Accepted Manuscript
Genera of phytopathogenic fungi: GOPHY 2
Y. Marin-Felix, M. Hernández-Restrepo, M.J. Wingfield, A. Akulov, A.J. Carnegie,
R. Cheewangkoon, D. Gramaje, J.Z. Groenewald, V. Guarnaccia, F. Halleen, L.
Lombard, J. Luangsa-ard, S. Marincowitz, A. Moslemi, L. Mostert, W. Quaedvlieg,
R.K. Schumacher, C.F.J. Spies, R. Thangavel, P.W.J. Taylor, A.M. Wilson, B.D.
Wingfield, A.R. Wood, P.W. Crous
PII:
S0166-0616(18)30019-8
DOI:
10.1016/j.simyco.2018.04.002
Reference:
SIMYCO 76
To appear in:
Studies in Mycology
Please cite this article as: Marin-Felix Y, Hernández-Restrepo M, Wingfield MJ, Akulov A, Carnegie AJ,
Cheewangkoon R, Gramaje D, Groenewald JZ, Guarnaccia V, Halleen F, Lombard L, Luangsa-ard J,
Marincowitz S, Moslemi A, Mostert L, Quaedvlieg W, Schumacher RK, Spies CFJ, Thangavel R, Taylor
PWJ, Wilson AM, Wingfield BD, Wood AR, Crous PW, Genera of phytopathogenic fungi: GOPHY 2,
Studies in Mycology (2018), doi: 10.1016/j.simyco.2018.04.002.
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Genera of phytopathogenic fungi: GOPHY 2
1,2*
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Y. Marin-Felix , M. Hernández-Restrepo , M.J. Wingfield , A. Akulov , A.J. Carnegie , R.
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Cheewangkoon , D. Gramaje , J.Z. Groenewald , V. Guarnaccia , F. Halleen , L. Lombard , J. Luangsa9
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ard , S. Marincowitz , A. Moslemi , L. Mostert , W. Quaedvlieg , R.K. Schumacher , C.F.J. Spies , R.
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Thangavel , P.W.J. Taylor , A.M. Wilson , B.D. Wingfield , A.R. Wood , and P.W. Crous
Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands; 2Department of Biochemistry,
Genetics and Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, P. Bag X20, Pretoria
0028, Pretoria, 0002, South Africa; 3V.N. Karasin National University of Kharkiv, Svobody sq. 4, Kharkiv 61077, Ukraine; 4Forest
Science, NSW Department of Primary Industries, Locked Bag 5123, Parramatta, New South Wales 2124; 5Department of Plant
Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200, Thailand; 6Instituto de Ciencias de la Vid y del Vino,
Consejo Superior de Investigaciones Científicas - Universidad de la Rioja - Gobierno de La Rioja, 26071 Logroño, La Rioja, Spain;
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Department of Plant Pathology, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa; 8Plant Protection
Division, ARC Infruitec-Nietvoorbij, Private Bag X5026, Stellenbosch 7599, South Africa; 9Microbe Interaction and Ecology
Laboratory, Biodiversity and Biotechnological Resource Research Unit (BBR), BIOTEC, NSTDA 113 Thailand Science Park
Phahonyothin Rd., Khlong Nueng, Khlong Luang, Pathum Thani 12120, Thailand; 10Faculty of Veterinary and Agricultural Sciences,
University of Melbourne 3010, Melbourne, Victoria, Australia; 11Naktuinbouw, Sotaweg 22, 2371 GD Roelofarendsveen, the
Netherlands; 12Hölderlinstraße 25, 15517 Fürstenwalde/Spree, Germany; 13Plant Health and Environment Laboratory, Ministry for
Primary Industries, P.O. Box 2095, Auckland 1140, New Zealand; 14ARC – Plant Protection Research Institute, Private Bag X5017,
Stellenbosch 7599, South Africa; 15Wageningen University and Research Centre (WUR), Laboratory of Phytopathology,
Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands.
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*Correspondence: Y. Marin-Felix, y.marin@westerdijkinstitute.nl; P.W. Crous, p.crous@westerdijkinstitute.nl
Running title: Genera of Phytopathogenic Fungi 2
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Abstract: This paper represents the second contribution in the Genera of Phytopathogenic Fungi (GOPHY) series. The series
provides morphological descriptions and information regarding the pathology, distribution, hosts and disease symptoms for the
treated genera. In addition, primary and secondary DNA barcodes for the currently accepted species are inlcuded. This second
paper in the GOPHY series treats 20 genera of phytopathogenic fungi and their relatives including: Allantophomopsiella,
Apoharknessia, Cylindrocladiella, Diaporthe, Dichotomophthora, Gaeumannomyces, Harknessia, Huntiella, Macgarvieomyces,
Metulocladosporiella, Microdochium, Oculimacula, Paraphoma, Phaeoacremonium, Phyllosticta, Proxypiricularia, Pyricularia,
Stenocarpella, Utrechtiana and Wojnowiciella. This study includes the new genus Pyriculariomyces, 20 new species, five new
combinations, and six typifications for older names.
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Key words: DNA barcodes, fungal systematics, 26 new taxa, six new typifications
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Taxonomic novelties: New genera: Pyriculariomyces Y. Marín, M.J. Wingf. & Crous; New species: Apoharknessia eucalypti
Crous & M.J. Wingf., Cylindrocladiella addiensis L. Lombard & Crous, Cylindrocladiella nauliensis L. Lombard & Crous, Diaporthe
heterophyllae Guarnaccia & Crous, Diaporthe racemosae A.R. Wood, Guarnaccia & Crous, Dichotomophthora basellae Hern.Restr., Cheew. & Crous, Dichotomophthora brunnea Hern.-Restr. & Crous,, Harknessia bourbonica Crous & M.J. Wingf.,
Harknessia corymbiae Crous & A.J. Carnegie, Harknessia cupressi Crous & R.K. Schumach., Harknessia pilularis Crous & A.J.
Carnegie, Huntiella abstrusa A.M. Wilson, Marinc., M.J. Wingf., Metulocladosporiella chiangmaiensis Y. Marín, Cheew. & Crous,
Metulocladosporiella malaysiana Y. Marín & Crous, Metulocladosporiella musigena Y. Marín, Cheew. & Crous, Metulocladosporiella
samutensis Y. Marín, Luangsa-ard & Crous, Microdochium novae-zelandiae Hern.-Restr., Thangavel & Crous, Phaeoacremonium
pravum C.F.J. Spies, L. Mostert & Halleen, Phyllosticta iridigena Y. Marín & Crous; Phyllosticta persooniae Y. Marín & Crous; New
combinations: Macgarvieomyces luzulae (Ondřej) Y. Marín, Akulov & Crous, Pyriculariomyces asari (Crous & M.J. Wingf.) Y. Marín,
M.J. Wingf. & Crous, Utrechtiana arundinacea (Corda) Crous, Quaedvl. & Y. Marín, Utrechtiana constantinescui (Melnik & Shabunin)
Crous & Y. Marín; New status and combination: Oculimacula acuformis (Nirenberg) Y. Marín & Crous; Typification:
Epitypification: Helminthosporium arundinaceum Corda, Phomopsis pseudotsugae M. Wilson, Pyricularia luzulae Ondřej,
Pyricularia zingiberis Y. Nishik.; Lectotypification: Phomopsis pseudotsugae M. Wilson, Pyricularia zingiberis Y. Nishik.
INTRODUCTION
The series, Genera of Phytopathogenic Fungi (GOPHY), was launched by Marin-Felix et al. (2017) to
provide a stable platform for the taxonomy of phytopathogenic fungi. The common denominator for the
genera included in this series is their association with plant diseases. The authors recognise that many
species treated are not well-known plant pathogens or where Koch’s postulates have not been proven for
them. The focus of the series is mainly to resolve generic and species concepts of the fungi studied. This
is particulary important because many taxa have been shown to represent species complexes, or to be
accommodated in genera that are poly- or paraphyletic (Crous et al. 2015b). The Series links to a larger
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MATERIAL AND METHODS
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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). Some of the main problems are that for many genera and species type material has not been
designated or/and that the vast majority of these taxa were described before the DNA phylogenetic era
(Hibbett et al. 2011) and thus lack DNA barcodes (Schoch et al. 2012). Another important aim of this
project is to secure the application of names by generating DNA barcodes of type species of genera and
type specimens of species. In those 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). The ultimate objective is to move to a single scientific name for
fungi (Crous et al. 2015b) for which sexual-asexual links have been resolved.
For each paper in the GOPHY series, morphological descriptions and information regarding the
pathology, distribution, hosts and disease symptoms are provided for the treated genera. In addition, this
information is linked to primary and secondary DNA barcodes of the current accepted species in each
genus. These DNA barcodes are critically important because of problems relating to generic delimitation
and species identification based solely on morphology. A clear example is the delimitation of the genera
Bipolaris and Curvularia, treated in the first paper of the GOPHY series (Marin-Felix et al. 2017). These
two genera share many morphological similarities, and intermediate conidial characters (Manamgoda et
al. 2012). Species delimitation in both genera based on morphology alone is of limited value because
many species have overlapping characters (Sivanesan 1987, Madrid et al. 2014, Manamgoda et al.
2014). Some genera include species that do not produce reproductive structures and their identification
must rely on DNA data. For some phytopathogenic genera, the DNA barcodes for species delimitation
have been established in previous studies, but for the vast majority, these data remain unavailable.
Mycologists wishing to contribute to future issues in the GOPHY series are encouraged to contact
Pedro Crous (p.crous@westerdijkinstitute.nl) before submitting their contributions. This will ensure there
is no overlap with activities arising from other research groups. Preference will be given to genera that
include novel DNA data and/or novel species, combinations or typifications. The generic contributions,
apart from being published in this series of papers, will also be placed in the database displayed on
www.plantpathogen.org.
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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
Westerdijk Fungal Biodiversity Institute in Utrecht, The Netherlands (CBS), the working collection of P.W.
Crous (CPC), housed at the Westerdijk Fungal Biodiversity Institute, and the culture collection (CMW) of
the Forestry and Agricultual Biotechnology Institute (FABI), at the University of Pretoria, South Africa. For
fresh collections, we followed the procedures previously described in Crous et al. (1991). Colonies were
transferred to different media, i.e. cherry decoction agar (CHA), carnation leaf agar (CLA), cornmeal agar
(CMA), 2 % malt extract agar (MEA), 2 % potato dextrose agar (PDA), synthetic nutrient-poor agar (SNA),
oatmeal agar (OA), water agar (WA) (Crous et al. 2009), autoclaved pieces of grapevine canes placed on
water agar (grapevine water agar; GWA), pine needle agar (PNA; Smith et al. 1996), or malt extract
peptone agar (MPA; Speakman 1984), and incubated at different conditions depending on the taxon to
induce sporulation. Requirements of media and conditions of incubations are specified in each genus.
Reference strains and specimens are maintained at the CBS, CMW and CPC.
Vegetative and reproductive structures were mounted in 100% lactic acid either directly from
specimens or from colonies sporulating on MEA, PDA, PNA, OA or SNA. For cultural characterisation,
isolates were grown and incubated on different culture media and temperatures as stipulated for each
genus. Colour notations were rated according to the colour charts of Rayner (1970). Taxonomic novelties
were deposited in MycoBank (www.MycoBank.org; Crous et al. 2004a).
DNA isolation, amplification and analyses
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Allantophomopsiella Crous, IMA Fungus 5: 180. 2014. Fig. 1.
Classification: Leotiomycetes, Leotiomycetidae, Phacidiales, Phacidiaceae.
