KX886998
KX887002
KX887003
KX887004
MH327858
KX887005
KX887011
KX887012
KX887013
KX887016
KX887018
KX887019
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Crous et al. (2018)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Crous et al. (2015c),
present study
Fan et al. (2017)
Fan et al. (2017)
Present study
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Fan et al. (2017)
Crous et al. (2016a)
Swart et al. (2001)
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
E. protearum
CBS 113618T
KX887275
KX887038
KX887156
KX886920
Fan et al. (2017)
E. punicae
CPC 19968
KX887276
KX887039
KX887157
KX886921
Fan et al. (2017)
E. quercus-ilicis
CBS 232.61T
KX887277
KX887040
–
KX886922
Fan et al. (2017)
IsoT
KX887278
KX887041
KX887158
KX886923
Fan et al. (2017)
E. randii
CBS 170.38
T
Fan et al. (2017)
E. rhois
CBS 519.50
KX887280
KX887043
KX887160
KX886925
E. ricini
CBS 403.63
KX887281
KX887044
KX887161
KX886926
Fan et al. (2017)
ET
E. rosarum
CBS 212.33
KX887283
KX887046
KX887163
KX886928
Fan et al. (2017)
E. salicina
CPC 17824T
KX887286
KX887049
KX887166
KX886931
Fan et al. (2017)
KX887287
KX887050
KX887167
KX886932
Fan et al. (2017)
E. semecarpi
CBS 477.62T
E. sesseae
CPC 18549
KX887288
KX887051
KX887168
KX886933
Fan et al. (2017)
T
E. sicula
CBS 398.59
KX887289
KX887052
KX887169
KX886934
Fan et al. (2017)
ET
E. solidaginis
CBS 191.37
KX887290
KX887053
KX887170
KX886935
Fan et al. (2017)
T
E. tectificae
CBS 124777
KX887292
KX887055
KX887172
KX886937
Fan et al. (2017)
ET
KX887293
KX887056
KX887173
–
Fan et al. (2017)
E. terminaliae
CBS 343.39
T
E. theae
CBS 228.50
KX887295
KX887058
KX887175
KX886939
Fan et al. (2017)
E. tiliae
CBS 350.73
KX887296
KX887059
KX887176
KX886940
Fan et al. (2017)
ET
E. veneta
CBS 164.29
KX887297
KX887060
KX887177
KX886941
Fan et al. (2017)
E. verbenae
CPC 18561ET
KX887298
KX887061
KX887178
KX886942
Fan et al. (2017)
MK539952
MK540023
MK540082
–
Present study
E. veronicae
CBS 145362T
T
E. violae
CBS 336.35
KX887302
KX887065
KX887182
KX886946
Fan et al. (2017)
T
E. zizyphi
CBS 378.62
KX887303
KX887066
KX887183
KX886947
Fan et al. (2017)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity
Institute; DAR: Plant Pathology Herbarium, New South Wales, Australia; STE-U: Department of Plant Pathology, Stellenbosch University, South Africa. T, ET
and IsoT indicate ex-type, ex-epitype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; rpb2: partial DNA-directed RNA polymerase II
second largest subunit gene; tef1: partial translation elongation factor 1-alpha gene.
ACCEPTED MANUSCRIPT
Table 5. DNA barcodes of accepted Exserohilum spp.
BRIP 11426T
GenBank accession numbers2
ITS
LSU
gapdh
rpb2
LT837453
LT883391
LT883533
LT852480
Hernández-Restrepo et al. (2018)
Ex. holmii
CBS 413.65IsoT
LT837459
LT715621
LT715890
LT852489
Hernández-Restrepo et al. (2018)
Ex. khartoumensis
IMI 249194IsoT
LT852490
Hernández-Restrepo et al. (2018)
LT852502
Hernández-Restrepo et al. (2018)
LT852507
Hernández-Restrepo et al. (2018)
LT715886
LT852508
Hernández-Restrepo et al. (2018)
LT715878
HF934851
LT715902
HF934854
Amaradasa et al. (2014), Hernández-Restrepo
et al. (2018)
Amaradasa et al. (2014), Chowdhary et al.
(2015), Hernández-Restrepo et al. (2018)
Hernández-Restrepo et al. (2018)
LT715619
LT715888
Ex. minor
BRIP 14616
LT837470
LT883403
LT883545
Ex. monoceras
BRIP 12271A
LT837475
LT883406
LT883548
Ex. neoregeliae
CBS 132832T
LT837476
LT715617
Ex. oryzicola
CBS 502.90IsoT
HF934949
HF934886
Ex. pedicellatum
CBS 322.64ET
KT265258
HF934889
Ex. protrudens
BRIP 14814T
LT631308
LT715611
LT715880
LT715741
Ex. rostratum
CBS 325.87
KT265237
LT715629
LT715898
LT852492
Ex. turcicum
CBS 690.71ET
LT837487
LT883415
LT882581
–
M
AN
U
SC
LT837461
T
D
References
RI
PT
Exserohilum corniculatum
TE
Isolates1
EP
1
Species
Chowdhary et al. (2015), Hernández-Restrepo
et al. (2018)
Hernández-Restrepo et al. (2018)
AC
C
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; IMI:
International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, UK. T, ET, IsoT and A indicate ex-type, ex-epitype, ex-isotype, and authentic
strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; gapdh: partial glyceraldehyde-3-phosphate
dehydrogenase gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene.
ACCEPTED MANUSCRIPT
Table 6. DNA barcodes of accepted Neosetophoma spp.
Isolates
1
GenBank accession numbers
ITS
rpb2
tef1
MK539953
MK540084
–
KP744450
–
–
KY496758
–
KY514402a
MH018134
–
–
MF684861
–
–
KP711356
–
–
KX306763
–
–
CBS 145363T
MFLUCC 13-0734T
MFLUCC 14-0528T
GZCC 18-0111T
IBRC-M 30176T
MFLUCC 13-0388T
CBS 141409T
2
References
tub2
–
–
–
–
–
–
–
RI
PT
Species
EP
TE
D
M
AN
U
SC
Present study
Liu et al. (2015)
Tibpromma et al. (2017)
Hyde et al. (2018)
Karunarathna et al. (2017)
Liu et al. (2015)
Hernández-Restrepo et al.
(2016a)
Nph. phragmitis
CBS 145364T
MK539954
MK540085
MK540148b
–
Present study
Nph. poaceicola
MFLUCC 16-0886T
KY568986
–
–
–
Thambugala et al. (2017)
Nph. rosae
MFLUCC 15-1073T
MG828925
–
MG829218a
–
Wanasinghe et al. (2018)
Nph. rosarum
MFLUCC 17-0308T
MG828927
–
–
–
Wanasinghe et al. (2018)
Nph. rosigena
MFLUCC 17-0768T
MG828928
–
–
–
Wanasinghe et al. (2018)
Nph. samarorum
CBS 138.96ET
KF251160
KF252168
KF253119b
KF252655 Quaedvlieg et al. (2013)
Nph. sambuci
CBS 145365T
MK539955
MK540086
MK540149b
–
Present study
Nph. shoemakeri
MFLUCC 17-0780
MG844346
–
MG844352a
–
Hyde et al. (2018)
Nph. xingrensis
GZCC 18-0110T
MH018135
–
–
–
Hyde et al. (2018)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; GZCC: Guizhou Academy of Agricultural Sciences Culture Collection,
Guiyang, China; IBRC: Herbarium of the Plant bank, Iranian Biological Resource Center, Karaj, Iran; MFLUCC: Mae Fah Luang University
Culture Collection, Chiang Ria, Thailand. T and ET indicate ex-type and ex-epitype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1:
a
b
partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene. and in tef1 column indicate the primers used for
Neosetophoma aseptata
Nph. clematidis
Nph. garethjonesii
Nph. guiyangensis
Nph. iranianum
Nph. italica
Nph. lunariae
AC
C
sequencing: a: EF1-983F / EF1-2218R; b: EF1-728F / EF-2.
ACCEPTED MANUSCRIPT
Table 7. DNA barcodes of accepted Neostagonospora spp.
Isolates
1
GenBank accession numbers
rpb2
tef1
–
MG520901a
2
References
MFLUCC 15-0464T
CBS 135092T
CBS 135101T
MFLUCC 16-0493T
KF251163
KF251164
KX926416
KF252171
KF252172
–
–
KF253122b
MG520902a
KF252658
KF252659
–
Nst. sorghi
Nst. spinificis
CBS 145366T
BCRC FU30120
MK539956
KP676045
MK540087
LC055104
MK540150b
–
MK540168
–
1
tub2
–
RI
PT
Neostagonospora
arrhenatheri
Nst. caricis
Nst. elegiae
Nst. phragmitis
ITS
KX926417
SC
Species
Phookamsak et al. (2017),
Thambugala et al. (2017)
Quaedvlieg et al. (2013)
Quaedvlieg et al. (2013)
Phookamsak et al. (2017),
Thambugala et al. (2017)
Present study
Yang et al. (2016)
AC
C
EP
TE
D
M
AN
U
BCRC: Bioresource Collection and Research Centre, Taiwan; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the
Netherlands; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand. T indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit
gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers
used for sequencing: a: EF1-983F / EF1-2218R, b: EF1-728F / EF-2.
ACCEPTED MANUSCRIPT
CBS 125.93
GU238043
GU237771
KT389656
Not. gossypiicola
CBS 377.67
GU238079
GU237845
KT389658
Not. infossa
CBS 123395NT
GU238089
FJ427025
Not. macrospora
CBS 140674T
LN880537
Not. quercina
CBS 633.92
GU237583
References
Aveskamp et al. (2010), Chen et al.
(2015)
Aveskamp et al. (2010), Chen et al.
(2015)
Aveskamp et al. (2010), Chen et al.
(2015)
KT389659
FJ427135
Aveskamp et al. (2009, 2010), Chen et al.
(2015)
LN880536
LT593073
LN880539
Crous et al. (2016b)
EU754127
GU237900
KT389657
GU237609
Aveskamp et al. (2010), Chen et al.
(2015)
CGMCC 3.19246
MK088581
MK088574
MK088588
MK088595
Present study
LC12187
MK088582
MK088575
MK088589
MK088596
Present study
Not. raii
MCC 1082T
–
MF664467
–
MF664468
Crous et al. (2017b)
Not. variabilis
CBS 142457T
LN907428
LT592939
LT593078
LT593008
Valenzuela-Lopez et al. (2018)
TE
EP
1
M
AN
U
GU237611
D
SC
Not. arachidis-hypogaeae
RI
PT
Table 8. DNA barcodes of accepted Nothophoma spp.
Species
Isolates1
GenBank accession numbers2
LSU
ITS
rpb2
tub2
ET
Nothophoma anigozanthi
CBS 381.91
GU238039 GU237852 KT389655 GU237580
AC
C
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbiological Culture Collection Center, Beijing,
China; LC: Dr Lei Cai's personal culture collection, housed at CAS, China; MCC: National Centre for Microbial Resources (formerly Microbial Culture
Collection), Pune, India. T, ET and NT indicate ex-type, ex-epitype and ex-neotype strains, respectively.
2
LSU: partial large subunit (28S) nrRNA gene; ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA
polymerase II second largest subunit gene; tub2: partial β-tubulin gene.
ACCEPTED MANUSCRIPT
Table 9. DNA barcodes of accepted Parastagonospora spp.
Parastagonospora
allouniseptata
P. avenae
Isolates1
ITS
KU058711
MFLUCC 13-0386T
GenBank accession numbers2
rpb2
tef1
–
MG520914a
References
tub2
–
RI
PT
Species
AC
C
EP
TE
D
M
AN
U
SC
Li et al. (2015), Phookamsak et al.
(2017)
CBS 289.69
KF252669
Quaedvlieg et al. (2013)
KF251174
KF252182
KF253132b
MFLUCC 13-0557T of P. forlicesenica
KY769660
–
–
–
Thambugala et al. (2017)
P. caricis
CBS 135671T
KF251176
KF252184
KF253134b
KF252671
Quaedvlieg et al. (2013)
P. dactylidis
MFLUCC 13-0375T
KU058712
–
–
–
Li et al. (2015)
MFLUCC 13-0376T of P. minima
KU058713
–
MG520916a
–
Li et al. (2015), Phookamsak et al.
(2017)
MFLUCC 13-0573T of P. cumpignensis
KU842388
–
–
–
Li et al. (2016a)
P. fusiformis
MFLUCC 13-0215T
KX926418
KX863711
–
–
Thambugala et al. (2017)
P. italica
MFLUCC 13-0377T
–
Li et al. (2015), Phookamsak et al.