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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 bibliography related to the phylogeny presented for each
respective genus. Phylogenetic analyses consisted of Maximum-Likelihood (ML), Bayesian Inference (BI),
and Maximum Parsimony (MP). The ML and the BI were carried out using methods described by
Hernández-Restrepo et al. (2016b), and the MP using those described by Crous et al. (2006b). Sequence
data generated in this study were deposited in GenBank and the alignments and trees in TreeBASE
(http://www.treebase.org).
DNA barcodes (genus): ITS, LSU.
DNA barcodes (species): ITS, rpb2. Table 1.
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Type species: Allantophomopsiella pseudotsugae (M. Wilson) Crous., basionym: Phomopsis
pseudotsugae M. Wilson. Lectotype designated here: material deposited in Royal Botanic Garden
Edinburgh, E00414771. Epitype and ex-epitype strain designated here: CBS H-23354, CBS 320.53.
Conidiomata up to 600 m diam, pycnidial, immersed, becoming erumpent, irregularly multilocular, dark
brown, ostiolate; conidiomatal wall composed of 3–4 layers of dark brown cells, textura angularis.
Conidiophores arising from inner layer of conidioma, branched, septate, at times reduced to conidiogenous
cells. Conidiogenous cells integrated or discrete, ampulliform to subcylindrical or lageniform, hyaline,
smooth with minute periclinal thickening at apex. Conidia inequilaterally fusiform or naviculate, hyaline,
smooth, aseptate, guttulate, bearing mucoid apical appendages, flabelliform to irregular in shape. Sexual
morph unknown (adapted from Crous et al. 2014b).
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Culture characteristics: Colonies spreading, flat with sparse aerial mycelium and feathery margins. On PDA
surface olivaceous grey, reverse iron-grey. On OA surface olivaceous grey with patches of iron-grey.
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Optimal media and cultivation conditions: PNA at 25 °C under continuous near-ultraviolet li ght to promote
sporulation.
Distribution: North America and Europe.
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Hosts: Conifers (Pinaceae).
Disease symptoms: Canker and dieback.
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Notes: This genus was recently introduced by Crous et al. (2014b) to accommodate A. pseudotsugae, a
pathogen of conifers that was found to be very damaging, especially after wounding during tree dormancy
(Roll Hansen 1992). In a study considering the pathogenicity of this fungus on Pinus sylvestris associated
with pruning wounds, it was observed that Allantophomopsis pseudotsugae occurred commonly in slash
of pine trees wounded during the autumn (Uotila 1990).
Allantophomopsiella is morphologically related to the phytopathogenic genera Apostrasseria and
Allantophomopsis. However, it can be easily differentiated from both genera by the lack of percurrent
proliferation on its conidiogenous cells, and by the production of inequilaterally fusiform or naviculate
conidia.
References: Uotila 1990 (pathogenicity); Crous et al. 2014b (morphology and phylogeny).
Allantophomopsiella pseudotsugae (M. Wilson) Crous, IMA Fungus 5: 180. 2014. Fig. 1.
Basionym: Phomopsis pseudotsugae M. Wilson, Transactions of the Royal Scottish Arboricultural Society
34: 147. 1920.
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Synonyms: Phacidiella coniferarum G.G. Hahn, Mycologia 49: 227. 1957.
Phacidium coniferarum (G.G. Hahn) DiCosmo, et al., Canad. J. Bot. 61: 37. 1983.
Allantophomopsis pseudotsugae (M. Wilson) Nag Raj, Coelomycetous anamorphs with appendagebearing conidia: 116. 1993.
Additional synonyms are provided in Nag Raj (1993).
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Materials examined: UK, Scotland, Murthly, on Pseudotsuga menziesii (Pinaceae), Apr. 1920, M. Wilson (lectotype of Phomopsis
pseudotsugae designated here: MBT379803, material deposited in Royal Botanic Garden Edinburgh, E00414771). Norway,
Førde in Sunnfjord, shoot of Pseudotsuga menziesii (Pinaceae), Apr. 1948, H. Robak (epitype of Phomopsis pseudotsugae
designated here CBS H-23354, MBT379804, culture ex-epitype CBS 320.53).
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Notes: Allantophomopsis pseudotsugae was introduced by Wilson (1920) as Phomopsis pseudotsugae
to accommodate a fungus that infects Pseudotsuga menziesii in Scotland. Type material was not
specified when it was described, but the author deposited original material used for the Royal Botanic
Garden Edinburgh. This material with the barcode number E00414771, which appears to be syntype
material of the species, is selected here as lectotype. To fix the use of the name, the strain CBS 320.53
is designated here as ex-epitype. This strain was collected in Norway, occurring on the type host (Wilson
1920), and fits well with the description of the species recently provided by Crous et al. (2014b).
Authors: Y. Marin-Felix & P.W. Crous
Apoharknessia Crous & S.J. Lee, Stud. Mycol. 50: 239. 2004. Fig. 2.
Classification: Sordariomycetes, Diaporthomycetidae, Diaporthales, Apoharknessiaceae.
Type species: Apoharknessia insueta (B. Sutton) Crous & S.J. Lee, basionym: Harknessia insueta B.
Sutton. Holotype of H. insueta: IMI 22697. Epitype and ex-epitype strain designated by Lee et al. (2004):
CBS H-9913, CBS 111377 = STE-U 1451.
DNA barcodes (genus): ITS, LSU.
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DNA barcodes (species): ITS, cal, tub2. Table 2.
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Conidiomata pycnidial, separate to gregarious, subepidermal, becoming erumpent, stromatic,
amphigenous, depressed globose or subglobose to irregular, unilocular, glabrous; opening irregularly,
with yellowish, furfuraceous cells, lacking a definite ostiole; conidiomatal wall of textura angularis.
Conidiophores reduced to conidiogenous cells lining cavity. Conidiogenous cells lageniform to ampulliform
or ampulliform to subcylindrical, hyaline, smooth, in mucilage, percurrently proliferating once or twice near
apex. Conidia broadly ellipsoidal to obovoid or obliquely gibbose, apex obtusely rounded, aseptate, nonapiculate, medium brown or brown, with or without a longitudinal hyaline band on flat surface, thickwalled, smooth, with or without striations along length of conidia, with prominent central guttule; basal
appendage absent or hyaline, tubular, smooth, thin-walled, devoid of cytoplasm; apical appendage
absent or hyaline, conical or tubular, short, smooth, thin-walled, devoid of cytoplasm. Sexual morph
unknown.
Culture characteristics: Colonies flat on MEA, PDA and OA, with sparse or moderate aerial mycelium,
smooth, lobate margins, fast sporulating. On MEA surface greenish black or olivaceous black in centre,
dirty white in outer region; reverse olivaceous grey in centre, dirty white in outer region.
Optimal media and cultivation conditions: MEA, PDA and OA under continuous near-ultraviolet light at 25 °C
to promote sporulation.
Distribution: Brazil, Colombia, Cuba, Malaysia and Mauritius.
Hosts: Eucalyptus (Myrtaceae).
Disease symptoms: Leaf spots.
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Notes: The genus Apoharknessia was established by Lee et al. (2004) for Ap. insueta, a presumed foliar
pathogen of eucalypts in Brazil, Colombia, Cuba and Mauritius. The genus included two species following
the description of Ap. eucalyptorum to accommodate a presumed endophyte of Eucalyptus pellita isolated
from collections made in Malaysia (Crous et al. 2017a).
Apoharknessia is distinguished from Harknessia by having an apical appendage on its conidia, and
cultures that lack fluffy aerial mycelium and sporulate abundantly on the aerial hyphae. Cultures of
Harknessia s. str. are slow to sporulate, and tend to form abundant pycnidial conidiomata in culture (Lee
et al. 2004).
Recently, the family Apoharknessiaceae was introduced based on a phylogenetic study of the
members of the Diaporthales, in order to accommodate Apoharknessia, designated as the type genus of
the family, together with Lasmenia, which is also a genus associated with plant disease (Senanayake et
al. 2017).
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References: Lee et al. 2004 (morphology and phylogeny); Crous et al. 2012c, 2017a, Senanayake et
al. 2017 (phylogeny).
Apoharknessia eucalypti Crous & M.J. Wingf., sp. nov. MycoBank MB820945. Fig. 3.
Etymology: Name refers to Eucalyptus, the host genus from which this fungus was collected.
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Foliicolous, isolated from leaves incubated in moist chambers (presumed endophytic). Conidiomata up to
250 µm diam, pycnidial, solitary to gregarious, subepidermal, becoming erumpent, stromatic,
amphigenous, depressed globose; opening irregularly, with yellowish, furfuraceous cells; conidiomatal
wall of textura angularis. Conidiophores reduced to conidiogenous cells lining cavity. Conidiogenous cells
8–12 × 3–5 µm, ampulliform to subcylindrical, hyaline, smooth, in mucilage, percurrently proliferating once
or twice near apex. Conidia (7–)8–10(–11) × (5–)6(–7) µm, broadly ellipsoid to obovoid, apex obtusely
rounded, aseptate, non-apiculate, medium brown, thick-walled, smooth, striations along length of
conidium body, with prominent central guttule. Basal appendage absent or 0–2 µm long, 2 µm diam,
hyaline, tubular, smooth, thin-walled, devoid of cytoplasm.
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Culture characteristics: Colonies on MEA, PDA and OA, with moderate aerial mycelium and smooth,
lobate margins, reaching 60 mm diam after 2 wk at 25 °C. On MEA surface greenish black in middle, dirt y
white in outer region, reverse olivaceous grey in centre, dirty white in outer region. On PDA surface and
reverse olivaceous grey in centre, dirty white in outer region. On OA surface greenish black in middle,
outer region dirty white.
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Material examined: Malaysia, Sabah, isolated from leaves of Eucalyptus pellita (Myrtaceae), May 2015, M.J. Wingfield (holotype
CBS H-23081, culture ex-type CPC 27550 = CBS 142518).
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Notes: Apoharknessia eucalypti is an odd member of the genus in that its conidia generally lack an apical
appendage. The latter was observed on one or two conidia, irrespective of the media used for cultivation.
The ITS sequence of Ap. eucalypti is only 92 % similar to that of Ap. insueta [GenBank JQ706083;
Identities = 569/618 (92 %), 33 gaps (5 %)] and 97 % similar to that of Ap. eucalyptorum [GenBank
KY979752; Identities = 604/621 (97 %), 5 gaps (0%)].
Authors: P.W. Crous, Y. Marin-Felix, J.Z. Groenewald & M.J. Wingfield
Cylindrocladiella Boesew., Canad. J. Bot. 60: 2289. 1982. Fig. 4.
Synonym: Nectricladiella Crous & C.L. Schoch, Stud. Mycol. 45: 54. 2000.
Classification: Sordariomycetes, Hypocreomycetidae, Hypocreales, Nectriaceae.
Type species: Cylindrocladiella parva (P.J. Anderson) Boesew., basionym: Cylindrocladium parvum P.J.
Anderson. Representative strain: CBS 114524 = ATCC 28272.
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DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): his3, tef1, tub2. Table 3. Fig. 5.
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Ascomata perithecial, superficial, solitary, basal stroma absent, globose to obpyriform, collapsing laterally
when dry, smooth, with minute, brown setae arising from ascomatal wall surface, red, changing colour in
KOH, ostiole consisting of clavate cells, lined with inconspicuous periphyses. Asci unitunicate, 8-spored,
cylindrical, sessile, thin-walled, with a flattened apex, and a refractive apical apparatus. Ascospores
uniseriate, overlapping, hyaline, ellipsoid to fusoid with obtuse ends, smooth, 1-septate. Conidiophores
monomorphic, penicillate, or dimorphic (penicillate and subverticillate), mononematous, hyaline.
Penicillate conidiophores consist of a stipe, a penicillate arrangement of fertile branches, a stipe
extension, and a terminal vesicle. Subverticillate conidiophores consist of a stipe, and one or two series of
phialides. Stipe septate, hyaline, smooth. Stipe extensions aseptate, straight, thick-walled, with one basal
septum, terminating in a thin-walled vesicle of characteristic shape. Conidiogenous apparatus with
primary branches 0–1-septate, secondary branches aseptate, terminating in 2–4 phialides. Phialides
cylindrical, straight or doliiform to reniform to cymbiform, hyaline, aseptate, apex with minute periclinal
thickening and collarette. Conidia cylindrical, rounded at both ends, straight, (0–)1(–3)-septate, frequently
slightly flattened at base, held in asymmetrical clusters by hyaline slime. Chlamydospores brown, thickwalled, more frequently arranged in chains than clusters (adapted from Boesewinkel 1982 and Lombard
et al. 2012).
Culture characteristics: Colonies on MEA white to pale brick when young, becoming pale brick to dark
sepia when mature, fluffy, cottony, effuse to convex with papillate surface, margin entire, undulate, lobate,
or fimbriate, sometimes with abundant chlamydospores forming microsclerotia within medium.
Optimal media and cultivation conditions: CLA to induce sporulation of the asexual morph at 25 °C, while
for the sexual morph sterile toothpicks placed on SNA is used at 20 °C.
Distribution: Worldwide.
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Hosts: Soil-borne, weak pathogen of forestry, agricultural and horticultural crops (Crous 2002, Lombard et
al. 2012).
Disease symptoms: Leaf spots, cutting rot, stem cankers, damping-off and root rot.
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Notes: Boesewinkel (1982) established the asexual genus Cylindrocladiella, based on C. parva, to
accommodate several cylindrocladium-like species characterised by small (< 20 µm long), 1-septate,
cylindrical conidia and aseptate stipe extensions. Initially, Cylindrocladiella spp. were linked to the sexual
genus Nectricladiella (Crous & Wingfield 1993, Schoch et al. 2000). Following the implementation of the
International Code of Nomenclature for algae, fungi and plants (ICN; McNeill et al. 2012), Rossman et al.
(2013) proposed that the generic name Cylindrocladiella be retained over Nectricladiella. Recently,
Lombard et al. (2015) showed that the genus Cylindrocladiella formed a monophyletic group in the
Nectriaceae, closely related to the genera Aquanectria and Gliocladiopsis. To date, 36 species of
Cylindrocladiella have been recognised (Crous & Wingfield 1993, van Coller et al. 2005, Inderbitzin et al.
2012, Lombard et al. 2012, 2015, 2017; Crous et al. 2017a), of which two are linked to nectricladiella-like
sexual morphs (Schoch et al. 2000, Crous 2002, Lombard et al. 2012, 2015). These fungi are generally
soil-borne and regarded as saprobes or weak pathogens of numerous plant hosts (Crous 2002, van
Coller et al. 2005, Scattolin & Montecchio 2007, Lombard et al. 2012). Disease symptoms associated with
Cylindrocladiella infection include leaf spots and root, stem and cutting rots (Crous et al. 1991, Peerally
1991, Crous & Wingfield 1993, Crous 2002, van Coller et al. 2005, Scattolin & Montecchio 2007, Lombard
et al. 2012).
References: Boesewinkel 1982 (morphology and pathogenicity); Crous & Wingfield 1993 (morphology);
Schoch et al. 2000, Lombard et al. 2012, 2017 (morphology and phylogeny); Crous 2002 (morphology,
pathogenicity and monograph); van Coller et al. 2005 (morphology, pathogenicity and phylogeny).
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Cylindrocladiella addiensis L. Lombard & Crous, sp. nov. MycoBank MB824497.
Etymology: Name refers to Addis Abeba, Ethiopia, from where this fungus was collected.
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Cultures sterile. Cylindrocladiella addiensis differs from its closest phylogenetic neighbours, C. elegans
and C. noveazelandica, by unique fixed alleles in three loci based on the alignments deposited in
TreeBASE (S22340): ITS position 92(T); tef1 positions 32(T), 80(C), 84(C), 154(indel), 155(A), 156(G),
157(indel), 199(A), 244(G), 261(G), 368(T), 398(T), 458(G/C), 466(G), 467(C), 475(T), 478(C), 483(T),
485(T), 487(T), 490(T) and 492(G); tub2 position 174(indel).
Culture characteristics: Colonies convex, cottony, with smooth margins, cream; reverse pale luteous;
chlamydospores not seen; reaching 45–65 mm after 1 wk on MEA at 24 °C in ambient light.
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Materials examined: Ethiopia, Addis Ababa, from soil, 2010, coll. P.W. Crous, isol. L. Lombard [holotype CBS 143794 (maintained
as metabolically inactive culture), isotype cultures CBS 143793, CBS 143795 (also maintained as metabolically inactive cultures).
Note: None of the three isolates of C. addiensis could be induced to sporulate on MEA, PDA, OA, SNA or
SNA amended with carnation leaf pieces.
Cylindrocladiella nauliensis L. Lombard & Crous, sp. nov. MycoBank MB824500. Fig. 6.
Etymology: Name refers to the area Aek Nauli, Indonesia, from where this fungus was collected.
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Conidiophores 35–55 × 3–6 m, dimorphic, penicillate and subverticillate, mononematous and hyaline,
comprising a stipe, a penicillate arrangement of fertile branches, a stipe extension and a terminal vesicle;
stipe septate, hyaline, smooth; stipe extension 100–135 m long, 4–5 m wide, aseptate, straight, thickwalled with one basal septum, terminating in thin-walled, broadly clavate to ellipsoidal vesicles sometimes
with papillate apex. Penicillate conidiogenous apparatus 10–22 × 2–4 m, with primary branches
aseptate, secondary branches 8–15 × 2–4 m, aseptate, each terminal branch producing 2–4 phialides;
phialides 8–13 × 2–3 m, elongate doliiform to reniform to cymbiform, hyaline, aseptate, apex with minute
periclinal thickening and collarette. Subverticillate conidiophores abundant, comprised of a septate stipe
and rarely primary branches terminating in 2–4 phialides; primary branches 25–45 × 2–4 m, straight,
hyaline, 0–1-septate; phialides 12–32 × 2–3 m, cymbiform to cylindrical, hyaline, aseptate, apex with
minute periclinal thickening and collarette. Conidia (10–)11–13(–14) × 2–3 m (av. = 12 × 2 m),
cylindrical, rounded at both ends, straight, 1-septate, frequently slightly flattened at base, held in
asymmetrical clusters by hyaline slime. Sexual morph unknown.
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Culture characteristics: Colonies convex, cottony, with smooth margins, cream with pale luteous to brick
centre; reverse pale luteous to honey with sepia centre; chlamydospores moderate throughout medium
arranged in chains; reaching 60–70 mm after 1 wk on MEA at 24 °C in ambient light.
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Materials examined: Indonesia, Sumatra Utara, Aek Nauli, from soil, May 2005, coll. M.J. Wingfield, isol. L. Lombard (holotype
CBS H-23400, culture ex-type CBS 143792); ibid., isotype culture CBS 143791 (metabolically inactive).
Notes: Cylindrocladiella nauliensis is closely related to C. longistipitata. The stipe extensions of C.
nauliensis (up to 135 m long) are shorter than those of C. longistipitata (up to 216 m long; Lombard et
al. 2012). Additionally, the conidia of C. nauliensis are smaller [(10–)11–13(–14) × 2–3 m (av. = 12 × 2
m)] than those of C. longistipitata [(12–)14–16(–17) × 2–4 m (av. = 15 × 3 m); Lombard et al. 2012].
Authors: L. Lombard & P.W. Crous
Diaporthe Nitschke, Pyrenomyc. Germ. 2: 240. 1870. Figs 7, 8.
Synonym: Phomopsis, Sacc., Syll. fung. (Abellini) 2: 484. 1883.
Classification: Sordariomycetes, Sordariomycetidae, Diaporthales, Diaporthaceae.
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Type species: Diaporthe eres Nitschke. Lectotype designated by Udayanga et al. (2014a): B 70 0009145.
Epitype and ex-epitype strain designated by Udayanga et al. (2014a): BPI 892912, AR5193 = CBS
138594.
DNA barcodes (genus): ITS.
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DNA barcodes (species): cal, his3, tef1, tub2. Table 4. Fig. 9.
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Ascomata immersed in substrate, subglobose or irregular, solitary or clustered in groups, often erumpent
through a pseudostroma mostly surrounding ascomata with more or less elongated ascomatal necks.
Pseudostroma distinct, often delimited with dark lines. Asci unitunicate, 8-spored, sessile, elongate to
clavate or cylindrical, loosening from ascogenous cells at an early stage and floating free in ascomata.
Ascospores biseriate to uniseriate in ascus, fusoid, ellipsoid to cylindrical, straight, inequilateral or curved,
septate, hyaline, sometimes with appendages. Conidiomata pycnidial, deeply embedded in culture on
several media, globose to conical, eustromatic, multilocular, occasionally with ostiolate necks, scattered
or aggregated, brown to black, surface covered with hyphae, cream to pale luteous or yellowish, conidial
droplets or cirrus exuding from central ostioles; conidiomatal wall consisting of pale brown, thick walled
textura angularis. Conidiophores cylindrical to clavate, straight to sinuous, densely aggregated, branched,
0–6-septate, smooth, hyaline in upper region, pale brown at base. Conidiogenous cells phialidic, hyaline,
cylindrical, terminal and lateral, tapering slightly towards apex. Paraphyses occasionally produced,
intermingled among conidiophores, hyaline, smooth, 1–3-septate. Alpha conidia aseptate, generally
hyaline, smooth, fusiform to ellipsoidal, with obtuse or acute to rounded ends, non- to multi-guttulate, but
often bi-guttulate. Beta conidia aseptate, hyaline, filiform, smooth, straight or more often hooked,
eguttulate, tapering or truncated towards ends. Gamma conidia rarely produced, hyaline, smooth, non- to
multi-guttulate, fusiform to subcylindrical with acute or rounded apex (adapted from Gomes et al. 2013,
Udayanga et al. 2014a).
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Culture characteristics: Colonies on MEA, PDA and OA producing abundant compact, flattened, aerial
mycelium, sometimes in rings, with an entire to irregular margin, white, cream to yellowish or pale
olivaceous grey, smoke grey to grey, cottony. Reverse pale brown to grey, dark green, producing
brownish dots with age, with solitary or aggregated conidiomata at maturity.
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Optimal media and cultivation conditions: On MEA, PDA and OA at 25 °C, or sterile pine need les placed
on SNA at 25 °C under near-ultraviolet light (12 h light, 12 h dark) to induce sporulation of the asexual
morph.
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Distribution: Worldwide.
Hosts: On a wide range of plant families.
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Disease symptoms: Root and fruit rots, dieback, stem cankers, leaf spots, leaf and pod blights, and seed
decay.
Notes: The genus Diaporthe presently includes 213 species supported by ex-type cultures and
supplementary DNA barcodes, which include endophytes, saprobes and important plant pathogenic
species. Recent phylogenetic analyses of the genus Diaporthe grouped some of those species into
complexes, such as D. arecae, D. eres and D. sojae (Huang et al. 2013, Udayanga et al. 2014a, 2015).
Several pathology studies confirmed Diaporthe species to be associated with diverse suites of diseases
(Fig. 7) on a broad range of economically important agricultural crops (Udayanga et al. 2011). More than
one Diaporthe species is frequently reported as causative agents of the same disease (Thomson et al.
2011, Guarnaccia et al. 2016).