KU058714
–
MG520915a
(2017)
P. nodorum
CBS 110109
KF251177
KF252185
KF253135b
KF252672
Quaedvlieg et al. (2013)
P. novozelandica
CBS 145416T
MK539957
MK540088
MK540151
MK540169
Present study
P. phragmitis
CBS 143446T
MK539958
MK540089
MK540152
–
Present study
P. poaceicola
MFLUCC 15-0471T
KX926419
KX880499
–
–
Thambugala et al. (2017)
P. poae
CBS 135089T
KF252673
Quaedvlieg et al. (2013)
KF251178
KF252186
KF253136b
P. poagena
CBS 136776T
KJ869116
–
–
–
Crous et al. (2014b)
P. uniseptata
KU058715
–
MG520917a
–
Li et al. (2015), Phookamsak et al.
MFLUCC 13-0387T
(2017)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand. T
indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial
translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used in sequencing: a: EF1-983F / EF1-2218R,
b
: EF1-728F / EF-2.
ACCEPTED MANUSCRIPT
Table 10. DNA barcodes of accepted Phaeosphaeriopsis spp.
Species
Isolates
1
GenBank accession numbers
rpb2
tef1
MK540094
MK540157a
2
References
CBS 141287T
CBS 142110T
KY173430
KY173591
MK540158a
KY173610
Phs. agavensis
Phs. aloes
Phs. aloicola
Phs. amblyospora
Phs. dracaenicola
Phs. glaucopunctata
CBS 102206
CBS 145367T
CBS 145368T
CBS 110131T
MFLUCC 11-0157T
MFLUCC 13-0265ET
KY090635
MK539959
MK539960
AY188993
KM434273
KJ522473
KY090685
MK540090
MK540091
–
KM434309
–
–
MK540153a
MK540154a
–
KM434301b
MG520918b
–
–
MK540170
–
–
–
Phs. grevilleae
Phs. nolinae
Phs. obtusispora
Phs. phacidiomorpha
Phs. pseudoagavacearum
Phs. triseptata
CBS 145369T
CBS 102205
CBS 102204
T111
CBS 145370T
MFLUCC 13-0271T
MK539961
KY090637
KY090636
FJ462742
MK539962
KJ522475
MK540092
KY090686
KY090687
–
MK540093
KJ522485
MK540155a
–
–
–
MK540156a
MG520919b
MK540171
–
–
–
MK540172
–
Phs. yuccae
MFLUCC 16-0558T
KY554482
–
MG520920b
–
SC
M
AN
U
D
TE
1
tub2
MK540173
RI
PT
Phaeosphaeriopsis
agapanthi
Phs. agavacearum
ITS
KX228260
Crous et al. (2016b),
present study
Crous et al. (2016a),
present study
Ahmed et al. (2017)
Present study
Present study
Câmara et al. (2003)
Phookamsak et al. (2014b)
Thambugala et al. (2014),
Phookamsak et al. (2017)
Present study
Ahmed et al. (2017)
Ahmed et al. (2017)
Zhang et al. (unpubl. data)
Present study
Thambugala et al. (2014),
Phookamsak et al. (2017)
Phookamsak et al. (2017)
AC
C
EP
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLUCC: Mae Fah Luang University Culture Collection,
Chiang Ria, Thailand; T: isolate housed in China. T and ET indicate ex-type and ex-epitype strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit
gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers
used in sequencing: a: EF1-728F / EF-2, b: EF1-983F / EF1-2218R.
ACCEPTED MANUSCRIPT
Table 11. DNA barcodes of accepted Pleiocarpon spp.
Pleiocarpon livistonae
Pl. strelitziae
Isolates
1
CBS 145030T
CBS 142251T
2
ITS
MK539963
KY304644
GenBank accession numbers
his3
rpb2
tef1
MK540234
MK540095
MK540165
KY304616
KY304697
KY304722
RI
PT
Species
Present study
Aiello et al. (2017)
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; his3: partial histone H3 gene; rpb2: partial DNA-directed RNA
polymerase II second largest subunit gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
AC
C
EP
TE
D
M
AN
U
SC
1
References
tub2
MK540179
KY304750
ACCEPTED MANUSCRIPT
Table 12. DNA barcodes of accepted Pyrenophora spp.
1
GenBank accession numbers
LSU
gapdh
MK540042
MK540180
MK540043
MK540181
MK540044
MK540182
JN712532
MK540183
Pyrenophora avenicola
Py. biseptata
CBS 307.84
CBS 307.69
CBS 319.69
CBS 108963
ITS
MK539972
MK539973
MK539974
MK539975
Py. bromi
CBS 311.68
MK539976
MH870851
DAOMC 127414
JN943666
JN940074
CBS 279.31
CBS 195.31
A
MK539977
MK539978
MK540184
–
AY004839
–
MK540045
MH866633
MK540185
MK540186
MK540103
MK540104
MK540046
MK540047
JN940087
MK540187
MK540188
AY004812
MK540105
MK540106
–
M
AN
U
Py. chaetomioides
T
References
rpb2
–
–
MK540102
–
RI
PT
Isolates
SC
Species
CBS 314.68
T
CBS 127918
DAOMC 92161
MK539979
MK539980
JN943667
Py. dictyoides
DAOMC 63666
JN943653
JN940080
AY004836
–
CBS 258.80
CBS 967.87
CBS 127933
MK539981
MK539982
MH877971
MK540048
MK540049
MK540050
MK540189
MK540190
MK540191
MK540107
MK540108
MK540109
CBS 312.69
CBS 108941
CBS 509.77
CBS 315.69
CBS 128043
MK539983
MK539984
MK539985
MK539986
MK539987
MK540051
MK540052
MK540053
MK540054
MH876230
MK540192
MK540193
MK540194
MK540195
MK540196
–
MK540110
MK540111
–
MK540112
CBS 128044
MK539988
MH876231
MK540197
MK540113
JN712467
JN712533
MK540198
MK540114
Py. fugax
Py. grahamii
Py. leucospermi
TE
EP
AC
C
Py. erythrospila
D
Py. cynosuri
Py. dactylidis
T
CBS 111083
Present study
Present study
Present study
Crous et al. (2011),
present study
Vu et al. (2019), present
study
Zhang & Berbee (2001),
Hambleton (unpubl.
data)
Present study
Vu et al. (2019), present
study
Present study
Present study
Zhang & Berbee (2001),
Hambleton (unpubl.
data)
Zhang & Berbee (2001),
Hambleton (unpubl.
data)
Present study
Present study
Vu et al. (2019), present
study
Present study
Present study
Present study
Present study
Vu et al. (2019), present
study
Vu et al. (2019), present
study
Crous et al. (2011),
ACCEPTED MANUSCRIPT
Py. nobleae
MK540115
CBS 114493
MK539990
JN712545
MK540200
MK540116
CBS 240.48
CBS 318.69
MK539991
MK539992
MK540055
MH871050
MK540201
MK540202
MK540117
MK540118
CBS 128046
MK539993
MH876233
MK540203
MK540119
KM257054
KM243296
CBS 119213
EU552124
MK540056
CBS 127912
MH877963
CBS 259.80
CBS 966.87
CBS 127936
T
CBS 127934
DAOMC 222769
MK539994
MK539995
MK539996
MK539997
JN943649
CBS 190.29
ET
RI
PT
MK540199
KM257057
–
MK540204
MK540120
MK540057
MK540205
MK540121
MK540058
MK540059
MK540060
MK540061
JN940093
MK540206
MK540207
MK540208
MK540209
–
MK540122
MK540123
MK540124
MK540125
DQ497614
MK540062
MH876232
MK540210
MK540211
MK540126
MK540127
JN943650
JN940083
AY004832
JN993632
A
CBS 319.68
CBS 128045
AC
C
DAOMC 145373
MK539998
MK539999
EP
Py. poae
TE
D
Py. novozelandica
Py. phaeocomes
JN712542
SC
Py. nisikadoi
MK539989
M
AN
U
Py. lolii
CBS 111505
Py. pseudoerythrospila
Py. semeniperda
CBS 127931T
DAOMC 213153
MK540000
JN943665
MK540063
JN940088
MK540212
AY004826
–
–
Py. sieglingiae
Py. teres
BRIP 10941
CBS 127927
CBS 127930
T of Py. teres f.
CBS 228.76
KJ415564
MK540001
MK540002
MK540003
KJ415518
MK540064
MK540065
MK540066
KJ415382
MK540213
MK540214
MK540215
–
MK540128
MK540129
MK540130
present study
Crous et al. (2011),
present study
Crous et al. (2011),
present study
Present study
Vu et al. (2019), present
study
Vu et al. (2019), present
study
Manamgoda et al.
(2014)
Marincowitz et al.
(2008), present study
Vu et al. (2019), present
study
Present study
Present study
Present study
Present study
Hambleton (unpubl.
data), James et al.
(unpubl. data)
Present study
Vu et al. (2019), present
study
Zhang & Berbee (2001),
Schoch et al. (2012),
Hambleton (unpubl.
data)
Present study
Zhang & Berbee (2001),
Hambleton (unpubl.
data)
Tan et al. (2014)
Present study
Present study
Present study
ACCEPTED MANUSCRIPT
maculata
MK540131
MK540132
MK540133
MK540134
CBS 123929
CBS 123932
DAOMC 171966
MK540008
MK540009
JN943663
MK540070
MK540071
JN940090
MK540220
MK540221
–
MK540135
MK540136
JN993620
CBS 127915
MK540010
MH877964
CBS 127924
MK540011
MH877965
CBS 328.53
CBS 391.54
CBS 392.54
CBS 128047
MK540012
MK540013
MK540014
MK540015
CBS 128048
MK540016
SynT of Py.
Py. tritici-repentis
CBS 259.59
Py. variabilis
Py. wirreganensis
CBS 191.29
CBS 127922
T
CBS 127920
CBS 109896
tritici-vulgaris
SC
RI
PT
MK540216
MK540217
MK540218
MK540219
MK540222
MK540137
MK540223
MK540138
MK540072
MK540073
MK540074
MH877983
MK540224
MK540225
MK540226
MK540227
MK540139
–
MK540140
MK540141
MH876234
MK540228
MK540142
MK540017
MK540075
AM884276
MK540143
MK540018
MK540019
MK540020
MK540021
MK540076
MK540077
MK540078
MK540079
MK540229
MK540230
MK540231
MK540232
MK540144
MK540145
MK540146
MK540147
M
AN
U
Py. triseptata
MK540067
MK540068
MK540069
MH877692
D
Py. trichostoma
MK540004
MK540005
MK540006
MK540007
EP
Py. tetrarrhenae
A of Py. japonica
TE
CBS 281.31
CBS 282.31
CBS 314.69
CBS 336.29
Present study
Present study
Present study
Vu et al. (2019), present
study
Present study
Present study
Schoch et al. (2012),
Hambleton (unpubl.
data)
Vu et al. (2019), present
study
Vu et al. (2019), present
study
Present study
Present study
Present study
Vu et al. (2019), present
study
Vu et al. (2019), present
study
Lepoint et al. (2010),
present study
Present study
Present study
Present study
Present study
AC
C
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The
Netherlands; DAOMC: Plant Research Institute, Department of Agriculture (Mycology), Ottawa, Canada. A, ET, SynT and T indicate
authentic, ex-epitype, ex-syntype and ex-type strains, respectively.
1
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; gapdh: partial
glyceraldehyde-3-phosphate dehydrogenase gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene.
ACCEPTED MANUSCRIPT
Table 13. DNA barcodes of accepted Ramichloridium spp.
Ramichloridium apiculatum
Isolates1
T
CBS 156.59
ITS
EU041791
GenBank accession numbers2
rpb2
tef1
LSU
EU041848
MF951416
–
References
Arzanlou et al. (2007), Videira et al.
(2017)
Li et al. (2012)
Li et al. (2012), Videira et al. (2017)
Zhang et al. (2007)
Li et al. (2012)
the work in which it was introduced. T indicates
RI
PT
Species
AC
C
EP
TE
D
M
AN
U
SC
R. cucurbitae
CBS 132087T
JQ622087
JQ622095
–
JQ622112
T
R. luteum
CBS 132088
EU329730
JQ622099
MF951417
JQ622116
T
R. malus
LQ73
EF627452
–
–
–
JQ622086
JQ622094
–
JQ622111
R. punctatum
CBS 132090T
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; LQ: collection not specified in
ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S)nrDNA; rpb2: partial DNA-directed RNA polymerase II second
largest subunit gene; tef1: partial elongation factor 1-alpha gene.
ACCEPTED MANUSCRIPT
Table 14. DNA barcodes of accepted Seifertia spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
GenBank accession numbers2
References
LSU
ITS
tef1
Seifertia azaleae
DAOM 239136
EU030276
–
–
Seifert et al. (2007)
CPC 35017
MK540034
MK539964 MK540166 Present study
Sei. shangrilaensis MFLUCC 16-0238T
KU954100
–
KU954101
Li et al. (2016b)
1
CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute; DAOM: Plant Research Institute,
Department of Agriculture (Mycology), Ottawa, Canada; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria,
Thailand. T indicates ex-type strains.
2
LSU: partial large subunit (28S) nrRNA gene; ITS: internal transcribed spacers and intervening 5.8S nrDNA; tef1: partial translation
elongation factor 1-alpha gene.
Species
ACCEPTED MANUSCRIPT
Table 15. DNA barcodes of accepted Seiridium spp.
Isolates.1
MFLUCC 12-0647T
CBS 226.55T
CBS 909.85R
PHSI2001Pathcw07
CBS 224.55ET
CBS 343.97ET
CBS 142629T
CBS 228.55T
ITS
JQ683725
LT853089
LT853064
AY687314
LT853083
LT853099
LT853100
LT853098
GenBank accession numbers2
rpb2
tef1
tub2
–
JQ683741
JQ683709
LT853137
LT853186
LT853236
LT853113
LT853161
LT853211
–
–
DQ534043
LT853131
LT853180
LT853230
LT853146
LT853196
LT853246
LT853147
LT853197
LT853247
LT853145
LT853195
LT853245
RI
PT
Species
References
AC
C
EP
TE
D
M
AN
U
SC
Maharachchikumbura et al. (2015)
Bonthond et al. (2018)
Bonthond et al. (2018)
Liu et al. (2007)
Bonthond et al. (2018)
Bonthond et al. (2018)
Bonthond et al. (2018)
Bonthond et al. (2018)
Jaklitsch et al. (2016), Bonthond et al.
S. marginatum
CBS 140403GT
KT949914
LT853149
LT853199
LT853249
(2018)
S. neocupressi
CBS 142625T
LT853079
LT853127
LT853176
LT853226
Bonthond et al. (2018)
S. papillatum
CBS 340.97T
LT853102
LT853150
LT853200
LT853250
Bonthond et al. (2018)
S. persooniae
CBS 143445T
MG386033
–
–
MG386163
Crous et al. (2017b)
S. pezizoides
CBS 145115
MK079342
MK058475
MK058480
MK058485
Present study
S. phylicae
CBS 133587T
LT853091
LT853139
LT853188
LT853238
Bonthond et al. (2018)
S. podocarpi
CBS 137995T
LT853101
LT853148
LT853198
LT853248
Bonthond et al. (2018)
S. pseudocardinale
MFLUCC 13-0525T
KU848210
–
–
–
Wijayawardene et al. (2016)
CBS 145114
MK079341
MK058479
MK058484
MK058489
Present study
T
S. rosarum
MFLUCC 17-0654
MG828961
–
–
–
Wanasinghe et al. (2018)
S. spyridicola
CBS 142628T
LT853095
LT853142
LT853192
LT853242
Bonthond et al. (2018)
S. unicorne
CBS 143871ET
MK079339
MK058477
MK058482
MK058487
Present study
CBS 143872
MK079338
MK058476
MK058481
MK058486
Present study
CBS 143873
MK079340
MK058478
MK058483
MK058488
Present study
CBS 538.82R
LT853088
LT853136
LT853185
LT853235
Bonthond et al. (2018)
S. venetum
MFLU 15-0369R
KT438836
–
–
KT438837
Maharachchikumbura et al. (2015)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLU and MFLUCC: Herbarium and culture collection of Mae Fah Luang
University, Chiang Rai, Thailand, respectively; PHSI: from Liu et al. (2007). T, ET, GT and R indicate type or ex-type, ex-epitype, ex-generic type and reference
strains, respectively.
Seiridium camelliae
S. cancrinum
S. cardinale
S. ceratosporum
S. cupressi
S. eucalypti
S. kartense
S. kenyanum
ACCEPTED MANUSCRIPT
2
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial
translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
ACCEPTED MANUSCRIPT
Table 16. DNA barcodes of accepted Septoriella spp.
Species
Isolates
1
GenBank accession numbers
rpb2
tef1
MG829227a
MG520935a
2
References
MFLU 18-0113HT
MFLUCC 15-0701T
Sep. artemisiae
Sep. arundinicola
Sep. arundinis
MFLUCC 17-0693T
MFLU 16-0225HT
MFLUCC 15-0702T
MG828929
MG828946
KU058716
MG829261
-
MG829228a
MG520921a
-
Sep. bromi
Sep. chlamydospora
Sep. dactylidicola3
Sep. dactylidis
Sep. elongata
Sep. forlicesenica
MFLUCC 13-0739T
MFLUCC 15-0177T
MFLUCC 14-0002T
MFLU 15-2720HT
MFLUCC 12-4444T
MFLUCC 15-0470T
KU058717
KU163658
KU163657
KM491546
KX926422
KY131966
MG520922a
-
Sep. garethjonesii
MFLUCC 15-0469T
KX926425
KX898363
MG520923a
-
Sep. germanica
Sep. hibernica
Sep. hirta
Sep. hollandica
Sep. hubertusii
Sep. italica
CBS 145372T
CBS 145371T
CBS 536.77ET
CBS 145374T
CBS 338.86T
MFLUCC 13-0267T
MK539965
MK539966
KR873249
MK539967
KF251230
KX926421
MK540096
MK540097
KR873324
MK540098
KF252235
KX891169
MK540159b
MK540160b
MG520924a
MK540174
MK540175
KF252717
-
Sep. leuchtmannii
Sep. muriformis
Sep. neoarundinis
CBS 459.84IsoT
MFLUCC 13-0277T
MFLUCC 15-0027T
KF251188
KX926415
KY706139
KF252195
KX863710
-
KF253144b
MG520936a
KF252682
-
Sep. neodactylidis
Sep. oudemansii
Sep. phragmitis
Sep. poae
Sep. pseudophragmitis
MFLUCC 13-0618T
CBS 138012T
CBS 140065ET
CBS 136766T
CBS 145417T
KP744432
KR873250
KR873251
KJ869111
MK560161
KJ869233
MK559450
MK559452b
MK559451
SC
M
AN
U
D
TE
EP
AC
C
tub2
-
RI
PT
Septoriella agrostina
Sep. allojunci
ITS
MG828945
KU058718
Wanasinghe et al. (2018)
Li et al. (2015), Phookamsak
et al. (2017)
Wanasinghe et al. (2018)
Wanasinghe et al. (2018)
Li et al. (2015), Phookamsak
et al. (2017)
Li et al. (2015)
Hyde et al. (2018)
Jayasiri et al. (2015)
Li et al. (2015)
Phookamsak et al. (2017),
Thambugala et al. (2017)
Phookamsak et al. (2017),
Thambugala et al. (2017)
Present study
Present study
Crous et al. (2015a)
Present study
Quaedvlieg et al. (2013)
Phookamsak et al. (2017),
Thambugala et al. (2017)
Quaedvlieg et al. (2013)
Thambugala et al. (2017)
Phookamsak et al. (2017),
Thambugala et al. (2017)
Liu et al. (2015)
Crous et al. (2015a)
Crous et al. (2015a)
Crous et al. (2014b)
Present study
ACCEPTED MANUSCRIPT
MFLU 18-0114HT
MFLUCC 13-0380T
MFLUCC 15-0475T
CBS 604.86
MG828948
KT314184
KX926424
KF251193
KX891171
KF252200
MG829230a
KF253149b
KF252687
RI
PT
Sep. rosae
Sep. subcylindrospora
Sep. tridentina
Sep. vagans
1
Wanasinghe et al. (2018)
Ariyawansa et al. (2015a)
Thambugala et al. (2017)
Quaedvlieg et al. (2013)
AC
C
EP
TE
D
M
AN
U
SC
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLU, MFLUCC: Mae Fah Luang University Culture
Collection, Chiang Rai, Thailand. T, ET, HT and IsoT indicate ex-type, ex-epitype, holotype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial RNA polymerase II second largest subunit gene; tef1:
partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used in
sequencing: a: EF1-983F, EF1-2218R, b: EF1-728F, EF-2.
3
Only LSU and SSU sequences available: KY657264 and KY657265, respectively (Thambugala et al. 2017).
ACCEPTED MANUSCRIPT
Table 17. DNA barcodes of accepted Setophoma spp.
Isolates1
CBS 145418T
CBS 135105T
CBS 145373T
CBS 333.39ET
CBS 335.29LT
CBS 137988T
ITS
MK539968
KF251244
MK539969
KF251245
KF251246
KJ869141
GenBank accession numbers2
rpb2
tef1
MK540099 MK540161
KF252249
KF253195
MK540100 –
KF252250
–
KF252251
KF253196
–
MK540162
References
tub2
–
KF252728
MK540176
–
KF252729
MK540177
RI
PT
Species
AC
C
EP
TE
D
M
AN
U
SC
Present study
Quaedvlieg et al. (2013)
Present study
Quaedvlieg et al. (2013)
Quaedvlieg et al. (2013)
Crous et al. (2014b),
present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T, ET and LT indicate ex-type strains, ex-epitype and exlectotype, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit
gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
Setophoma brachypodii
Set. chromolaenae
Set. pseudosacchari
Set. sacchari
Set. terrestris
Set. vernoniae
ACCEPTED MANUSCRIPT
Table 18. DNA barcodes of the accepted Wingfieldomyces sp.
Isolates1
Wingfieldomyces cyperi CBS 141450T
ITS
KX228286
GenBank accession numbers2
rpb2
tef1
tub2
MK540101
MK540163
MK540178
RI
PT
Species
1
References
Crous et al. (2016b),
present study
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T indicates ex-type strain.
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit
gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
AC
C
EP
TE
D
M
AN
U
SC
2
ACCEPTED MANUSCRIPT
Table 19. DNA barcodes of accepted Stagonosporopsis spp.
Strain1
GenBank accession numbers2
T
ITS
LSU
act
rpb2
References
RI
PT
Species
tub2
CBS 106.96
GU237734
GU238166
JN251974
KT389672
Sta. ailanthicola
Sta. ajacis
MFLUCC 16-1439T
CBS 177.93NT
KY100872
GU237791
KY100874
GU238168
–
JN251962
KY100876
KT389673
Sta. andigena
Sta. artemisiicola
CBS 101.80
CBS 102636
GU237714
GU237728
GU238169
GU238171
JN251958
JN251971
–
KT389674
Sta. astragali
Sta. bomiensis
Sta. centaureae
Sta. citrulli
Sta. caricae
Sta. chrysanthemi
Sta. crystalliniformis
CBS 178.25
CGMCC 3.18366T
MFLUCC 16-0787T
ATCC TSD-2T
CBS 248.90
DAR 28714
ATCC 10748ET
CBS 713.85T
GU237792
KY742123
KX611240
KJ855546
GU237807
MK253236
JQ897484
GU237903
GU238172
KY742277
KX611238
–
GU238175
MK253237
JQ897460
GU238178
JN251963
–
–
–
JN251969
MK255065
JQ897508
JN251960
Sta. cucurbitacearum
CBS 133.96
GU237780
GU238181
Sta. dennisii
Sta. dorenboschii
CBS 631.68ET
CBS 426.90T
GU237899
GU237862
Sta. helianthi
Sta. heliopsidis
CBS 200.87T
CBS 109182
KT389545
GU237747
Sta. hortensis
CBS 572.85
Sta. inoxydabilis
Sta. loticola
GU237671
KY100878
GU237673
GU237674
GU237676
GU237677
KY742365
–
KJ855602
GU237680
MK255067
JQ897504
GU237683
JN251968
KT389676
GU237686
GU238182
GU238185
Q-bank3
JN251980
KT389677
KT389678
GU237687
GU237690
KT389761
GU238186
–
JN251980
KT389683
KT389679
KT389848
GU237691
GU237730
GU238198
JN251966
KT389680
GU237703
CBS 425.90T
GU237861
GU238188
JN251972
KT389682
GU237693
CBS 562.81IsoT
GU237890
GU238192
JN251978
KT389684
GU237697
EP
TE
D
–
KY742189
–
–
–
MK255066
–
KT389675
AC
C
M
AN
U
SC
Stagonosporopsis actaeae
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Tibpromma et al. (2017)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012)
Chen et al. (2017)
Hyde et al. (2016)
Stewart et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012)
Present study
Vaghefi et al. (2012)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
ACCEPTED MANUSCRIPT
CBS 101494T
GU237724
GU238194
JN251967
KT389685
GU237699
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Sta. oculo-hominis
CBS 634.92T
GU237901
GU238196
JN251976
KT389686
GU237701
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Sta. papillata
CGMCC 3.18367T
KY742125
KY742279
–
KY742191
KY742367
Chen et al. (2017)
Sta. rudbeckiae
CBS 109180
GU237745
GU238197
Q-bank3
–
GU237702
Aveskamp et al. (2010)
Sta. tanaceti
CBS 131484T
JQ897481
JQ897461
JQ897512
–
JQ897496
Vaghefi et al. (2012)
Sta. trachelii
CBS 379.91
GU237850
GU238173
JN251977
KT389687
GU237678
Aveskamp et al. (2010), de Gruyter et al. (2012),
Chen et al. (2015)
Sta. valerianellae
CBS 329.67IsoT
GU237832
GU238201
JN251965
–
GU237706
Aveskamp et al. (2010), de Gruyter et al. (2012)
1
ATCC: American Type Culture Collection, Virginia, USA; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbiological Culture
Collection Center, Beijing, China; DAR: New South Wales Plant Pathology Herbarium, NSW, Australia; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand. T,
ET, IsoT
and NT indicate ex-type, ex-epitype, ex-isotype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; act: partial actin gene; rpb2: partial DNA-directed RNA polymerase II second
largest subunit; tub2: partial β-tubulin gene.