Although Diaporthe was historically considered monophyletic based on the typical phomopsis-like
asexual morph, the paraphyletic nature of this genus was recently revealed (Gao et al. 2017, Senanayake
et al. 2017). Most of the known species in early literature were described in relation to their host
association and morphological characters. However, a single species of Diaporthe can be found on
diverse hosts, and can co-occur on the same host or lesion in different life modes. Phylogenetic studies
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demonstrated that morphological characters are not always reliable for species level identification due to
their variability under changing environmental conditions (Gomes et al. 2013). As a consequence,
identification and description of species based on host association alone is no longer tenable. For
accurate species delimitation, phylogenetic inference of the ITS, cal, his3, tef1 and tub2 or combinations
of these is required.
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References: Mostert et al. 2001, Van Niekerk et al. 2005, Thomson et al. 2011, Guarnaccia et al. 2016,
2018 (morphology, pathogenicity and phylogeny); Udayanga et al. 2011, 2014a, 2015, Gomes et al. 2013
(morphology and phylogeny); Dissanayake et al. 2017b, c, Gao et al. 2017 (phylogeny).
Diaporthe heterophyllae Guarnaccia & Crous, sp. nov. MycoBank MB823830. Fig. 10.
Etymology: Name refers to Acacia heterophylla, the host from which this fungus was collected.
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On PNA: Conidiomata 250–350 m diam, pycnidial, globose or irregular, solitary, deeply embedded in
media, erumpent, dark brown to black, whitish translucent to yellow conidial drops and/or cirrus exuded
from ostioles; conidiomatal wall consisting of 3–4 layers of medium brown textura angularis.
Conidiophores 7–22 × 1.5–4 m, hyaline, smooth, 0–1-septate, densely aggregated, cylindrical, straight.
Conidiogenous cells 6–9 × 1–2 m, phialidic, hyaline, terminal, cylindrical, tapered towards apex.
Paraphyses not observed. Alpha conidia 6–10.5 × 2.5–4.5 m, mean ± SD = 8.4 ± 1.1 × 3.2 ± 0.4 m,
L/W ratio = 2.6, aseptate, ovate to ellipsoidal, hyaline, multi-guttulate and acute or rounded at both ends.
Beta conidia 17–24 × 1–2 m, mean ± SD = 21.7 ± 1.8 × 1.5 ± 0.3 m, L/W ratio = 14.5, hyaline,
aseptate, eguttulate, filiform, curved, tapering towards both ends. Gamma conidia not observed.
Culture characteristics: Colonies covering medium within 10 d at 21 °C, wi th surface mycelium flattened,
dense and felty. Colony on MEA, PDA and OA at first white, becoming cream to yellowish, flat on MEA
and OA, dense, felted on PDA. Reverse grey with brownish dots with age, with visible solitary
conidiomata at maturity on all media.
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Material examined: France, La Rèunion, on Acacia heterophylla (Fabaceae), 8 Mar. 2015, P.W. Crous (holotype CBS H-23376,
culture ex-type CBS 143769 = CPC 26215).
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Notes: Diaporthe heterophyllae is phylogenetically close but clearly differentiated from D. eres based on
ITS, tef1, tub2, his3 and cal sequence similarity (98 %, 88 %, 97 %, 95 %, and 97 %, respectively).
Morphologically, D. heterophyllae differs from D. eres in its longer alpha conidia (6.5–10.5 vs. 6–8.5 m)
and in its shorter beta conidia (17–24 vs. 22–28 m) (Udayanga et al. 2014a).
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Diaporthe racemosae A.R. Wood, Guarnaccia & Crous, sp. nov. MycoBank MB823831. Fig. 11.
Etymology: Name refers to Euclea racemosa, the host from which this fungus was collected.
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On PNA: Conidiomata 350–600 m diam, pycnidial, globose or irregular, solitary, deeply embedded in
media, erumpent, dark brown to black, yellowish translucent to pale brown conidial drops and/or cirrus
exuded from ostioles; conidiomatal wall consisting of 3–4 layers of pale brown textura angularis.
Conidiophores 7–17 × 2–4 m, hyaline, smooth, 0–1-septate, densely aggregated, cylindrical, straight.
Conidiogenous cells 5.5–8 × 1–2 m, phialidic, hyaline, terminal, subcylindrical, tapered towards apex.
Paraphyses not observed. Alpha conidia 4–6.5 × 2–3 m, mean ± SD = 5.7 ± 0.6 × 2.3 ± 0.3 m, L/W
ratio = 2.5, aseptate, ellipsoidal to subcylindrical, hyaline, non- to multi-guttulate and acute or rounded at
both ends. Beta and gamma conidia not observed.
Culture characteristics: Colonies covering medium within 10 d at 21 °C, wi th surface mycelium flattened,
dense and felty. Colony on MEA and OA at first white, becoming olivaceous to dark grey. On PDA at first
white, becoming white to yellowish. Reverse grey with brownish dots with age, with visible solitary
conidiomata at maturity on all media.
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Material examined: South Africa, Western Cape, Bot River, from Euclea racemosa (Ebenaceae), 29 Dec. 2014, A.R. Wood
(holotype CBS H-23377, culture ex-type CBS 143770 = CPC 26646).
Authors: V. Guarnaccia, A.R. Wood & P.W. Crous
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Notes: Diaporthe racemosae is phylogenetically close but clearly differentiated from D. schini based on
ITS, tef1, tub2, his3 and cal sequence similarity (98 %, 94 %, 98 %, 94 %, and 96 %, respectively).
Moreover, D. racemosa produces only alpha conidia, while D. schini produces only beta conidia (Gomes
et al. 2013).
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Dichotomophthora Mehrl. & Fitzp. ex M.B. Ellis, Dematiaceous Hyphomycetes (Kew): 388. 1971. Fig
12.
Synonyms: Dichotomophthora Mehrl. & Fitzp., Mycologia 27: 550. 1935. (nom. inval., Art. 39.1,
Melbourne).
Dichotomophthora Mehrl. & Fitzp. ex P.N. Rao, Mycopath. Mycol. appl. 28: 139. 1966. (nom. inval., Art.
39.1, Melbourne).
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae.
Type species: Dichotomophthora portulacae Mehrl. & Fitzp. ex M.B. Ellis. Type specimen and exparatype strain: IMI 8742, CBS 174.35.
DNA barcodes (genus): ITS.
DNA barcodes (species): ITS, rpb2, gpdh. Table 5. Fig. 13.
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Conidiophores macronematous, mononematous, unbranched or irregularly branched, sometimes
swollen and repeatedly dichotomously or trichotomously branched or lobed at apex, forming a stipe and
head; stipe hyaline to brown; branches usually short. Conidiogenous cells mono- or polytretic, integrated,
terminal, lobed, cicatrized. Conidia solitary, dry, simple, ellipsoidal to cylindrical, rounded at ends,
subhyaline to brown, multi-distoseptate. Microconidia ovoid, 0–2-distoseptate. Sclerotia often formed in
culture resembling immature perithecia, semi- or immersed in agar, subglobose, ellipsoidal, ovoid, dark
brown or black. Sexual morph unknown (adapted from Ellis 1971).
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Culture characteristics: Colonies on PDA and OA white, hazel, orange, or dark grey to olivaceous,
cottony, velvety, somewhat fluffy, or flat, margin irregular, effuse; reverse centre hazel, dark brown,
periphery hazel, orange to luteous. Diffusible pigment luteous to orange (produced in some strains).
Optimal media and cultivation conditions: On PDA and OA at 25 °C under near-ultraviolet lig ht (12 h
light, 12 h dark). Some strains are sterile in culture.
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Distribution: Worldwide.
Hosts: Anredera and Basella (Basellaceae), Beta vulgaris (Chenopodiaceae), Gymnocalycium
mihanovichii var. friedrichii and Myrtillocactus geometrizans (Cactaceae), Portulaca (Portulacaceae), and
soil.
Disease symptoms: Leaf spots, foliar abscission, stem blight, seed rot and damping-off.
Notes: Dichotomophthora was introduced as a monotypic genus with Di. portulacae isolated from
Portulaca oleracea in Hawaii (Mehrlich & Fitzpatrick 1935). However, the publication lacked a Latin
diagnosis of the fungus and the name was therefore invalid. Later, Rao (1966) provided a Latin
description of Di. portulacae and introduced a new species, Di. indica, but the validation was
misapplied and both names were regarded as invalid (de Hoog & Oorschot 1983). Ellis (1971)
validated the genus and the species based on the holotype specimen of Di. portulacae (IMI 8742). De
Hoog & van Oorschot (1983) revised the taxonomy of Dichotomophthora and included Di. portulacae
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and Di. lutea. Dichotomophthora portulacae was restricted to species with dichotomously branched
conidiophores, and conidia with 2–3 septa, 45–75 × 20–30 µm. Dichotomophthora lutea was
introduced based on Dactylaria lutea, which is characterised by unbranched or irregularly branched
conidiophores, and conidia with 1–5 septa, 30–115 × 10–20 µm.
Dichotomophthora species are mainly known as plant pathogens with a wide host range as well as
soil-borne fungi or saprobes (Mehrlich & Fitzpatrick 1935, Routien 1957, Rao 1966, Ellis 1971, Klisiewicz
1985, Baudoin 1986, Pfeiffer et al. 1989, Eken 2003, Farr & Rossman 2017, Soares & Nechet 2017).
However, a case of human keratitis caused by Di. portulacae was reported from subtropical Africa (de
Hoog et al. 2000). Since many records of Di. portulacae may represent Di. lutea due to previous
taxonomic confusion, host and distribution data need to be re-evaluated (de Hoog & van Oorschot 1983,
Farr & Rossman 2017, Soares & Nechet 2017).
This is the first time that numerous isolates, including the ex-type strains of both species of
Dichotomophthora, have been subjected to phylogenetic analyses. Our results suggest that
Dichotomophthora belongs in the Pleosporaceae (Pleosporales), closely related to Curvularia. The
phylogenetic analysis and subtle morphological evidence revealed two additional new species, introduced
here as Di. basellae and Di. brunnea. For an accurate identification at the species level, a DNA sequence
analysis is recommended, since Dichotomophthora species are morphologically variable in culture and on
natural substrates.
References: Mehrlich & Fitzpatrick 1935, Routien 1957, Rao 1966, Ellis 1971, de Hoog & van
Oorschot 1983 (taxonomy and morphology); Baudoin 1986, Klisiewicz 1985, Pfeiffer et al. 1989, Eken
2003, Soares & Nechet 2017 (pathogenicity).
Dichotomophthora basellae Hern.-Restr., Cheew. & Crous, sp. nov. MycoBank MB824604. Fig. 14.
Etymology: Name reflects the substrate from which this fungus was isolated, Basella alba.
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Hyphae 3–7.5 µm wide, hyaline to brown, septate, smooth to verruculose. Conidiophores
macronematous, mononematous, unbranched or irregularly branched, sometimes swollen and
repeatedly dichotomously or trichotomously branched or lobed at apex, forming a stipe and head; stipe
970–1370 × 10–12(–14) µm, pale brown, smooth; branches usually short; head 23–65 µm wide, pale
brown to brown. Conidiogenous cells polytretic, integrated and terminal, lobed, cicatrized, individual
lobes 6–14 × 6–9.5 µm. Conidia 32–86 × 10–18 µm, solitary, dry, ellipsoidal to cylindrical rounded at
ends, subhyaline to yellow brown, 2–5-distoseptate. Microconidia 11–30 × 9–13(–15) µm, obovoid to
ellipsoidal, 0–2-distoseptate. Sclerotia 295–444 × 234–409 µm, resembling immature perithecia, semi- or
immersed in agar, globose, subglobose, ellipsoidal, ovoid, dark brown or black. Sexual morph unknown.
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Culture characteristics: Colonies at 25 °C under near-ultraviolet light (1 2 h light, 12 h dark), on PDA and
OA reaching 45–50 mm after 1 wk, centre black, periphery luteous, velvety, flat, margin regular, effuse;
reverse centre olivaceous, dark brown, periphery pale luteous. Diffusible pigment luteous.