3
Q-bank: Sequences retrieved from Q-bank Fungi database (http://www.q-bank.eu/fungi/).
Sta. lupini
ACCEPTED MANUSCRIPT
Table 20. DNA barcodes of accepted Stemphylium spp.
Ste. drummondii
Ste. eturmiunum
Ste. gracilariae
Ste. halophilum
Ste. ixeridis
Ste. lancipes
Ste. loti
Ste. lucomagnoense
Ste. lycii
Ste. lycopersici
References
RI
PT
SC
M
AN
U
Ste. botryosum
Ste. callistephi
Ste. canadense
Ste. chrysanthemicola
D
CBS 124746T
CBS 338.73
CBS 116583ET
CBS 141024T
VPRI 42502
CBS 714.68T
CBS 527.50T
CBS 116602T
CBS 117255T
VPRI 10316
CBS 346.83T
CBS 109845T
BRIP 27557
BRIP 27560
BRIP 55702
BRIP 60383
BRIP 62759
BRIP 63689
CBS 482.90T
CBS 337.73T
CBS 124748T
CBS 133314ET
CBS 407.54T
CBS 116601T
CBS 125241T
CBS 122639NT
BRIP 13821
BRIP 13902
BRIP 15113
TE
Stemphylium amaranthi
Ste. armeriae
Ste. astragali
Ste. beticola
GenBank accession numbers2
ITS
gapdh
cmdA
KU850505
KU850652
KU850793
KU850511
KU850658
KU850799
KU850512
KU850659
KU850800
KU850520
KU850667
KU850808
MK336834
MK336880
MK336857
KC584238
AF443881
KU850826
KU850539
KU850686
KU850828
KU850641
KU850782
KU850932
KU850640
KU850781
KU850931
MK336835
MK336881
MK336858
GQ395365
KU850687
KU850829
KU850831
KU850689
KU850541
MK336843
MK336866
MK336820
MK336844
MK336867
MK336821
MK336849
MK336872
MK336826
MK336850
MK336873
MK336827
MK336853
MK336876
MK336830
MK336854
MK336877
MK336831
KU850549
AF443883
KU850839
KU850553
KU850700
KU850843
KU850590
KU850737
KU850881
KU850596
KU850742
KU850887
KU850597
KU850743
KU850888
KU850629
KU850770
KU850920
KU850602
KU850748
KU850893
KU850611
KU850756
KU850902
MK336813
MK336859
MK336836
MK336814
MK336860
MK336837
MK336817
MK336863
MK336840
EP
Isolates1
AC
C
Species
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Present study
Present study
Present study
Present study
Present study
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Present study
Present study
ACCEPTED MANUSCRIPT
MK336851
Present study
CBS 717.68T
CBS 138295T
CBS 109842T
BRIP 27486HT
CBS 110049
CBS 133518T
CBS 116586ET
CBS 115268T
CBS 116580T
CBS 718.68T
BRIP 14850HT
CBS 715.68
BRIP 5891
BRIP 14857
BRIP 26714
BRIP 40125
BRIP 40155
BRIP 52457
BRIP 53984
VPRI 31963
VPRI 21969HT
KU850618
KU850631
KU850620
MK336819
KU850591
KU850637
KU850627
KU850643
KU850647
KU850648
MK336815
KU850565
MK336829
MK336816
MK336818
MK336822
MK336823
MK336824
MK336825
MK336833
MK336832
AF443891
KU850772
KU850762
MK336865
KU850738
KU850778
KU850768
KU850784
KU850788
KU850789
MK336861
KU850712
MK336875
MK336862
MK336864
MK336868
MK336869
MK336870
MK336871
MK336879
MK336878
KU850909
KU850922
KU850911
MK336842
KU850882
KU850928
KU850918
KU850934
KU850938
KU850939
MK336838
KU850855
MK336852
MK336839
MK336841
MK336845
MK336846
MK336847
MK336848
MK336856
MK336855
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Woudenberg et al. (2017)
Present study
Woudenberg et al. (2017)
Present study
Present study
Present study
Present study
Present study
Present study
Present study
Present study
Present study
D
M
AN
U
SC
RI
PT
MK336874
TE
Ste. waikerieanum
1
MK336828
EP
Ste. majusculum
Ste. novae-zelandiae
Ste. paludiscirpi
Ste. rombundicum
Ste. sarciniforme
Ste. simmonsii
Ste. solani
Ste. symphyti
Ste. trifolii
Ste. triglochinicola
Ste. truncatulae
Ste. vesicarium
BRIP 62317
AC
C
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Queensland, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the
Netherlands; VPRI: Victorian Plant Pathology Herbarium, Bundoora, Victoria, Australia. T, ET, HT and NT indicate ex-type, ex-epitype, ex-holotype
and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; cmdA: partial
calmodulin gene.
ACCEPTED MANUSCRIPT
Table 21. DNA barcodes of accepted Tubakia spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
GenBank accession number2
References
rpb2
tef1
tub2
ITS
Tubakia americana CBS 129014
MG591873
MG976449 MG592058 MG592152 Braun et al. (2018)
T
T. braunii
CBS 115011
MG591912
MG976488 MG592101 MG592193 Braun et al. (2018)
T. californica
CBS 143670T
MG591835
MG976451 MG592023 MG592117 Braun et al. (2018)
ET
T. dryina
CBS 112097
MG591851
MG976455 MG592039 MG592133 Braun et al. (2018)
T
MG591878
MG976461 MG592063 MG592157 Braun et al. (2018)
T. dryinoides
NBRC 9267
T
T. hallii
CBS 129013
MG591880
MG976462 MG592065 MG592159 Braun et al. (2018)
T
T. iowensis
CBS 129012
MG591879
–
MG592064 MG592158 Braun et al. (2018)
MG591886
MG976465 MG592071 MG592165 Braun et al. (2018)
T. japonica
NBRC 9268ET
T. liquidambaris
CBS 139744
MG605068
–
MG603578 –
Harrington & McNew (2018)
T
MG605069
–
MG603579 –
Harrington & McNew (2018)
T. macnabbii
CBS 137349
T
T. melnikiana
CPC 32255
MG591893
MG976472 MG592080 MG592174 Braun et al. (2018)
T
T. oblongispora
NBRC 9885
MG591897
MG976474 MG592084 MG592178 Braun et al. (2018)
T
NBRC 9884
MG591898
MG976475 MG592085 MG592179 Braun et al. (2018)
T. paradryinoides
IsoT
T. seoraksanensis
CBS 127492
MG591908
MG976485 MG592096 MG592188 Braun et al. (2018)
T
CPC 33020
MG591910
MG976486 MG592099 MG592191 Braun et al. (2018)
T. sierrafriensis
IsoT
T. suttoniana
CBS 639.93
MG591921
MG976493 MG592110 MG592202 Braun et al. (2018)
T
T. tiffanyae
CBS 137345
MG605081
–
–
–
Harrington & McNew (2018)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at the Westerdijk Fungal
Biodiversity Institute; NBRC: Biological Resource Center, NITE, Chiba, Japan. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial
translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
Species
ACCEPTED MANUSCRIPT
Table 22. DNA barcodes of accepted Zasmidium spp.
Species
Zasmidium angulare
Isolates1
T
CBS 132094
T
ITS
JQ622088
GenBank accession numbers2
rpb2
act
tef1
LSU
JQ622096 MF951690 –
JQ622113
tub2
–
References
CBS 118742
FJ839626
FJ839662
MF951691
–
KF253229
KF252763
Z. aporosae
Z. arcuatum
MFLU 12-2206HT
CBS 113477T
KC677912
EU041779
–
EU041836
–
MF951692
–
–
–
–
–
–
Z. aucklandicum
Z. biverticillatum
Z. cellare
CPC 13569
CBS 335.36
CBS 146.36NT
MF951409
EU041796
EU041821
MF951280
EU041853
EU041878
MF951733
–
MF951693
–
–
–
Z. cerophillum
CBS 103.59T
EU041798
GU214485
MF951694
Z. citri-griseum
Z. commune
Z. cyatheae
Z. dasypogonis
Z. daviesiae
CBS 139467ET
CBS 142530T
CPC 24725T
CBS 143397T
CBS 116002
KF901796
KY979765
KT037530
MG386048
FJ839633
KF902155
KY979820
KT037571
MG386101
FJ839669
Z. ducassei
Z. elaeocarpi
Z. eucalypticola
Z. eucalyptigenum
BRIP 53367T
CPC 32929
CBS 142187T
CBS 142186T
CBS 138860T
HQ149687
MK539971
MF951398
MF951400
KP004458
Z. eucalyptorum
CBS 118500T
Z. fructicola
Z. fructigenum
SC
RI
PT
Z. anthuriicola
–
–
–
–
–
–
KF902518
–
–
–
MF951698
–
KY979860
KT037629
–
KF903477
KF903385
–
KT037490
–
KF903373
KF903079
–
–
–
KF903069
–
MK540041
MF951263
MF951265
KP004486
–
–
MF951699
MF951701
–
–
MK540233
–
–
KT037630
–
MK540167
–
–
–
–
–
–
–
KF901652
MF951266
MF951702
KF903495
KF903101
–
CBS 139625T
KP896052
KP895922
MF951703
KP896003
KP896099
–
CBS 139626T
KP896056
KP895926
MF951704
KP896007
KP896103
KP896149
AC
C
EP
TE
D
M
AN
U
–
–
–
Li et al. (2012), Videira et al. (2017)
Crous et al. (2009a), Quaedvlieg et
(2013), Videira et al. (2017)
Phengsintham et al. (2013)
Arzanlou et al. (2007), Videira et
(2017)
Videira et al. (2017)
Arzanlou et al. (2007)
Arzanlou et al. (2007), Videira et
(2017)
Arzanlou et al. (2007), Crous et
(2009b), Videira et al. (2017)
Quaedvlieg et al. (2014)
Crous et al. (2017a)
Guatimosim et al. (2016)
Crous et al. (2017b)
Crous et al. (2009a), Quaedvlieg et
(2014), Videira et al. (2017)
Shivas et al. (2011)
Present study
Videira et al. (2017)
Videira et al. (2017)
Crous et al. (2014c), Guatimosim et
(2016)
Quaedvlieg et al. (2014), Videira et
(2017)
Huang et al. (2015), Videira et
(2017)
Huang et al. (2015), Videira et
al.
al.
al.
al.
al.
al.
al.
al.
al.
ACCEPTED MANUSCRIPT
T
CBS 143422
CBS 124107T
MG386050 MG386103 –
FJ839634
FJ839670
MF951705
–
–
–
–
–
–
Z. gupoyu
Z. hakeae
Z. indonesianum
CBS 122099
CBS 142185T
CBS 139627T
MF951401
MF951402
KF901739
MF951267
MF951268
KF902086
MF951706
MF951707
MF951710
–
–
–
–
–
KF903377
–
–
–
Z. iteae
Z. lonicericola
CBS 113094T
CBS 125008ET
MF951405
KF251283
MF951271
KF251787
MF951711
MF951712
–
–
–
KF253231
–
KF252765
Z. macluricola
Z. musae
Z. musae-banksii
BRIP 52143T
CBS 122477T
CBS 121710T
GU108499
EU514291
EU041795
–
–
EU041852
–
–
MF951716
–
EU514346
–
–
–
–
–
–
–
Z. musicola
CBS 122479T
EU514294
MF951275
MF951717
–
–
–
Z. musigenum
Z. nocoxi
CBS 365.36T
CBS 125009T
EU041801
KF251284
EU041858
KF251788
–
MF951719
–
–
–
KF253232
–
KF252766
Z. pitospori
Z. podocarpi
Z. proteacearum
CBS 122274
CBS 142529T
CBS 116003
MF951406
KY979766
FJ839635
MF951276
KY979821
FJ839671
MF951720
–
MF951721
–
KY979861
KF903478
–
–
–
–
KY979930
KF903070
Z. pseudoparkii
CBS 110999T
DQ303023
JF700965
MF951723
KF903419
KF903273
KF902977
Z. pseudotsugae
Z. pseudovespa
rapssd
CBS 121159T
EF114687
MF951407
EF114704
KF901836
–
MF951724
–
–
–
KF902812
Z. queenslandicum
CBS 122475T
EU514295
MF951277
MF951725
–
–
–
Z. rothmanniae
Z. scaevolicola
CBS 137983T
CBS 127009T
KJ869135
KF251285
MH878613 –
KF251789 MF951726
–
–
–
KF253233
–
KF252767
SC
M
AN
U
D
TE
EP
AC
C
RI
PT
Z. gahniicola
Z. grevilleae
(2017)
Crous et al. (2017b)
Crous et al. (2009a), Videira et al.
(2017)
Videira et al. (2017)
Videira et al. (2017)
Quaedvlieg et al. (2014), Videira et al.
(2017)
Videira et al. (2017)
Quaedvlieg et al. (2013), Videira et al.
(2017)
Shivas et al. (2009)
Arzanlou et al. (2008)
Arzanlou et al. (2007), Videira et al.
(2017)
Arzanlou et al. (2008), Videira et al.
(2017)
Arzanlou et al. (2007)
Quaedvlieg et al. (2013), Videira et al.
(2017)
Videira et al. (2017)
Crous et al. (2017a)
Crous et al. (2009a), Quaedvlieg et al.
(2014), Videira et al. (2017)
Crous et al. (2006), Quaedvlieg et al.
(2011, 2013), Videira et al. (2017)
Winton et al. (2007)
Quaedvlieg et al. (2013, 2014), Videira
et al. (2017)
Arzanlou et al. (2008), Videira et al.
(2017)
Crous et al. (2014c), Vu et al. (2019)
Quaedvlieg et al. (2013), Videira et al.
ACCEPTED MANUSCRIPT
T
CBS 142188
CBS 121711T
MF951408
EU041803
MF951278
EU041860
MF951727
MF951729
–
–
–
–
–
–
Z. suregadae
Z. syzygii
MFLU 12-2212HT
CBS 133580T
KC677914
KC005777
KC677939
KC005798
–
MF951730
–
–
–
–
–
–
Z. thailandicum
Z. tsugae
Z. velutinum
CBS 145027T
ratstk
CBS 101948ET
MK539970 MK540040 –
EF114688 EF114705 –
EU041781 EU041838 MF951731
–
–
–
–
–
–
Z. xenoparkii
CBS 111185T
DQ303028
KF903438
SC
MF951732
M
AN
U
JF700966
1
RI
PT
Z. schini
Z. strelitziae
KF903274
–
–
–
KF902978
(2017)
Videira et al. (2017)
Arzanlou et al. (2007), Videira et
(2017)
Phengsintham et al. (2013)
Crous et al. (2012), Videira et
(2017)
Present study
Winton et al. (2007)
Arzanlou et al. (2007), Videira et
(2017)
Crous et al. (2006), Quaedvlieg et
(2011, 2014), Videira et al. (2017)
al.
al.
al.
al.
AC
C
EP
TE
D
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture
collection of Pedro Crous, housed at the Westerdijk Fungal Biodiversity Institute; MFLU: Mae Fah Luang University herbarium, Chiang Rai, Thailand; rapssd
and ratstk were not specified in the original publications. T, ET, HT and NT indicate ex-type, ex-epitype, holotype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; rpb2: partial DNA-directed RNA polymerase II
second largest subunit gene; act: partial actin gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
ACCEPTED MANUSCRIPT
Fig. 1. Allophoma spp. A, B. Disease symptoms. A. Symptoms caused by Allophoma hayatii
(ex-type CBS 142859) on Lantana camara. B. Symptoms caused by Allophoma
pterospermicola (ex-type CGMCC 3.19245) on Pterospermum xylocarpum. C–O. Asexual
morph. C. Conidiomata of Allophoma pterospermicola (LC12181) sporulating on Maesa
montana. D. Conidiomata of Allophoma oligotrophica (ex-type CGMCC 3.18114) sporulating
on OA. E. Conidiomata of Allophoma minor (ex-type CBS 325.82). F. Section of the
RI
PT
conidiomatal wall of Allophoma minor (ex-type CBS 325.82). G, H. Conidiogenous cells. G.
Allophoma piperis (ex-epitype CBS 268.93). H. Allophoma oligotrophica (ex-type CGMCC
3.18114). I–M. Conidia. I. Allophoma minor (ex-type CBS 325.82). J. Allophoma piperis (exepitype CBS 268.93). K. Allophoma oligotrophica (ex-type CGMCC 3.18114). L. Allophoma
M
AN
US
C
cylindrispora (ex-type CBS 142453). M. Allophoma nicaraguensis (ex-type CBS 506.91). N.
Swollen cells of Allophoma hayatii (ex-type CBS 142859). O. Chlamydospores of Allophoma
hayatii (ex-type CBS 142859). Scale bars: D, E = 100 m; O = 50 m; N = 20 m; F, I–M = 10
m; G, H = 5 m. Pictures A, N–O taken from Babaahmadi et al. (2018); D, H, K, from Chen et
al. (2017); E, F, I from Aveskamp et al. (2010); G, J, M from Chen et al. (2015); L from
Valenzuela-Lopez et al. (2018).
Fig. 2. Phylogenetic tree generated from a maximum parsimony analysis based on the combined
LSU (860 bp), ITS (480 bp), tub2 (333 bp) and rpb2 (803 bp) sequences of all accepted species
D
of Allophoma. The tree was rooted to Phoma herbarum CBS 615.75. Values above the branches
represent parsimony bootstrap support values (> 50%). Novel sequences and novel taxon are
TE
printed in bold. GenBank accession numbers are indicated in Table 1. T, IsoT and NT indicate ex-
EP
type, ex-isotype and ex-neotype strains, respectively. TreeBASE: S23493.
Fig. 3. Allophoma pterospermicola (ex-type CGMCC 3.19245). A, B. Colony on OA (front and
reverse). C, D. Colony on MEA (front and reverse). E, F. Colony on PDA (front and reverse).
AC
C
G. Conidiomata sporulating on OA. H, I. Conidiomata. J. Section of conidioma. K. Section of
conidiomatal wall. L, M. Conidiogenous cells. N. Conidia. Scale bars: H, J = 20 m; I = 40 m;
K, N = 10 m; L, M = 5 m.
Fig. 4. Alternaria spp. A–D. Disease symptoms. A. Alternaria dauci on Daucus carota. B.
Alternaria linariae on Solanum lycopersicum. C. Alternaria neoipomoeae on Ipomoeae batatas
(Photo A.H. Thompson, ARC, South Africa). D. Alternaria solani on Solanum tuberosum
(Photo J.E. van der Waals, University of Pretoria, South Africa). E–V. Asexual morph. E–O.
Conidiophores. E. Alternaria caricis. F. Alternaria chartarum. G. Alternaria cinerariae. H.
Alternaria conjuncta. I. Alternaria elegans. J. Alternaria embellisia. K. Alternaria indefessa. L.
Alternaria japonica. M. Alternaria penicillata. N. Alternaria proteae. O. Alternaria tenuissima.
ACCEPTED MANUSCRIPT
P–T. Conidia. P. Alternaria blumeae. Q. Alternaria calendulae. R. Alternaria perpunctulata. S.
Alternaria carotiincultae. T. Alternaria triglochinicola. U, V. Conidia producing secondary
conidia. U. Alternaria mimicola. V. Alternaria molesta. Scale bars: 10 µm. Pictures A–D, P, Q
taken from Woudenberg et al. (2014); E–O, R–V from Woudenberg et al. (2013).
Fig. 5. Maximum Likelihood (ML) tree constructed with ITS (529 bp), ATPase (1180 bp),
RI
PT
gapdh (489 bp), rpb2 (573 bp) and tef1 (239 bp) sequences of ex-type strains of the species in
section Infectoriae. The phylogenetic tree was rooted to Alternaria abundans CBS 534.83 and
Alternaria breviramosa CBS 121331 (section Chalastospora). Bootstrap support values above
70 % are shown at the nodes. GenBank accession numbers are indicated in Table 2. The novel
T
indicates ex-type strain. TreeBASE:
M
AN
US
C
species described in this study are indicated in bold.
S23786.
Fig. 6. Maximum Likelihood (ML) tree constructed with ITS (576 bp), ATPase (1198 bp) and
gapdh (491 bp) sequences of ex-type strains of species in the sections Pseudoalternaria and
Chalastospora. The phylogenetic tree was rooted to Alternaria caricis CBS 480.90 and A.
scirpicola CBS 481.90 (section Nimbya). Bootstrap support values above 70 % are shown at the
nodes. GenBank accession numbers are indicated in Table 2. Names of species newly described
here are indicated in bold. The novel species described in this study are indicated in bold.
T
D
indicates ex-type strain. TreeBASE: S23787.
TE
Fig. 7. Maximum Likelihood (ML) tree constructed with ITS (523 bp), gapdh (503 bp), rpb2
(860 bp) and tef1 (247 bp) sequences of ex-type strains of species in section Radicina. The
EP
phylogenetic tree was rooted to Alternaria caricis CBS 480.90 and A. scirpicola CBS 481.90
(section Nimbya). Bootstrap support values above 70 % are shown at the nodes. GenBank
accession numbers are indicated in Table 2. Names of species newly described here are
AC
C
indicated in bold. The novel species described in this study are indicated in bold. T indicates extype strain. TreeBASE: S23788.
Fig. 8. Alternaria aconidiophora (ex-type FMR 17111). A. Colonies on PDA. B. Colonies on
PCA. C. Colonies on OA. D–F. Conidia. Scale bars = 10 µm.
Fig. 9. Alternaria altcampina (ex-type FMR 16476). A. Colonies on PDA. B. Colonies on PCA.
C. Colonies on OA. D–I. Conidiophores and conidia. Scale bars: D–E = 20 µm; F–I = 10 µm.
ACCEPTED MANUSCRIPT
Fig. 10. Alternaria chlamydosporifera (ex-type FMR 17360). A. Colonies on PDA. B. Colonies
on PCA. C. Colonies on OA. D–H. Conidiophores and conidia. I. Chlamydospores. Scale bars:
D = 50 µm; E, I = 20 µm; F–H = 10 µm.
Fig. 11. Alternaria curvata (ex-type FMR 16901). A. Colonies on PDA. B. Colonies on PCA.
RI
PT
C. Colonies on OA. D–F. Conidiophore and conidia. Scale bars = 10 µm.
Fig. 12. Alternaria fimeti (ex-type FMR 17110). A. Colonies on PDA. B. Colonies on PCA. C.
Colonies on OA. D–G. Conidiophores and conidia. Scale bars: D = 20 µm; E–G = 10 µm.
M
AN
US
C
Fig. 13. Alternaria inflata (ex-type FMR 16477). A. Colonies on PDA. B. Colonies on PCA. C.
Colonies on OA. D, F–G. Conidiophores and conidia. E. Chlamydospores. Scale bars = 10 µm.
Fig. 14. Alternaria lawrencei (ex-type FMR 17004). A. Colonies on PDA. B. Colonies on PCA.
C. Colonies on OA. D–K. Conidiophores and conidia. Scale bars: D = 20 µm; E–K = 10 µm.
Fig. 15. Alternaria montsantina (ex-type FMR 17060). A. Colonies on PDA. B. Colonies on
PCA. C. Colonies on OA. D–H. Conidiophores and conidia. Scale bars: D = 50 µm; E–H = 10
D
µm.
Fig. 16. Alternaria pobletensis (ex-type FMR 16448). A. Colonies on PDA. B. Colonies on
TE
PCA. C. Colonies on OA. D–I. Conidiophores and conidia. Scale bars: D = 20 µm; E–I = 10
EP
µm.
Fig. 17. Alternaria pseudoventricosa (ex-type FMR 16900). A. Colonies on PDA. B. Colonies
on PCA. C. Colonies on OA. D–F. Conidiophores and conidia. Scale bars: D = 20 µm; E, F =
AC
C
10 µm.
Fig. 18. Brunneosphaerella spp. A–E. Disease symptoms. A, B. Brunneosphaerella protearum
(epitype CBS H-20335). C. Brunneosphaerella nitidae (holotype CBS H-20334). D, E.
Ascomata visible on lesion surface. D. Brunneosphaerella protearum (CBS H-20335). E.