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Material examined: Thailand, Chiang Mai, Chiang Mai university experimental farm, on leaves of Basella alba (Basellaceae), 2010,
R. Cheewangkoon (holotype CBS H-23383, culture ex-type CPC 33016).
Notes: Dichotomophthora basellae is represented by one strain isolated from leaf spots on Basella rubra
in Thailand. This species is morphologically similar to Di. lutea in having multi-lobed conidiogenous cells
producing pale brown conidia, and colonies that produce a luteous to orange diffusible pigment in
culture. In our study, Di. basellae produces larger conidia than Di. lutea (32–86 × 10–18 µm vs. 14–65.5
× 7.5–13 µm). However, de Hoog & van Oorschot (1983) described larger conidia in Di. lutea (30–115
× 10–20 µm). Additional studies with more isolates are thus required to compare these morphological
differences and substrate preferences of both species.
Dichotomophthora brunnea Hern.-Restr. & Crous, sp. nov. MycoBank MB824605. Fig. 15.
Etymology: From the Latin brunnea meaning brown, because of the brown colour of the conidia.
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Hyphae 2.5–7 µm wide, hyaline to dark brown, septate, slightly constricted at septa, smooth.
Conidiophores macronematous, mononematous, repeatedly dichotomously or irregularly branched,
lobed at apex, forming a stipe and head; stipe 42–536 × 4.5–7.5 µm, pale brown to brown, smooth;
branches usually short; head 10–28 µm wide, brown to pale brown. Conidiogenous cells mono- or
polytretic, integrated, terminal, lobed, cicatrized, individual lobes 6.5–17 × 4–9 µm. Conidia 29–56.5 × 6–
10 µm, solitary, dry, ellipsoidal to cylindrical rounded at ends, brown to dark brown, 2–6(–8)-distoseptate,
straight or slightly curved. Microconidia 13–19.5 × 7–9.5 µm, obovoid to ellipsoidal, 0–1-distoseptate.
Sclerotia not observed. Sexual morph unknown.
Culture characteristics: Colonies on PDA and OA at 25 °C under near-ultrav iolet light (12 h light, 12 h
dark), after 1 wk, reaching 90 mm, dark olivaceous, velvety, margin irregular, rhizoid (PDA) or entire
(OA); reverse black. Diffusible pigment not produced.
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Material examined: Unknown country, unknown substrate, date and collector (holotype CBS H-23382, culture ex-type dep. A.
Arambarri LPS 325 = CBS 149.94).
Notes: Dichotomophthora brunnea was previously maintained as Di. portulacae in the CBS collection.
However, the phylogenetic analysis suggests that the strain CBS 149.94 is a distinct species.
Morphologically, the new species differs from Di. basellae and Di. lutea in having dark brown conidia and
conidiogenous cells with 1–3 lobes (vs. yellowish or pale brown conidia and conidiogenous cells with
usually more than three lobes). Neither pigment nor sclerotia were observed on the media tested.
Dichotomophthora lutea (Routien) de Hoog & Oorschot, Proc. Kon. Ned. Akad. Wetensch., Sect. C
86: 56. 1983. Fig. 12.
Basionym: Dactylaria lutea Routien, Mycologia 49: 191. 1957.
Synonym: Dichotomophthora indica Rao, Mycopath. Mycol. Appl. 28: 139. 1966. (nom. inval., Art. 35.1,
Melbourne).
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Hyphae 3–7.5 µm wide, hyaline to brown, septate, smooth to verruculose. Conidiophores
macronematous, mononematous, unbranched or irregularly branched, sometimes swollen and
repeatedly dichotomously or trichotomously branched or lobed at apex, forming a stipe and head; stipe
7.5–10(–12) µm wide, hyaline to brown, smooth; branches usually short; head 16.5–62 µm wide, pale
brown to brown. Conidiogenous cells polytretic, integrated and terminal, discrete, lobed, cicatrized,
individual lobes 8–13 × 4–11 µm. Conidia 14–65.5 × 7.5–13 µm, solitary, dry, ellipsoidal to cylindrical
rounded at ends, straight to slightly curved, subhyaline to yellow brown, 0–4-distoseptate, sometimes
constricted at septa, sometimes anastomosing conidia observed. Microconidia 12–27 × 7–13 µm,
obovoid, 0–2-distoseptate. Sclerotia 146–325 × 197–370 µm, present or absent, often formed in culture,
resembling immature perithecia, semi- or immersed in agar, globose, subglobose, ellipsoidal or ovoid,
dark brown or black. Sexual morph unknown.
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Culture characteristics: Colonies on PDA and OA at 25 °C under near-ultrav iolet light (12 h light, 12 h
dark), reaching 20–80 mm after 1 wk, white, hazel, orange, or dark grey to olivaceous, cottony, velvety,
somewhat fluffy, or flat, margin irregular, effuse; reverse centre hazel, dark brown, periphery hazel,
orange to luteous. Diffusible pigment luteous to orange (produced in some strains).
Materials examined: Argentina, isolated from soil, unknown date, J.B. Routien, (culture ex-type of Dactylaria lutea CBS 145.57).
Cuba, Santiago de las Vegas, on leaves of Portulaca oleracea (Portulacaceae), 9 Mar. 1980, G. Arnold, INIFAT A80/85 = CBS
132.81. Italy, isolated from seedbed of Pinus radiata (Pinaceae), unknown date, G. Magnani, CBS 584.71. The Netherlands, on
leaves of Portulaca oleracea (Portulacaceae), unknown date and collector, CBS 585.71; The Hague, on leaves of Portulaca
oleracea (Portulacaceae), Jul. 1978, G.H. Boerema, CBS 518.78.
Notes: In the phylogenetic tree (Fig. 13), Di. lutea is represented by five strains isolated from soil and
leaves of Po. oleraceae from Argentina, Cuba, Italy and the Netherlands. This species shows
morphological variation among strains, with different colour and aspect of the colonies, production or
absence of diffusible pigment and sclerotia. The above description is based on CBS 584.71, CBS
585.71, CBS 518.78 and CBS 132.81. The conidia were smaller than those described by de Hoog & van
Oorschot (1983) based on the ex-type strain CBS 145.57 (14–65.5 × 7.5–13 vs. 30–115 × 10–20 µm).
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Unfortunately, the ex-type strain was sterile under the culture media and conditions tested.
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Dichotomophthora portulacae Mehrl. & Fitzp. ex M.B. Ellis, Dematiaceous Hyphomycetes (Kew):
388. 1971.
Synonyms: Dichotomophthora portulacae Mehrl. & Fitzp., Mycologia 27: 550. 1935. (nom. inval., Art.
39.1, Melbourne).
Dichotomophthora portulacae Mehrl. & Fitzp. ex P.N. Rao, Mycopath. Mycol. Appl. 28: 139. 1966.
(nom. inval., Art. 38.5(a), Melbourne).
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Hyphae 1.5–6 µm wide, subhyaline to pale brown, septate, slightly constricted at septa, smooth to
verruculose. Conidiophores macronematous, mononematous, branched more or less dichotomously in
apical region, forming a stipe and head; stipe 120–220 µm long, up to 14 µm wide at apex, reddish
brown, smooth, verruculose near base, terminal branches up to 110 µm long, each ending in two slightly
swollen, rounded or angular lobes. Conidiogenous cells mono- or polytretic, integrated and terminal,
cicatrized. Conidia 45–75 × 20–30 µm, solitary, dry, ellipsoidal to cylindrical rounded at ends, dark
reddish-brown, smooth, 2–3-distoseptate. Sclerotia 120–170 µm diam, often formed in culture,
resembling perithecia, subglobose to globose or ovoid, dark reddish-brown. Sexual morph unknown
(adapted from de Hoog & van Oorschot 1983).
Culture characteristics: Colonies at 25 °C under near-ultraviolet light (1 2 h light, 12 h dark) after 1 wk, on
PDA reaching 60 mm, centre white, periphery olivaceous, cottony, margin irregular, effuse, white;
reverse centre black, periphery olive. On OA reaching 45 mm, pale greenish grey, cottony, margin
effuse, buff; reverse greenish olivaceous. Diffusible pigment not produced.
Material examined: USA, Hawaii, on Portulaca oleracea (Portulacaceae), unknown date and collector, isol. F.P. Mehrlich (exparatype culture CBS 174.35).
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Notes: In this study Di. portulacae was represented only by the ex-paratype strain, which together with
Di. brunnea (CBS 149.94) formed a subclade in Dichotomophthora. Both species have dark brown or
reddish brown conidia and conidiogenous cells with 1–3 lobes. Nevertheless, Di. portulacae has
shorter conidia with fewer septa, (45–75 × 20–30 µm; 2–3-distoseptate vs. 29–56.5 × 6–10 µm; 2–6distoseptate).
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Authors: M. Hernández-Restrepo, R. Cheewangkoon & P.W. Crous
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Gaeumannomyces Arx & D.L. Olivier, Trans. Brit. Mycol. Soc. 35: 32. 1952. Fig 16.
Synonyms: Rhaphidospora Fr., Summa veg. Scand. 2: 401. 1849.
Rhaphidophora Ces. & De Not., Sfer. Ital.: 79. 1863.
Classification: Sordariomycetes, Sordariomycetidae, Magnaporthales, Magnaporthaceae.
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Type species: Gaeumannomyces graminis (Sacc.) Arx & D.L. Olivier, basionym: Rhaphidophora
graminis Sacc. Representative strain: CPC 26020 = CBS 141384.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, tef1, rpb1. Table 6. Fig. 17.
Ascomata perithecial, superficial, submerged, globose, subglobose to elliptical, with a central, ostiolate,
cylindrical neck, dark brown to black; ascomatal wall comprised of pseudoparenchymatous cells, light or
brown. Hamathecium comprised of septate, often constricted at septa, hyaline paraphyses, widest at
base and gradually narrow at apex, exceeding asci, dissolving at maturity. Asci numerous, unitunicate,
cylindrical to elongated clavate, short stalked, with apical refractile ring, 8-spored. Ascospores cylindrical,
slightly curved to sinuate, widest in middle, ends rounded, vacuolated, septate, septa often indistinct,
hyaline to pale brown, faintly tinted yellowish in mass. Conidiophores branched, verticillate,
indeterminate, brown, often reduced to conidiogenous cells, hyaline to brown. Conidiogenous cells
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phialidic, solitary or in dense clusters, lageniform, cylindrical, straight or slightly curved tapering to a short
cylindrical to funnel-shaped or hardly visible collarette. Conidia dimorphic (A) according to Wong &
Walker (1975) “germinating phialidic conidia”: solitary, grouped in slimy heads, ovoid to cylindrical,
straight or slightly curved, tapering to an often acute base, hyaline, and/or (B) according to Wong &
Walker (1975) “non-germinating phialidic conidia”: solitary, arranged in heads, hyaline, falcate to lunate,
usually strongly curved in a semicircle with varying degrees of curvature. Hyphopodia when present
hyaline becoming brown when mature, simple or lobed. Sclerotia present or absent (adapted from
Hernández-Restrepo et al. 2016b).
Culture characteristics: Colonies on PDA mycelium mostly submerged, dark (grey olivaceous, greyish
sepia, isabelline) aerial mycelium scarce, or sometimes cottony, white; margin effuse irregular to rhizoid.
On MEA elevated, cottony to funiculose, aerial mycelium white or pale i.e. pale greenish grey, smoke
grey, submerged mycelium black, margin effuse to rhizoid. Cultures of Gaeumannomyces vary in colour,
growth rate and amount of aerial mycelium, dark hyphal strands and black sclerotia.
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Optimal media and cultivation conditions: MEA and PDA incubated at 15–30 °C depending of sp ecies.
Other methods described for production of perithecia include PDA with wheat seedlings (Speakman
1982) and flooded cultures in MPA (Speakman 1984).
Distribution: Worldwide.
Hosts: Mainly pathogens on grasses (Poaceae on Avena, Hordeum, Oryza & Leersia, Secale, Sorghum,
Triticum, xTriticale, Zea, turf grasses, buffalo grass and other grasses) and Cyperaceae, but some occur
on non-grass hosts as saprobes or endophytes.
Disease symptoms: Take-all, crown black sheath rot, dieback, root decline, patches of white heads after
flowering, stem- and root rot.
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Notes: Gaeumannomyces comprises about 20 species (Hernández-Restrepo et al. 2016b) that are
mainly pathogenic to grasses, but some species are also regarded as saprobic or endophytic. The
generic type Gaeumannomyces graminis included four varieties based on ascospore size,
hyphopodial morphology and host preferences i.e. G. graminis var. graminis, G. graminis var. avenae,
G. graminis var. tritici and G. graminis var. maydis (Turner 1940, Dennis 1960, Walker 1972, Yao et al.
1992). After a wide range of isolates were subjected to DNA sequence analyses, it was demonstrated
that these established varieties and cryptic species represent different, phylogenetically supported
species (Ward & Bateman 1999, Ulrich et al. 2000, Freeman & Ward 2004, Hernández-Restrepo
2016b). Gaeumannomyces tritici and G. avenae, the causal agents of take-all of wheat and oat
respectively, are more aggressive pathogens than G. graminis and other species in the genus.
Species of Gaeumannomyces are morphologically difficult to distinguish because of their simple
morphology, overlapping morphological features and considerable intraspecific variation.
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References: von Arx & Olivier 1952, Deacon 1973, 1974 (taxonomy); Walker 1972, 1975, 1980, 1981
(taxonomy, morphology, pathogenicity); Asher & Shipton 1981 (biology and control); Elliott 1991, Elliott
et al. 1993 (pathogenicity); Bateman et al. 1992, Augustin et al. 1999, Ulrich et al. 2000, Rachdawong et
al. 2002 (molecular data); Freeman & Ward 2004 (review); Hernández-Restrepo et al. 2016b
(morphology and phylogeny).
Authors: M. Hernández-Restrepo & P.W. Crous
Harknessia Cooke, Grevillea 9: 85. 1881. Fig. 18.
Synonyms: Caudosporella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 123: 135. 1914.
Mastigonetron Kleb., Mykol. Zentbl. 4: 17. 1914.
Cymbothyrium Petr., Sydowia 1: 148. 1947.
Classification: Sordariomycetes, Sordariomycetidae, Diaporthales, Harknessiaceae.
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Type species: Harknessia eucalypti Cooke. Representative strain: CBS 342.97.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, cal, tub2. Table 7. Fig. 19.
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Ascomata perithecial, single or aggregated, immersed, brown; necks emergent to depressed; ascomatal
wall of 3–5 layers of brown cells of textura angularis. Paraphyses hyaline, septate, dispersed between asci.
Asci 8-spored, unitunicate, cylindrical to clavate, short pedicellate, with J- apical ring. Ascospores uni- to
biseriate, ellipsoid to fusoid, hyaline, aseptate, thick-walled, guttulate, smooth-walled. Conidiomata
erumpent, scattered, pycnidial, unilocular, globose to subglobose, brown; conidiomatal wall comprising 3–4
layers of brown-walled cells of textura angularis. Macroconidiophores lining cavity or limited to a basal layer
in some species; usually reduced to conidiogenous cells, rarely septate and branched; commonly invested
in mucus. Macroconidiogenesis cells ampulliform, subcylindrical or cylindrical, hyaline, proliferating
percurrently. Macroconidia consisting of a body with a basal appendage, delimited by a septum; conidium
body unicellular, ellipsoid to fusoid, subcylindrical, globose, broadly ventricose, broadly ellipsoid or broadly
fusoid, thick-walled, smooth, brown, with or without pale and dark coloured longitudinal bands, occasionally
longitudinally striate, guttulate; basal appendages hyaline, tubular, smooth, thin-walled, often collapsing.
Microconidiophores absent or present, in same conidioma, reduced to microconidiogenous cells.
Microconidiogenous cells ampulliform or subcylindrical to lageniform, hyaline, smooth, with apical periclinal
thickening. Microconidia hyaline, smooth, aseptate, oval to ellipsoid.
Culture characteristics: Colonies spreading, fluffy, with moderate to abundant aerial mycelium, covering
plate in 1 mo. On MEA surface dirty white to cream or pale luteous; reverse cream; sometimes sporulating
with black conidiomata, oozing black masses. These culture characteristics also apply to the new taxa
described below.
Optimal media and cultivation conditions: MEA, PDA and OA under continuous near-ultraviolet light at 25 °C
to promote sporulation.
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Distribution: Worldwide.
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Hosts: On diverse gymnosperm and dicotyledonous hosts, especially on Eucalyptus (Myrtaceae), which is
host to 27 of the currently accepted 38 species.
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Disease symptoms: Associated with leaf spots, leaf tip dieback or leaf scorch and stem cankers, but
pathogenicity has not been established definitively (Crous et al. 2012c).
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Notes: Harknessia is characterised by having stromatic to pycnidial conidiomata, and dark brown conidia
with tube-shaped basal appendages, longitudinal striations, and rhexolytic secession (Crous et al. 2012c).
Sexual morphs were initially described in Cryptosporella (Nag Raj & DiCosmo 1981), which was rejected
in favour of the older genus Wuestneia (Reid & Booth 1989). However, the type species of Wuestneia,
Wu. aurea (= Wuestneia xanthostroma), was located in the Cryphonectriaceae and was associated with a
coelomycete asexual morph having hyaline conidia. Wuestneia is therefore not considered as synonym of
Harknessia, and only species placed in the Harknessiaceae and linked to Harknessia morphs were thus
transferred to Harknessia (Crous et al. 2012c).
The family Harknessiaceae was introduced based on LSU sequences of taxa belonging to
Diaporthales in order to accommodate Harknessia (Crous et al. 2012c).
References: Lee et al. 2004 (morphology and phylogeny); Crous et al. 2012c (morphology and
phylogeny).
Harknessia bourbonica Crous & M.J. Wingf., sp. nov. MycoBank MB824016. Fig. 20.
Etymology: Name refers to Île Bourbon, the original name of La Réunion Island.
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Caulicolous and foliicolous, isolated from leaves and twigs incubated in moist chambers (presumed
endophyte). Conidiomata up to 300 µm diam, pycnidial, separate to gregarious, subepidermal, becoming
erumpent, stromatic, amphigenous, depressed globose; with irregular opening and border of yellowish,
furfuraceous cells; conidiomatal wall of textura angularis. Conidiophores reduced to conidiogenous cells
lining conidiomatal cavity. Conidiogenous cells 8–10 × 4–8 µm, ampulliform to subcylindrical, hyaline,
smooth, invested in mucilage, percurrently proliferating once or twice near apex. Conidia (12–)13–14(–
15) × (8–)9–10 µm in vitro, broadly ventricose to ellipsoid, apex subobtusely rounded, aseptate, nonapiculate, yellow-brown, thick-walled, striations in restricted areas, multi-guttulate. Basal appendage (5–
)8–12 × 2–2.5 µm in vitro, hyaline, tubular, smooth, thin-walled, devoid of cytoplasm. Microconidia not
seen.
Material examined: France, La Réunion, 21°15'5.4"S 55°36'3.3"E, on leaf litt er of Eucalyptus robusta (Myrtaceae), 8 Mar. 2015,
P.W. Crous & M.J. Wingfield (holotype CBS H-23387, culture ex-type CBS 143913 = CPC 26533).
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Notes: Harknessia bourbonica is related to Ha. ravenstreetina, which was also isolated from Eucalyptus
leaves. The two species are distinguished in that Ha. ravenstreetina has longer conidia (14−20 m) that
lack striations and has shorter basal appendages (1.5−5 × 2–2.5 m).
Harknessia corymbiae Crous & A.J. Carnegie, sp. nov. MycoBank MB824017. Fig. 21.
Etymology: Name refers to the host genus, Corymbia.
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Caulicolous and foliicolous, isolated from leaves and twigs incubated in moist chambers (presumed
endophyte). Conidiomata up to 250 µm diam, pycnidial, separate to gregarious, subepidermal, becoming
erumpent, stromatic, amphigenous, depressed globose; with irregular opening and border of yellowish,
furfuraceous cells; conidiomatal wall of textura angularis. Conidiophores reduced to conidiogenous cells
lining conidiomatal cavity. Conidiogenous cells 6–10 × 4–5 µm, ampulliform to subcylindrical, hyaline,
smooth, invested in mucilage, percurrently proliferating once or twice near apex. Conidia (23–)25–28(–
30) × (8–)9 µm in vitro, subcylindrical, apex apiculate, aseptate, yellow-brown, thick-walled, lacking
striations, granular; in lactic acid some conidia appear to have a central line of paler pigment. Basal
appendage (50–)65–80(–100) × 3–4 µm in vitro, hyaline, tubular, smooth, thin-walled, devoid of
cytoplasm. Microconidia 3–4 × 1.5–2 µm, hyaline, smooth, guttulate, aseptate, subcylindrical with obtuse
ends.
Material examined: Australia, New South Wales, Bom Bom State Forest, on leaf litter of Corymbia maculata (Myrtaceae), 13 Mar.
2017, A.J. Carnegie (holotype CBS H-23388, culture ex-type CPC 33289).
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Notes: Harknessia corymbiae was located in a distinct clade distant from the other species of the genus.
The only accepted species presently known from Corymbia is Ha. rhabdosphaera. Both species were
collected from Australia, but Ha. rhabdosphaera has smaller, striated conidia [(13–)15–17 × (13–)14–15
µm] with short basal appendages (up to 5 µm long).
Harknessia cupressi Crous & R.K. Schumach., sp. nov. MycoBank MB824018. Fig. 22.
Etymology: Name refers to the host genus, Cupressus.
Caulicolous and foliicolous, isolated from needles incubated in moist chambers (presumed endophyte).
Conidiomata up to 250 µm diam, pycnidial, separate to gregarious, subepidermal, becoming erumpent,
stromatic, amphigenous, depressed globose; with irregular opening and border of yellowish, furfuraceous
cells; conidiomatal wall of textura angularis. Conidiophores reduced to conidiogenous cells lining
conidiomatal cavity. Conidiogenous cells 5–10 × 3–5 µm, ampulliform to subcylindrical, hyaline, smooth,
invested in mucilage, percurrently proliferating once or twice near apex. Conidia (20–)21–23(–25) × (8–
)9–11(–13) µm in vitro, broadly ventricose, apex apiculate, aseptate, yellow-brown, thick-walled, striations
in restricted areas, multi-guttulate. Basal appendage 2–5(–12) × 2–2.5 µm in vitro, hyaline, tubular,
smooth, thin-walled, devoid of cytoplasm. Microconidia 4–7 × 3–4 µm, hyaline, smooth, aseptate, broadly
ellipsoid.
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Materials examined: Spain, Zaragoza, Carretera El Frago, on needles of Cupressus sempervirens (Cupressaceae), 7 Jan.
2016, coll. R. Blasco, det. R.K. Schumacher (holotype CBS H-23389, culture ex-type CBS 143914 = CPC 30192); ibid., CPC
30174.
Notes: Harknessia cupressi was located in an independent clade distant from the other species of the
genus. Harknessia cupressi is the only species known from Cupressus sempervirens.
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Harknessia pilularis Crous & A.J. Carnegie, sp. nov. MycoBank MB824020. Fig. 23.
Etymology: Name refers to Eucalyptus pilularis, the host species from which this fungus was isolated.