Brunneosphaerella nitidae (holotype CBS H-20334). F–H. Vertical sections through ascomata
showing wall structure. F. Brunneosphaerella jonkershoekensis (holotype PREM 59447). G,
H. Brunneosphaerella protearum (CBS H-20335).
I–M. Asci. I. Brunneosphaerella
jonkershoekensis (holotype PREM 59447). J, K. Brunneosphaerella nitidae (holotype CBS H20334). L, M. Brunneosphaerella protearum (CBS H-20335). N–Q. Ascospores. N, O.
Brunneosphaerella jonkershoekensis (holotype PREM 59447). P. Brunneosphaerella nitidae
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(holotype CBS H-20334). Q. Brunneosphaerella protearum (CBS H-20335).
R, S.
Germinating ascospores. R. Brunneosphaerella nitidae (holotype CBS H-20334). S.
Brunneosphaerella protearum (CBS H-20335). Scales bars: G = 75 µm; F, I = 50 µm; H, J–
M, P–S = 10 µm; N, O = 5 µm. Pictures A, B, D, F–I, L–O, Q, S taken from Crous et al.
(2009b); C, E, J, K, P, R from Crous et al. (2011).
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Fig. 19. Brunneosphaerella roupeliae (ex-type CBS 144602). A. Close-up of leaf spot with
ascomata. B, C. Asci with ascospores. D. Germinating ascospores. Scale bars = 10 µm.
Fig. 20. Disease symptoms of Elsinoe spp. A. Elsinoe ampelina on Vitis vinifera. B. Elsinoe
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asclepiadea on Asclepias mellodora. C. Elsinoe bidentis on Bidens segetum. D. Elsinoe
erythrinae on Erythrina sp. E. Elsinoe eucalypticola on Eucalyptus sp. F. Elsinoe fawcettii on
Citrus sp. G. Elsinoe freyliniae on Freylinia lanceolata. H. Elsinoe perseae on Persea
americana. I. Elsinoe othonnae on Othonna quinquedentata. J. Elsinoe poinsettiae on
Euphorbia sp. K. Elsinoe punicae on Punica granatum. L. Elsinoe terminaliae on Terminalia
catappa. Pictures taken from Fan et al. (2017).
Fig. 21. Elsinoe spp. A–J. Sexual morph. A. Subcutilar ascoma of Elsinoe fecunda. B–D. Asci.
B. Elsinoe eucalypticola (ex-type CBS 124765). C, D. Elsinoe fecunda (holotype, PREM
D
56503). E–J. Ascospores. E–H. Elsinoe eucalypticola (ex-type CBS 124765). I, J. Elsinoe
fecunda. K–P. Asexual morph. K, L. Conidiophores. K. Elsinoe asclepiadea (ex-type CPC
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18544). L. Elsinoe othonnae (ex-type CBS 139910). M–P. Conidia. M. Elsinoe asclepiadea
(ex-type CPC 18544). N. Elsinoe erythrinae (ex-epitype CPC 18542). O. Elsinoe tectificae (ex-
EP
type CBS 124777). P. Elsinoe othonnae (ex-type CBS 139910). Scale bars: A = 100 m; B–D,
K–U = 10 m; E–J = 5 m; C applies to C and B; J applies I and J. Pictures taken from Fan et
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al. (2017).
Fig. 22. RAxML phylogram obtained from the combined ITS (609 bp), LSU (741 bp), rpb2 (747
bp) and tef1 (422 bp) sequence alignment of all accepted species of Elsinoe. The tree was rooted
to Myriangium hispanicum CBS 247.33. The novelties proposed in this study are indicated in
bold. RAxML bootstrap support (BS) values above 70 % are shown at the nodes. GenBank
accession numbers are listed in Table 4. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype
strains, respectively. TreeBASE: S23834.
Fig. 23. Elsinoe picconiae (ex-type CBS 145026). A. Colony on OA. B, C. Conidiogenous
cells. D. Conidia. Scale bars = 10 m.
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Fig. 24. Elsinoe veronicae (ex-type CBS 145362). A. Colony on OA. B, C. Conidiogenous
cells. D. Conidia. Scale bars = 10 m.
Fig. 25. Exserohilum spp. A–E. Sexual morph. A. Ascomata of Exserohilum minor (ex-isotype
IMI 294530). B, C. Asci of Exserohilum minor (ex-isotype IMI 294530). D, E. Ascospores. D.
Exserohilum minor (ex-isotype IMI 294530). E. Exserohilum khartoumensis (ex-isotype CBS
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132708). F–AA. Asexual morph. F–N. Conidiophores and conidia. F, L. Exserohilum oryzicola
(ex-isotype CBS 502.90). G. Exserohilum turcicum (ex-epitype CBS 690.71). H, N.
Exserohilum holmii (ex-isotype CBS 413.65 and BRIP 12679). I. Exserohilum pedicellatum
(CBS 375.76). J, M. Exserohilum rostratum (CBS 120380, CBS 196.29). K. Exserohilum
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monoceras (CBS 198.29). O. Detail of the conidial hilum of Exserohilum oryzicola (ex-isotype
CBS 502.90). P–AA. Conidia. P, Q, Z, AA. Exserohilum rostratum (CBS 128054, CBS
120380, BRIP 11422). R. Exserohilum holmii (BRIP 12679). S. Exserohilum pedicellatum
(BRIP 12040). T, U. Exserohilum turcicum (BPI 431157 holotype). V. Exserohilum oryzicola
(BRIP 16229). W. Exserohilum protrudens (BRIP 14816). X. Exserohilum corniculatum (extype BRIP 11426). Y. Exserohilum neoregelia (CBS 132833). Scale bars A = 50 m; others =
10 m; C applies to B and C; E applies to D and E. Pictures taken from Hernández-Restrepo et
al. (2018).
D
Fig. 26. RAxML phylogram obtained from the combined ITS (793 bp), gapdh (578 bp) and
rpb2 (860 bp) sequence alignment of all the accepted species of Exserohilum. The tree was
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rooted to Curvularia and Bipolaris. RAxML bootstrap support (BS) values above 70 % are
shown in the nodes. GenBank accession numbers are indicated in Table 5. T, ET, IsoT and A indicate
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ex-type, ex-epitype, ex-isotype, and authentic strains. TreeBASE: S23834.
Fig. 27. Neosetophoma lunariae (ex-type CBS 141409). A. Conidiomata on OA. B. Conidiomata
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showing ostiolar region. C, D. Conidiogenous cells. E. Conidia. Scale bars = 10 m; C applies to
C and D. Pictures taken from Hernández-Restrepo et al. (2016a).
Fig. 28. RAxML phylogram obtained from the combined ITS (626 bp) and LSU (852 bp)
sequence alignment of members of the family Phaeosphaeriaceae. The tree was rooted to
Coniothyrium glycines CBS 124141 and Coniothyrium sidae CBS 135108. The novelties
proposed in this study are indicated in bold. RAxML bootstrap support (BS) values above 70 %
and Bayesian posterior probability scores above 0.95 are shown at the nodes. Numbers between
parentheses correspond to GenBank accession numbers for ITS and LSU sequences,
respectively.
T, ET, HT, IsoT, LT
and
NT
indicate ex-type, ex-epitype, holotype, ex-isotype, ex-
lectotype and ex-neotype strains, respectively. TreeBASE: S23834.
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Fig. 29. RAxML phylogram obtained from the combined ITS (585 bp), LSU (848 bp) and rpb2
(744 bp) sequence alignment of all accepted species of Neosetophoma. The tree was rooted to
Phaeosphaeriopsis glaucopunctata MFLUCC 13-0265 and Phaeosphaeriopsis agavacearum
CPC 29122. The novelties proposed in this study are indicated in bold. RAxML bootstrap
support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are
loci in Table 6 and Table 10.
T
and
ET
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shown at the nodes. GenBank accession numbers of LSU are listed in Fig. 28, and of the other
indicate ex-type and ex-epitype strains, respectively.
TreeBASE: S23834.
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Fig. 30. Neosetophoma aseptata (ex-type CBS 145363). A. Conidiomata sporulating on OA. B,
C. Conidiogenous cells. D. Conidia. Scale bars: A = 300 m; others = 10 m.
Fig. 31. Neosetophoma phragmitis (ex-type CBS 145364). A. Conidiomata sporulating on SNA.
B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 200 m; others = 10 m.
Fig. 32. Neosetophoma sambuci (ex-type CBS 145365). A. Conidiomata sporulating on PNA. B.
Conidioma on SNA showing papillate neck. C. Conidiogenous cells. D. Conidia. Scale bars: A, B
D
= 200 m; others = 10 m.
Fig. 33. Neostagonospora spp. A, B. Conidioma forming in culture. A. Neostagonospora
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caricis (ex-type CBS 135092). B. Neostagonospora elegiae (ex-type CBS 135101). C–G.
Conidiogenous cells. C, D. Neostagonospora caricis (ex-type CBS 135092). E–G.
EP
Neostagonospora elegiae (ex-type CBS 135101). H, I. Conidia. H. Neostagonospora caricis
(ex-type CBS 135092). I. Neostagonospora elegiae (ex-type CBS 135101). Scale bars: B = 150
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µm; all others = 10 µm; C applies to C and D. Pictures taken from Quaedvlieg et al. (2013).
Fig. 34. RAxML phylogram obtained from the combined ITS (571 bp), LSU (847 bp), rpb2
(337 bp) and tub2 (304 bp) sequence alignment of all accepted species of Neostagonospora.
The tree was rooted to Parastagonospora avenae CBS 289.69 and Parastagonospora
nodorum CBS 110109. The novelty proposed in this study is indicated in bold. RAxML
bootstrap support (BS) values above 70 % and Bayesian posterior probability scores above
0.95 are shown at the nodes. GenBank accession numbers of LSU are listed in Fig. 28, and of
the other loci in Table 7 and Table 9. T indicates ex-type strains. TreeBASE: S23834.
Fig. 35. Neostagonospora sorghi (ex-type CBS 145366). A. Conidiomata sporulating on MEA.
B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 200 m; others = 10 m.
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Fig. 36. Nothophoma spp. A, B. Disease symptoms. A. Nothophoma quercina (CGMCC
3.19246) on Osmanthus fragrans. B. Nothophoma quercina (LC12187) on Jasminum mesnyi.
C–I. Asexual morph. C, D. Conidiomata of Nothophoma anigozanthi (ex-epitype CBS 381.91)
sporulating on OA. E–H. Conidia. E. Nothophoma infossa (ex-neotype CBS 123395). F.
Nothophoma macrospora (ex-type CBS 140674). G. Nothophoma quercina (CGMCC 3.19246).
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H. Nothophoma variabilis (ex-type CBS 142457). I. Conidiogenous cells of Nothophoma
macrospora (ex-type CBS 140674). Scale bars: C = 200 m; D = 20 m; E–I = 10 m. Pictures
C, D taken from Chen et al. (2015); E from Aveskamp et al. (2009); F, I from Crous et al.
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(2016b); H from Valenzuela-Lopez et al. (2018).
Fig. 37. Phylogenetic tree generated from a maximum parsimony analysis based on the
combined LSU (868 bp), ITS (490 bp), tub2 (336 bp) and rpb2 (845 bp) sequences of all
accepted species of Nothophoma. The tree was rooted to Phoma herbarum CBS 615.75. Values
above the branches represent parsimony bootstrap support values (> 50%). GenBank accession
numbers are indicated in Table 8.
T, ET
strains, respectively. TreeBASE: S23494.
and
NT
indicate ex-type, ex-epitype and ex-neotype
Fig. 38. Parastagonospora spp. A–D. Sexual morph of Parastagonospora nodorum (CBS H-
D
13909). A, B. Ascomata. C, D. Asci and ascospores. E–M. Asexual morph. E, F.
Conidiomata. E. Parastagonospora poagena (ex-type CBS 136776). F. Parastagonospora
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poae (CBS 135091). G–I. Conidiogenous cells. G. Parastagonospora caricis (ex-type CBS H21304). H. Parastagonospora poae (CBS 135091). I. Parastagonospora poagena (ex-type
EP
CBS 136776). J–M. Conidia. J. Parastagonospora caricis (ex-type CBS H-21304). K.
Parastagonospora nodorum (CBS H-13909). L. Parastagonospora poae (CBS 135091). M.
Parastagonospora poagena (ex-type CBS 136776). Scale bars: C, D, J–M = 10 m; G–I = 5
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m. Pictures A–D, F–H, J–L taken from Quaedvlieg et al. (2013); E, I, M from Crous et al.
(2014b).
Fig. 39. RAxML phylogram obtained from the combined ITS (575 bp), LSU (848 bp), rpb2
(337 bp) and tef1 (866 bp) sequence alignment of all accepted species of Parastagonospora.
The tree was rooted to Neostagonospora carici CBS 135092 and Neostagonospora elegiae
CBS 135101. The novelties proposed in this study are indicated in bold. RAxML bootstrap
support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are
shown at the nodes. GenBank accession numbers of LSU are listed in Fig. 28, and of the others
loci are indicated in Table 7 and Table 9. T indicates ex-type strains. TreeBASE: S23834.