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Caulicolous and foliicolous, isolated from leaves and twigs incubated in moist chambers (presumed
endophyte). Conidiomata up to 250 µm diam, pycnidial, separate to gregarious, subepidermal, becoming
erumpent, stromatic, amphigenous, depressed globose; with irregular opening and border of yellowish,
furfuraceous cells; conidiomatal wall of textura angularis. Conidiophores reduced to conidiogenous cells
lining conidiomatal cavity. Conidiogenous cells 4–7 × 2–5 µm, ampulliform to subcylindrical, hyaline,
smooth, invested in mucilage, percurrently proliferating once or twice near apex. Conidia (13–)14–16(–
20) × (8–)11–12(–13) µm in vitro, globose to rarely broadly ellipsoid, apex obtusely rounded, aseptate,
non-apiculate, yellow-brown, thick-walled, striations covering entire conidial body, multi-guttulate. Basal
appendage (2–)3–5 × 2–2.5 µm in vitro, hyaline, tubular, smooth, thin-walled, devoid of cytoplasm.
Microconidia not seen.
Materials examined: Australia, New South Wales, Pine Creek State Forest, 30.405423S 152.932698E, on leaves of Eucalyptus
pilularis (Myrtaceae), 23 Jan. 2017, A.J. Carnegie (holotype CBS H-23391, culture ex-type CPC 33218); ibid., CPC 33356.
Notes: Harknessia pilularis is related to Ha. rhabdosphaera, but Ha. rhabdosphaera produces longer
conidiogenous cells (7–15 × 4–6 µm), and wider conidia (13–15 µm).
Authors: Y. Marin-Felix, A.J. Carnegie, M.J. Wingfield, R.K. Schumacher & P.W. Crous
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Huntiella Z.W. de Beer, et al., Stud. Mycol. 79: 211. 2014. Fig. 24.
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Classification: Sordariomycetes, Hypocreomycetidae, Microascales, Ceratocystidaceae.
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Type species: Huntiella moniliformis (Hedgc.) Z.W. de Beer, et al., basionym: Ceratostomella moniliformis
Hedgc. Holotype: BPI 595959.
DNA barcodes (genus): LSU, 60S, mcm7.
DNA barcodes (species): ITS, mcm7, tef1, tub2. Table 8. Fig. 25.
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Ascomata perithecial, ellipsoidal, subglobose, globose to obpyriform or ovoid; bases ornamented with
dark brown to black conical spines or pale brown, unbranched hyphae; necks long, straight or slightly
curved, tapering towards apex, dark brown to black at base, becoming paler towards apex, ostiolate, with
a disciform base; hyphae on neck hyaline, not divergent, straight or convergent. Asci evanescent.
Ascospores hyaline, aseptate, in face view subglobose, in side view ellipsoidal giving an impression of a
hat, with hood-like gelatinous sheath. Conidiophores macronematous, rarely branched, septate,
occasionally reduced to conidiogenous cells. Conidiogenous cells hyaline, enteroblastic, mostly of two
types, lageniform, producing rectangular-shaped conidia and cylindrical, producing barrel-shaped conidia.
Conidia aseptate, majority of species have two distinct shapes: bacilliform, hyaline, or barrel-shaped or
oblong to ellipsoidal, hyaline or subhyaline. Chlamydospores absent.
Culture characteristics: On 2 % MEA aerial mycelium abundant, colonies white, yellow green to brown,
smoke grey, dark olive to black. Some species produce aromas: Hu. bhutanensis produces an
unpleasant rotten odour, Hu. decipiens, Hu. moniliformis and Hu. salinaria a pleasant banana-oil aroma,
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Hu. moniliformopsis little to no distinct odour, and Hu. omanensis fruity aroma that turns to a fermented
odour with age.
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Optimal media and cultivation conditions: On 2 % MEA in dark, optimum growth varies between species:
Hu. sublaevis 20–30 °C; Hu. bhutanensis, Hu. oblonga, Hu. ceramica and Hu. tribiliformis 20–25 °C; Hu.
decipiens, Hu. chinaeucensis, Hu. inquinans, Hu. microbasis, Hu. salinaria and Hu. sumatrana 25–30 °C;
Hu. cryptoformis, Hu. omanensis, Hu. savannae and H. tyalla 30–35 °C.
Distribution: Australia, Bhutan, China, Ecuador, Indonesia, Malawi, Oman, South Africa and Tasmania.
Hosts: Acacia (Fabaceae), Combretum and Terminalia (Combretaceae), Eucalyptus (Myrtaceae),
Mangifera (Anacardiaceae) associated with Cryphalus scabricollis (bark beetle, Scolytinae), Picea
(Pinaceae) infested with Ips schmutzenhoferi (bark beetle, Scolytinae), Pinus (Pinaceae), Ziziphus
(Rhamnaceae), and Staphilinid (rove beetle, Staphylinidae)
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Disease symptoms: Huntiella species do not produce distinct disease symptoms on their hosts in nature.
They usually infect freshly made wounds on trees and infections are often associated with nitidulid
beetles (Coleoptera: Nitidulidae) and flies (Diptera). Pathogenicity tests using some of the species have
given rise to lesions under controlled environments, but, based on the size of lesions and failure to reisolate the fungus from these lesions, they are generally considered not to be primary pathogens (Tarigan
et al. 2010, van Wyk et al. 2011, de Beer et al. 2014).
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Notes: Huntiella is one of nine genera in the recently erected family, Ceratocystidaceae (De Beer et al.
2014, Mayers et al. 2015, Nel et al. 2017). The genus was proposed to accommodate Ceratocystis
moniliformis and related species that form a well-defined monophyletic lineage within the group previously
treated as Ceratocystis sensu lato (Wingfield et al. 2013). Huntiella currently includes 17 species.
Species of Huntiella have several features in common, namely conical spines on their ascomatal
bases, disk-like structures at the bases of the ascomatal necks, which break off easily, hat-shaped
ascospores and one to two types of conidia (De Beer et al. 2014). Although morphological and culture
characteristics overlap between the species, some species can be differentiated. These include Hu.
sublaevis that has a limited number of spines on the ascomata (Van Wyk et al. 2011), and Hu.
chinaeucensis and Hu. microbasis that have only rectangular-shaped conidia (Tarigan et al. 2010, Chen
et al. 2013). With the exception of Hu. ceramica that is known only from the asexual morph, all other
species have both sexual and asexual morphs. Different Huntiella species exhibit a variety of sexual
strategies, with Hu. omanensis undergoing heterothallic mating while MAT2 isolates of Hu. moniliformis
are able to reproduce unisexually (Wilson et al. 2015).
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References: Kamgan et al. 2008, Heath et al. 2009, Tarigan et al. 2010, Kamgan Nkuekam et al. 2012,
Chen et al. 2013, Kamgan Nkuekam et al. 2013 (pathogenicity); De Beer et al. 2013a (higher
classification); De Beer et al. 2013b (nomenclature); Wingfield et al. 2013, De Beer et al. 2014 (generic
definitions and phylogenetic relationships); Wilson et al. 2015 (mating strategies).
Huntiella abstrusa A.M. Wilson, Marinc., M.J. Wingf., sp. nov. MycoBank MB821072. Fig. 26.
Etymology: Name refers to the fact that this cryptic species was obscured by the name Ceratocystis
moniliformis for more than a decade.
On MEA: Ascomata 145–315 × 130–275 µm, perithecial, embedded in media or superficial in mycelial
mass, single or in groups, pale brown when young, becoming dark brown with age, ellipsoidal to
subglobose; asocomata wall textura epidermoidea to textura globulosa, covered with short sterile hyphae,
30–105 µm long, hyaline becoming pale brown with age, mostly unbranched, flexuous, tapering towards
apex, with conical spines; necks 595–1100 µm long, 21–71.5 µm wide near base, 11–18 µm wide near
apex, dark brown, tapering towards apex, becoming easily detached from ascomatal base when pressed,
with a disk-like structure near base; hyphae near apex 18.5–37 µm long, 1–2 µm wide near base, 0.5–1.5
µm wide, mostly straight, showing no distinct divergent nor convergent, aseptate, unbranched, hyaline,
tapering towards apex. Asci not observed. Ascospores 4.5–5.5 × 3.5–6 µm (av. 5.2 × 4.6 µm) without
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sheath, 2–3.5 µm thick in side-view, hyaline, oblong to subglobose, clothed with gelatinous sheath, 0.5–
1.5 µm thick, in side-view giving an impression of a hat. Conidiophores up to 80 µm long, septate,
flexuous, sometimes branched, occasionally reduced to conidiogenous cells, often lightly sub-hyaline to
pale brown near base. Conidiogenous cells hyaline, enteroblastic, in two shapes, originating from same
or different hyphae; lageniform, 15.5–33 µm long, 2–4.5 µm wide at base, gradually tapering towards
apex, 1–3 µm wide at apex, producing rectangular conidia; or cylindrical with wide-mouthed, 22–50 µm
long, 3.5–6.5 µm wide at apex, 3–4 µm wide at base, producing barrel-shaped conidia. Conidia in chains,
hyaline, aseptate, in two shapes, rectangular, 4–8.5 × 1.5–2.5 µm (av. 5.6 × 1.8 µm); or barrel-shaped, 5–
14.5 × 4–7 µm (av. 8.1 × 5.9 µm). Chlamydospores absent.
Culture characteristics: On 2 % MEA optimum growth at 30 °C reaching 82.7 mm in dark in 3 d, followed
by 25 °C reaching 77.3 mm, showing slow growth at 1 0, 15, 20, 35 °C. Cultures circular with smooth
margins, aerial mycelium fluffy to velvety, moderately dense, above and below dark brown fading towards
edge and with white margins.
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Material examined: Indonesia, Riau province, Teso East, S 0°04’33.00”, E 101°37 ’23.00”, on the bark of Eucalyptus sp.
(Myrtaceae), Oct. 2005, M. Tarigan (holotype PREM 61671, culture ex-type CBS 142243 = CMW 21092).
Notes: For more than a decade, Hu. abstrusa was thought to be Ceratocystis moniliformis (now Hu.
moniliformis). Phylogenetic analyses of ITS, LSU, 60S, mcm7 and tub2 have shown that Hu. abstrusa is
distinct from Hu. moniliformis and other Huntiella spp. It is most closely related to Hu. inquinans, Hu.
microbasis and Hu. sumatrana from Indonesia, Hu. chinaeucensis from China and Hu. bhutanensis from
Bhutan (De Beer et al. 2014, Van Wyk et al. 2004, Tarigan et al. 2010, Chen et al. 2013). Neither Hu.
abstrusa nor any of the other Huntiella species are primary pathogens and typically infect freshly made
wounds on trees. Despite only minor morphological differences between Huntiella species, Hu. abstrusa
can be distinguished from the other species by its longer necks and the presence of barrel-shaped
conidia. The Hu. abstrusa isolate examined is considered heterothallic.
Authors: A.M. Wilson, S. Marincowitz, M.J. Wingfield & B.D. Wingfield
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Macgarvieomyces Klaubauf et al., Stud. Mycol. 79: 106. 2014. Fig. 27.
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Classification: Sordariomycetes, Sordariomycetidae, Magnaporthales, Pyriculariaceae.
Type species: Macgarvieomyces borealis (de Hoog & Oorschot) Klaubauf et al., basionym: Pyricularia
borealis de Hoog & Oorschot. Holotype and ex-type strain: IMI 105288, CBS 461.65.
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DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, act, cal, rpb1. Table 9. Fig. 28.
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Mycelium consisting of smooth, hyaline, branched, septate hyphae. Conidiophores solitary, erect,
straight or curved, mostly unbranched, medium to dark brown, smooth or finely verruculose, septate.