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Fig. 40. Parastagonospora novozelandica (ex-type CPC 29613). A–C. Conidiogenous cells.
D. Conidia. Scale bars = 10 m.
Fig. 41. Parastagonospora phragmitis (ex-type CPC 32075). A, B. Conidiogenous cells. C, D.
Conidia. Scale bars = 10 m.
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Fig. 42. Phaeosphaeriopsis spp. A. Symptomatic leaves of Agapanthus precox caused by
Phaeosphaeriopsis agapanthi. B. Asci and ascospores of Phaeosphaeriopsis agavacearum
(ex-type CBS 142110). C–P. Asexual morph. C, D. Conidiomata sporulating on PNA and
OA, respectively, of Phaeosphaeriopsis agapanthi (ex-type CBS 141287). E. Conidiomata
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sporulating on OA of Phaeosphaeriopsis agavacearum (ex-type CBS 142110). F. Conidioma
of Phaeosphaeriopsis agavacearum (ex-type CBS 142110). G–L. Conidiogenous cells giving
rise to conidia. G, H. Phaeosphaeriopsis agapanthi (ex-type CBS 141287). I–L.
Phaeosphaeriopsis glaucopunctata (CBS 653.86). M–O. Conidia. M. Phaeosphaeriopsis
agapanthi (ex-type CBS 141287). N. Phaeosphaeriopsis agavacearum (ex-type CBS 142110).
O.
Phaeosphaeriopsis
glaucopunctata
(CBS
653.86).
P.
Chlamydospores
of
Phaeosphaeriopsis agavacearum (ex-type CBS 142110). Scale bars: F = 100 m; others = 10
m; I applies to I–L. Pictures A, C, D, G, H, M taken from Crous et al. (2016b); B, E, F, N, P
D
from Crous et al. (2016a); I–L, O from Quaedvlieg et al. (2013).
Fig. 43. RAxML phylogram obtained from the combined ITS (587 bp), LSU (849 bp), rpb2
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(838 bp), tef1 (601 bp) and tub2 (519 bp) sequence alignment of all accepted species of
Phaeosphaeriopsis. The tree was rooted to Neostagonospora caricis CBS 135092 and
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Neostagonospora elegiae CBS 135101. The novelties proposed in this study are indicated in
bold. RAxML bootstrap support (BS) values above 70 % and Bayesian posterior probability
scores above 0.95 are shown at the nodes. GenBank accession numbers of LSU are listed in
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Fig. 28, and of the other loci in Table 7 and Table 10.
T
and
ET
indicate ex-type and ex-
epitype strains, respectively. TreeBASE: S23834.
Fig. 44. Phaeosphaeriopsis aloes (ex-type CBS 145367). A. Conidiomata sporulating on
MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 180 µm; all others = 10 m.
Fig. 45. Phaeosphaeriopsis aloicola (ex-type CBS 145368). A. Ascomata sporulating on SNA.
B. Asci. C. Pseudoparaphyses. D. Ascospores. Scale bars: A = 200 µm; all others = 10 m.
Fig. 46. Phaeosphaeriopsis grevilleae (ex-type CBS 145369). A. Conidiomata sporulating on
PDA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 m.
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Fig. 47. Phaeosphaeriopsis pseudoagavacearum (ex-type CBS 145370). A. Conidiomata
sporulating on MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 m.
Fig. 48. Pleiocarpon strelitziae (ex-type CBS 142251). A–F. Disease symptoms. A, B.
Wilting and dying Strelitzia reginae plants in the nursery. C–F. Basal rot and wilting of plant
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induced during the pathogenicity test. G–L. Asexual morph. G, H. Simple conidiophores. I,
J. Sporodochia. K. Microconidia. L. Macroconidia. Scale bars: 10 µm; G applies to G–L.
Pictures taken from Aiello et al. (2017).
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Fig. 49. Pleiocarpon livistonae (ex-type CBS 145030). A. Sporodochium on SNA. B.
Conidiophores with conidiogenous cells. C. Conidia. D. Chlamydospores. Scale bars = 10 m.
Fig. 50. Pyrenophora spp. A–D. Sexual morph. A, B. Sterile ascomata of Pyrenophora
campanulata (CBS 127927). C. Protoascomata of Pyrenophora erythrospila on PDA (CBS
312.69). D. Protoascoma of Pyrenophora erythrospila (CBS 108941). E–K. Asexual morph.
E–H. Conidiophores. E. Pyrenophora fugax (CBS 509.77). F. Pyrenophora novozelandica
(CBS 127934). G. Pyrenophora erythrospila (CBS 312.69). H. Pyrenophora fugax (CBS
509.77). I, J. Conidia. I. Pyrenophora erythrospila (CBS 312.69). J. Pyrenophora fugax (CBS
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m; D = 20 m; G–K = 10 m.
D
509.77). K. Chlamydospores of Pyrenophora tetrarrhenae (CBS 127924). Scale bars: A = 50
Fig. 51. RAxML phylogram obtained from the combined ITS (788 bp), LSU (862 bp), gapdh
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(694 bp) and tef1 (860 bp), sequences of all the accepted species of Pyrenophora. Bipolaris
panici-miliacei CBS 199.29 and Bipolaris yamadae CBS 202.29 were used as outgroup. The
novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS) values
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above 70 % and Bayesian posterior probability scores ≥ 0.95 are shown in the nodes. GenBank
accession numbers were indicated in Table 12 and Manamgoda et al. (2014). A, ET, LT, SynT and T
indicate authentic, ex-epitype, ex-lectotype, ex-syntype and ex-type strains, respectively.
TreeBASE: S23834.
Fig. 52. Pyrenophora avenicola (ex-type CBS 307.84). A, B. Conidiophores and conidia. C–I.
Conidia. Scale bars: A, B = 10 m; C–I = 5 m; C applies to C–I.
Fig. 53. Pyrenophora cynosuri (ex-type CBS 127918). A–C. Conidiophores and conidia. D.
Conidium forming secondary conidium. E–M. Conidia. Scale bars: 10 m; E applies to E–M.
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Fig. 54. Pyrenophora nisikadoi (CBS 119213). A, B. Conidiophores and conidia. C.
Chlamydospores. D–K. Conidia. Scale bars: A, B = 10 m; C = 20 m; D = 5 m; D applies to
D–K.
Fig. 55. Pyrenophora novozelandica (ex-type CBS 127934). A–D. Conidiophores and conidia.
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E–M. Conidia. Scale bars: A–D = 10 m; E = 5 m; E applies to E–M.
Fig. 56. Pyrenophora pseudoerythrospila (ex-type CBS 127931). A. Protoascomata on OA. B,
C. Protoascomata. Scale bars = 10 m.
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Fig. 57. Pyrenophora sieglingiae (ex-type CBS 127930). A–C. Sterile ascomata. D. Neck of
ascoma. E, F. Conidiophores and conidia. G–O. Conidia. Scale bars: C = 100 m; D–F = 20
m; G = 10 m; G applies to G–O.
Fig. 58. Pyenophora variabilis (ex-type CBS 127920). A–C. Conidiophores and conidia. D–
N. Conidia. Scale bars = 10 m; C applies to A–C; N applies to D–N.
Fig. 59. Ramichloridium spp. A. Ramichloridium luteum on apple. B. Sporulating colonies of
Ramichloridium luteum (ex-type CBS 132088) on PDA. C–G. Macronematous conidiophores
D
with sympodially proliferating conidiogenous cells, which give rise to a conidium-bearing rachis
with crowded and prominent scars. C. Ramichloridium apiculatum (ex-type CBS 156.59). D.
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Ramichloridium cucurbitae (ex-type CBS 132087). E, F. Ramichloridium luteum (ex-type CBS
132088). G. Ramichloridium punctatum (ex-type CBS 132090). H, I. Scanning electron
EP
micrographs of Ramichloridium luteum (ex-type CBS 132088) showing sympodial proliferation
with scars on conidiogenous cells. J, K. Conidiophores reduced to conidiogenous cells. J.
Ramichloridium cucurbitae (ex-type CBS 132087). K. Ramichloridium luteum (ex-type CBS
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132088). L, M. Conidia. L. Ramichloridium apiculatum (ex-type CBS 156.59). M.
Ramichloridium punctatum (ex-type CBS 132090). Scale bars: H = 2 m; I = 1 m; all others =
10 m. Pictures C, L taken from Li et al. (2012); all others from Arzanlou et al. (2007).
Fig. 60. RAxML phylogram obtained from the combined ITS (594 bp), LSU (761 bp), rpb2 (819
bp) and tef1 (470 bp) sequence alignment of all accepted species of Ramichloridium and related
taxa. The tree was rooted to Parapenidiella tasmaniensis CBS 124991 and Stenella araguata CBS
105.75. The novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS)
values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes.
GenBank accession numbers are listed in Table 13, Li et al. (2012), Videira et al. (2017) and Chen
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& Kirschner (2018). T and NT indicate ex-type and ex-neotype strains, respectively. TreeBASE:
S23834.
Fig. 61. Seifertia azalae. A, B. Disease symptoms caused on Rhododendron. C–G. Synnemata.
H, I. Conidiogenous cells and conidia. J, K. Conidia. Scale bars: F = 100 m; G = 50 m; H–
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K = 10 m.
Fig. 62. Seiridium spp. A–F. Disease symptoms on Cupressaceae hosts. A–C. Flagging of
branches. D. Trunk canker with gummosis. E. Branch canker. F. Conidiomata. G–I. Seiridium
pezizoides (CPC 35011). G, H. Acervuli on Cupressaceae sp. I. Conidial masses on artificial
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media. J, K. Conidiophores and conidiogenous cells. J. Seiridium neocupressi (CBS 142625).
K. Seiridium eucalypti (CBS 343.97). L–R. Conidia. L. Seiridium cardinale (CBS 909.85). M.
Seiridium spyridicola (CBS 142628). N. Seiridium unicorne (CBS 538.82). O. Seiridium
neocupressi (CBS 142625). P. Seiridium eucalypti (CBS 343.97). Q. Seiridium kartense (CBS
142629). R. Seiridium pezizoides (CPC 35011). S–U. Seiridium cupressi (IMI 40096). S, T.
Ascomata. U. Ascospores. Scale bars: F = 2 mm; G, H = 50 m; J–U = 10 m. Pictures J–U
taken from Bonthond et al. (2018).
Fig. 63. A–D. Boxplots of conidial measurement data in µm from S. unicorne and other
D
Cupressaceae pathogens. The boxes show the lower and upper quantiles and whiskers extend
to 1.5x the interquartile range. Except for the new epitype all measurements are adapted from
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Bonthond et al. (2018). The ex-epitype strain (CBS 143871), holotype (IMI 5816) and
reference strain (CBS 538.82) of S. unicorne are highlighted in blue. E. The best Maximum
EP
Likelihood (ML) tree based on four loci (ITS: 616 bp, rpb2: 802 bp, tef1: 633 bp, tub2: 809
bp). Nodes are labelled with ML bootstrap values (BS > 49 %)/Bayesian posterior probabilities
(PP > 0.49) using the same model selection, settings and software as in Bonthond et al. (2018).
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Strains are displayed by number, host and country of collection. GenBank accession numbers
are listed in Bonthond et al. (2018) and in Table 15. T, ET, GT and R indicate ex-type, ex-epitype,
ex-generic type and reference strains, respectively. TreeBASE: S23390.
Fig. 64. Seiridium pezizoides (CBS 145115). A–D. Colony morphology in 90-mm-diam Petri
dishes after 10 d at 22 °C on MEA, SNA, PDA and CMA, respectively. E–K. Conidiomata on
Vitis vinifera. L, M. Conidiophores. N. Conidia. Scale bars: E = 1 mm; F–K = 100 m; L–N =
10 m.
Fig. 65. Seiridium unicorne (ex-epitype CBS 143871). A–D. Colony morphology in 90-mmdiam Petri dishes after 10 d at 22 °C on MEA, SNA, PDA and CMA, respectively. E.
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Symptoms on naturally infected host. F, G. Conidiomata on artificially infected Cupressaceae
sp. H. Sporulation on PDA. I. Conidioma on SNA partially immersed in agar. J–O.
Conidiophores and conidia. P. Conidia. Scale bars: F–H = 100 m; I–P = 10 m.
Fig. 66. Septoriella spp. A. Conidiomata on OA of Septoriella hirta (ex-neotype CBS 536.77).
B. Conidiomata in vivo of Septoriella phragmitis (ex-epitype CBS 140065). C. Conidial cirrhus
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of Septoriella phragmitis (ex-epitype CBS 140065). D. Conidioma of Septoriella hirta (exneotype CBS 536.77). E, F. Section through conidiomata of Septoriella hirta (ex-neotype CBS
536.77). G–I. Conidiogenous cells. G. Septoriella oudemansii (ex-type CBS 138012). H, I.
Septoriella phragmitis (ex-epitype CBS 140065). J. Developing conidia of Septoriella hirta (ex-
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neotype CBS 536.77). K–O. Conidia. K. Septoriella hirta (ex-neotype CBS 536.77). L.
Septoriella oudemansii (ex-type CBS 138012). M. Septoriella poae (ex-type CBS 136766). N,
O. Septoriella phragmitis (ex-epitype CBS 140065). Scale bars: D, E = 100 µm; F = 50 µm; all
others = 10 µm. Pictures A–F, H–K, N, O taken from Crous et al. (2015); G, L from Crous et al.
(2014b).
Fig. 67. RAxML phylogram obtained from the combined ITS (580 bp), LSU (849 bp) and
rpb2 (1083 bp) sequence alignment of all accepted species of Septoriella. The tree was
rooted to Neostagonospora caricis CBS 135092 and Neostagonospora elegiae CBS 135101.
D
The novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS)
values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the
Table 7 and Table 16.
TE
nodes. GenBank accession numbers of LSU are listed in Fig. 28, and of the other loci in
T, ET, HT
and IsoT indicate ex-type, ex-epitype, holotype and ex-isotype
EP
strains, respectively. TreeBASE: S23834.
Fig. 68. Septoriella germanica (ex-type CBS 145372). A. Conidiomata sporulating on PNA. B,
m.
AC
C
C. Conidiogenous cells. D. Conidia with mucoid caps. Scale bars: A = 200 µm; all others = 10
Fig. 69. Septoriella hollandica (ex-type CBS 145374). A. Ascomata sporulating on PNA. B,
C. Asci. D. Ascospores. Scale bars = 10 m.
Fig. 70. Septoriella pseudophragmitis (ex-type CPC 24166). A. Conidiomata sporulating on
MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 250 µm; all others = 10 m.
Fig. 71. Setophoma spp. A, B. Conidioma forming in culture. A. Setophoma chromolaenae (extype CBS 135105). B. Setophoma vernoniae (ex-type CBS 137988). C–E. Conidiomata with
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setae of Setophoma chromolaenae (ex-type CBS 135105). F–H. Conidiogenous cells. F, G.
Setophoma chromolaenae (ex-type CBS 135105). H. Setophoma vernoniae (ex-type CBS
137988). I, J. Conidia. I. Setophoma chromolaenae (ex-type CBS 135105). J. Setophoma
vernoniae (ex-type CBS 137988). Scale bars: C–E = 20 µm; all others = 10 µm; F applies to F
and G.
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Fig. 72. RAxML phylogram obtained from the combined ITS (589 bp), LSU (835 bp), tef1 (788
bp) and tub2 (532 bp) sequence alignment of all accepted species of Setophoma. The tree was
rooted to Neostagonospora caricis CBS 135092 and Neostagonospora elegiae CBS 135101.
The novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS)
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C
values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes.
GenBank accession numbers of LSU are listed in Fig. 28, and of the other loci in Table 7, Table
17 and Table 18. T, ET and LT indicate ex-type strains, ex-epitype and ex-lectotype, respectively.
TreeBASE: S23834.
Fig. 73. Setophoma pseudosacchari (ex-type CBS 145373). A. Ascomata sporulating on OA.
B, Asci with ascospores. C. Conidia. Scale bars: A = 300 µm; all others = 10 m.
Fig. 74. Wingfieldomyces cyperi (ex-type CBS 141450). A. Symptomatic leaf of Cyperus
D
sphaerocephala. B–D. Asci. E. Pseudoparaphyses. F–I. Ascospores. Scale bars: 10 µm; F
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applies to F–I. Pictures B, C, E–I taken from Crous et al. (2016b).
Fig. 75. Stagonosporopsis spp. A, B. Disease symptoms of Stagonosporopsis tanaceti (ex-type
EP
CBS 131484). A. Leaf necrosis. B. Drooping flower heads. C–G. Sexual morph of
Stagonosporopsis inoxydabilis (ex-type CBS 425.90). C. Close-up of ascoma with darkened
ostiolar area. D, E. Stipitate, bitunicate asci. F, G. Ascospores (arrows denote sheath). H–T.
AC
C
Asexual morph. H, I. Colony sporulating on OA. H. Stagonosporopsis chrysanthemi (CBS
500.63). I. Stagonosporopsis tanaceti (ex-type CBS 131484). J. Close-up of pycnidial
conidiomata of Stagonosporopsis tanaceti (ex-type CBS 131484). K. Close-up of darkened
ostiolar area of Stagonosporopsis chrysanthemi (CBS 500.63). L–Q. Conidiogenous cells. L–N.
Stagonosporopsis chrysanthemi (CBS 500.63). O–Q. Stagonosporopsis tanaceti (ex-type CBS
131484). R, S. Conidia. R. Stagonosporopsis chrysanthemi (CBS 500.63). S. Stagonosporopsis
tanaceti (ex-type CBS 131484). T. Chain of chlamydospores of Stagonosporopsis tanaceti (extype CBS 131484). Scale bars: C = 35 µm; J = 150 µm; all others = 10 µm; D applies to D and
E; L applies to L–N; O applies to O–Q. Pictures taken from Vaghefi et al. (2012).
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Fig. 76. The majority rule consensus phylogram of Stagonosporopsis spp. inferred from the
concatenated LSU (876 bp), ITS (459 pb), tub2 (299 bp) and rpb2 (596 bp) sequence alignment
using Bayesian Inference. The tree is rooted to Boeremia exigua var. exigua CBS 431.74.
Bootstrap support values > 75% and PP values > 0.90 are shown above or below the branches. T,
ET, IsoT
and
NT
indicate ex-type, ex-epitype, ex-isotype and ex-neotype strains, respectively.
Genbank accession numbers are indicated in Table 19, Aveskamp et al. (2010) and Vaghefi et
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al. (2012). TreeBASE: S23800.
Fig. 77. Stemphylium spp. A. Disease symptoms caused by Stemphylium vesicarium (BRIP
65181) on pyrethrum leaves. B–C. Pseudothecial ascomata of Stemphylium vesicarium (BRIP
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C
65181) on pyrethrum flower stems in vivo and in vitro respectively. D–H. Conidia of
Stemphylium spp. D, H. Stemphylium vesicarium (BRIP 65181). E. Stemphylium truncatulae
(holotype BRIP 14850). F, G. Stemphylium waikerieanum (ex-type VPRI 21969). I–K.
Conidiophores of Stemphylium spp. I. 1-branched conidiophore of Stemphylium vesicarium
(BRIP 65181). J. Simple conidiophore of Stemphylium truncatuale (holotype BRIP 14850). K.
Branched and immature conidiophores of Stemphylium waikerieanum (ex-type VPRI 21969).
L–N. Sexual morphs. L, M. Asci and ascospores of Stemphylium vesicarium isolated from the
dead flower stems of pyrethrum. N. Immature pseudothecial ascomata of Stemphylium
truncatulae on SNA after 1 wk. Scale bars: D–K = 20 µm; L, M = 100 µm. Pictures A–D, H, I,
D
M taken from Moslemi et al. (2017).
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Fig. 78. Bayesian phylogenetic tree inferred from ITS (565 bp), gapdh (613 bp) and cmdA (863)
using partitioned analysis with N92+G+I substituition model for ITS and gapdh and GTR+G+I
EP
for cmdA. Highest log likelihood -34098.05. The analysis involved 122 nucleotide sequences
including 98 sequences obtained from GenBank and 24 sequences obtained in the present study.
Scale bar indicates expected changes per site. The tree was rooted to Alternaria alternata
AC
C
(GV14 634a1). The novel species described in this study are shown in bold. Bayesian posterior
probability scores above 0.95 are shown at the nodes. GenBank accession numbers are indicated
in Table 20.
T, ET, HT
and
NT
indicate ex-type, ex-epitype, ex-holotype and ex-neotype strains,
respectively. TreeBASE:S23794.
Fig. 79. Stemphylium rombundicum (ex-type BRIP 27486). A–J. Conidia. K–S. Verrucose
conidiogenous cells and straight and simple conidiophores with conidia attached. Scale bars: A–
N, P = 20 µm; others 100 µm.
Fig. 80. Stemphylium truncatulae (ex-type BRIP 14850). A–H. Asexual morph. A–D. Conidia
on SNA. E–H. Straight and simple or multibranched conidiophores and conidiogenous cells. I–
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J. Sexual morph. I. Immature pseudothecium. J. Ascomatal wall. Scale bars: I = 100 µm; others
20 µm.
Fig. 81. Stemphylium waikerieanum (ex-type VPRI 21969). A. Dried leaf of Allium cepa
showing leaf lesions caused by the pathogen. B. Dried type culture. C. Revived colony after 1
wk on PDA. D–K. Asexual morph. D–H. Simple or 1-branched conidiophores on PDA and
wk. N. Ascomatal wall. Scale bars: N = 100 µm; others 20 µm.
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SNA. I–K. Phaeodictyospores. L–N. Sexual morph. L–M. Immature ascomata on SNA after 1
Fig. 82. Tubakia spp. A–E. Disease symptoms. A. Tubakia californica on Tanoak tree
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(Notholithocarpus densiflorus). B. Tubakia californica on California black oak (Quercus
kelloggii). C. Tubakia sierrafriensis (holotype CFNL 2944) on Quercus eduardi. D. Tubakia
japonica (epitype NBRC H-11611) on Castanea crenata. E. Tubakia melnikiana (holotype
HAL 3179 F) causing necrotic leaf lesion. F–N. Asexual morph. F. Scutellum of Tubakia
paradryinoides (holotype TFM:FPH 3923). G. Central columella of Tubakia oblongispora
(holotype NBRC H-11881). H–J. Conidiophores. H. Tubakia japonica (epitype NBRC H11611). I. Tubakia oblongispora (holotype NBRC H-11881). J. Tubakia paradryinoides
(holotype TFM:FPH 3923). K–M. Conidia. K. Tubakia dryinoides (holotype NBRC H11618). L. Tubakia oblongispora (holotype NBRC H-11881). M. Tubakia paradryinoides
D
(holotype TFM:FPH 3923). N. Microconidia of Tubakia dryinoides (holotype NBRC H-
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11618). Scale bars: 10 m. Pictures taken from Braun et al. (2018).
Fig. 83. Zasmidium spp. A–C. Disease symptoms caused by Zasmidium cyatheae (ex-type
EP
CPC 24725). A, B. Frond spots on Cyathea delgadii. C. Erumpent subcuticular ascomata,
fruiting epiphyllous. D–F. Sexual morph of Zasmidium cyatheae (ex-type CPC 24725). D.
Ascoma. E. Asci. F. Ascospores. G–R. Asexual morph. G–L. Conidiophores. G. Zasmidium
AC
C
biverticillatum (CBS 335.36). H. Zasmidium citri-griseum (ex-epitype CBS 139467). I.
Zasmidium fructigenum (ex-type CBS 139626). J. Zasmidium indonesianum (ex-type CBS
139627). K. Zasmidium musigenum (ex-type CBS 365.36). L. Zasmidium strelitziae (ex-type
CBS 121711). M–Q. Conidiophores. M. Zasmidium biverticillatum (CBS 335.36). N.
Zasmidium biverticillatum (CBS 335.36). O. Zasmidium citri-griseum (ex-epitype CBS
139467). P. Zasmidium fructicola (ex-type CBS 139625). Q. Zasmidium musigenum (ex-type
CBS 365.36). R. Primary and secondary conidia of Zasmidium cellare (ex-neotype CBS
146.36). Scale bars: 10 m. Pictures A–F taken from Guatimosim et al. (2016); G, K–N, Q, R
from Arzanlou et al. (2007); H–J, O, P from Huang et al. (2015).
Fig. 84. RAxML phylogram obtained from the combined ITS (534 bp), LSU (741 bp) and
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rpb2 (739 bp) sequence alignment of all accepted species of Zasmidium. The tree was rooted to
Anellosympodiella juniperi CBS 137992 and Neopenidiella nectandrae CBS 734.78. The
novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS) values
above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes.
GenBank accession numbers are listed in Table 22 and Videira et al. (2017).
T, ET, HT
and NT
indicate ex-type, ex-epitype, holotype and ex-neootype strains, respectively. TreeBASE:
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S23834.
Fig. 85. Zasmidium thailandicum (ex-type CBS 145027). A–C. Conidiophores sporulating on
AC
C
EP
TE
D
M
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C
SNA. D–F. Conidiogenous cells with apical rachis giving rise to conidia. Scale bars = 10 m.