Conidiogenous cells integrated, terminal, rarely intercalary, medium to dark brown, smooth or finely
verruculose, forming a rachis with protruding denticles, appearing flat-tipped. Conidia solitary, narrowly
obclavate to narrowly pyriform, hyaline, often becoming pale brown with age, smooth, granular, guttulate,
medianly 1-septate, apex obtusely rounded; hila somewhat thickened, refractive or not, not or slightly
darkened. Chlamydospores brown, ellipsoid, arranged in chains (adapted from Klaubauf et al. 2014).
Culture characteristics: Colonies on MEA buff to rosy buff, isabelline or pale luteous, with pale olivaceous
grey central mycelium, with entire, lobate or round and hairy edge, umbonate to conical or flat to slightly
raised colony with somewhat velvety or wool-like texture; reverse iron grey, ochreous and buff towards
edge or pale luteous. On CMA and OA transparent, pale luteous to olivaceous or grey olivaceous, flat,
smooth and velutinous surface, undulate margin. Colonies on PDA pale luteous, white with buff centre or
whitish to buff with honey centre, round, flat, fringed margin, reverse white with buff centre or whitish to
buff with honey centre.
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Optimal media and cultivation conditions: On OA at 25 °C under dark, or autoclaved barley s eeds placed
on SNA at 25 °C under near-ultraviolet light (12 h light, 12 h dark).
Distribution: Europe, New Zealand.
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Hosts: Primarily on Juncus effusus and Luzula spp. (Juncaceae), also reported on Carex sp. and
Kyllinga brevifolia (Cyperaceae).
Disease symptoms: Leaf spots.
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Notes: Macgarvieomyces was recently introduced to accommodate two species previously placed in
Pyricularia. Phylogenetic analyses based on LSU, ITS, act, cal and rpb1 demonstrated that these taxa
are not congeneric with Pyricularia s. str. (Klaubauf et al. 2014). Species in this genus were isolated
from Juncaceae in Europe and associated with leaf spots. Species have also been reported on
Cyperaceae and New Zealand (Farr & Rossman 2017); however, these host and distribution data
have not been corroborated based on DNA sequence analyses.
Reference: Klaubauf et al. 2014 (morphology and phylogeny).
Macgarvieomyces luzulae (Ondřej) Y. Marín, Akulov & Crous, comb. nov. MycoBank MB823764. Fig.
29.
Basionym: Pyricularia luzulae Ondřej, Česká Mykol. 42: 81. 1988.
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Leaf spots up to 17 mm long, ellipsoid to fusiform, grey to pale brown at middle, dark brown at margin.
On SNA: Mycelium with hyaline, smooth, septate, branched, 2–3 µm diam hyphae. Conidiophores 60–
120 × 4–7 µm, erect, dark brown, unbranched, subcylindrical, straight to flexuous, thick-walled, finely
verruculose, 2–3-septate. Conidiogenous cells 30–50 × 4–6 µm, terminal and subcylindrical, finely
verruculose, dark brown, tapering toward apex with numerous denticles pointing upwards, 1–4 × 1–1.5
µm; scars unthickened. Conidia (18–)20–22(–30) × (4–)5(–6) µm, solitary, narrowly pyriform, hyaline,
becoming pale brown with age, guttulate, 1(–2)-septate, apex obtusely rounded, base truncate, 2 µm
diam, slightly darkened, refractive.
Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and smooth,
lobate margins, covering plate after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse pale
luteous.
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Materials examined: Slovakia, on leaves of Luzula sylvatica (Juncaceae), Jul. 1984 (holotype of Pyricularia luzulae PRM
842743). Ukraine, on leaves of Luzula sp. (Juncaceae), 2016, A. Akulov (epitype of Pyricularia luzulae designated here
MBT379806, CBS H-23355, culture ex-epitype CBS 143401 = CPC 32458); ibid., CWU (Myc) AS 5966 / 6437; Carpathian
Biosphere reserve, on leaves of Luzula sylvatica (Juncaceae), Aug. 2016, A. Akulov, CPC 31555; ibid., CPC 31571.
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Notes: Pyricularia luzulae was introduced for a fungus isolated on Luzula sylvatica from Slovakia (Ondřej
1988). In the original description PRM 842743 was designated as holotype, but no living culture was
associated with this specimen. Therefore, a strain that closely fits the description of the protologue, and
isolated from the same host in a close country to Slovakia, namely Ukraine, is designated here as
epitype. The only difference observed was in the conidial size, since in the original description (in vivo)
the conidia were larger and 1–2-septate (17.5–36 × 3.5–7.5 µm). However, the measurements of our
isolate growing in vivo are 27.5–33 × 6.5–7.5 µm, and in vitro 18–30 × 4–6 µm, and conidia could
become 2-septate with age.
The phylogenetic study of Py. luzulae revealed that it belongs to Macgarvieomyces. As noted above,
the host of this genus is Juncus effusus in the same family as Luzula (Juncaceae).
Authors: Y. Marin-Felix, A. Akulov & P.W. Crous
Metulocladosporiella Crous et al., Mycol. Res. 110: 269. 2006. Fig. 30.
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Classification: Eurotiomycetes, Chaetothyriomycetidae, Chaetothyriales, Herpotrichiellaceae.
Type species: Metulocladosporiella musae (E.W. Mason) Crous et al., basionym: Cladosporium musae
E.W. Mason. Lectotype designated by Crous et al. (2006a): IMI 7521 (slide). Epitype and ex-epitype
strain designated by Crous et al. (2006a): CBS H-14788, CBS 161.74 = ATCC 36973.
DNA barcodes (species): ITS, tef1. Table 10. Fig. 31.
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DNA barcodes (genus): LSU, ITS.
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Mycelium internal and external on substrate, hyphae branched, septate, hyaline, subhyaline to pale
olivaceous, thin-walled. Stromata lacking. Conidiophores macronematous, mononematous,
occasionally with intermixed micronematous conidiophores, solitary or in loose groups, arising from
hyphae, erect, with a long, subcylindrical, simple stipe and a branched terminal part; stipe septate,
medium to brown, smooth or almost so, usually swollen at base; branched part loose to dense,
metuloid, of short to long branchlets and ramoconidia, tips paler than stipes, subhyaline to pale
olivaceous. Conidiogenous cells integrated, terminal, occasionally intercalary, polyblastic, sympodial;
conidiogenous loci (conidial scars) subconspicuous to conspicuous, subdenticulate, truncate,
unthickened to slightly thickened, and somewhat darkened-refractive. Conidia and ramoconidia in
simple and branched chains, ellipsoid, ovoid, subcylindrical, or fusiform, 0–1-septate, subhyaline to
pale olivaceous, thin-walled, smooth; hila truncate, unthickened to slightly thickened or slightly
darkened-refractive; secession schizolytic. Sexual morph unknown (adapted from Crous et al. 2006a).
Culture characteristics: Colonies on PDA and OA under near-ultraviolet light with smooth, regular
margins and sparse to moderate aerial mycelium; surface on PDA pale mouse grey to mouse grey or
dirty white-grey, greyish sepia, smoke grey to grey olivaceous, or olivaceous; reverse on PDA greenish
black, cinnamon to isabelline with centre fuscous black, or grey olivaceous to dark grey olivaceous or
olivaceous black.
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Optimal media and cultivation conditions: SNA under near-ultraviolet light at 25 °C to indu ce sporulation.
Hosts: Musa spp. (Musaceae).
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Distribution: Africa, America, Asia and Oceania.
Disease symptoms: Leaf spots; Cladosporium speckle disease of banana.
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Notes: Metulocladosporiella was introduced by Crous et al. (2006a) to accommodate two
cladosporium-like species causing speckle disease on banana. Metulocladosporiella can be
distinguished from Cladosporium and allied genera by the presence of apically branched, brown
conidiophores with paler tips and chains of pale, smooth, often subhyaline conidia.
The phylogenetic analysis based on the ITS and LSU sequences demonstrated that
Metulocladosporiella belongs in Chaetothyriales. Morphologically, the conidiogenous loci and conidial
hila resemble those of Cladophialophora, which is another member of this order. However,
Cladophialophora produces unbranched, micro- to semimacronematous conidiophores, and
concolourous conidia. Moreover, Cladophialophora includes human pathogenic species (Crous et al.
2006a). In the present study, the recommended barcodes for species delimitation are ITS and tef1.
Hitherto, all species described in Metulocladosporiella are pathogens of banana and occur in
countries where this crop is cultivated (Jones 2000, Crous et al. 2006a).
Reference: Crous et al. 2006a (morphology and phylogeny).
Metulocladosporiella chiangmaiensis Y. Marín, Cheew. & Crous, sp. nov. MycoBank MB824031.
Fig. 32.
Etymology: Name from Chiang Mai, the province in Thailand where this fungus was collected.
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Mycelium internal and external, superficial; hyphae 1.5–3.5 µm wide, branched, septate, occasionally
slightly constricted at septa, with small swellings, hyaline, subhyaline to pale olivaceous, thin-walled,
smooth, hyphae occasionally aggregated, forming ropes; with numerous intermixed micronematous
conidiophores, erect from vegetative mycelium, intercalary, straight to flexuous, unbranched, subhyaline,
usually with simple terminal conidial chains. Macronematous conidiophores 85–1050 µm long, arising
from superficial hyphae, erect, solitary to loosely aggregated, composed of a subcylindrical stipe, 5.5–
9.5(–10.5) µm wide, 3–18 septate, swollen or lobed at base, 8–17(–18.5) µm diam, with short, rhizoid,
subhyaline to pale brown hyphae growing from base, medium to brown in lower half, paler towards apex,
tips pale brown or even subhyaline, thick-walled below, thin-walled towards apex, smooth; apex
persistently branched, branched part composed of usually fairly compact, closely arranged subcylindrical
branchlets; primary branches (13–)17–45(–55) × 3–5.5 µm, 0(–1)-septate, giving rise to 1–3 secondary
branches, or to conidiogenous cells; secondary branches 13–28 × 2.5–5(–5.5) µm, 0(–1)-septate, giving
rise to 1–2(–3) conidiogenous cells; conidiogenous cells 9–17 × 3–5 µm, subcylindrical, terminal or
occasionally intercalary, sympodial, polyblastic, conidiogenous loci 1–1.5(–2) µm wide, subconspicuous
to conspicuous, subdenticulate, somewhat protuberant, truncate, wall unthickened to somewhat so,
darkened-refractive. Conidia 5.5–10(–12.5) × 2.5–4 µm, in simple and branched acropetal chains,
ellipsoid-ovoid, fusoid, subcylindrical, aseptate, subhyaline to pale brown, thin-walled, smooth, with 1–3(–
4) hila, 1–1.5 µm diam, up to 2 µm diam at base of ramoconidia, truncate, unthickened or almost so, and
somewhat darkened-refractive, secession schizolytic; ramoconidia 7–15(–17) × 3–4.5 µm.
Culture characteristics: Colonies on PDA reaching 25–28 mm diam after 2 wk at 25 °C, moderate
aerial mycelium, velvety, umbonate; surface greyish sepia, halo surrounding centre vinaceous buff,
and margins hazel; reverse cinnamon to isabelline, centre fuscous black. Colonies on OA reaching
32–35 mm diam after 2 wk at 25 °C, sparse aerial my celium, flat except margins due to aerial
mycelium; surface hazel, centre brown vinaceous, margins smoke grey; reverse livid vinaceous,
centre brown vinaceous, margins pale brown vinaceous.
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Material examined: Thailand, Chiang Mai Province, Mae Klang Luang, N 18° 32.46 5', E 98° 32.874', from leaves of Musa sp.
(Musaceae), 6 Oct. 2008, P.W. Crous & R. Cheewangkoon (holotype CBS H-23393, culture ex-type CBS 143918= CPC
18646).
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Notes: Metulocladosporiella chiangmaiensis is related to M. musigena. Both species are known from
Thailand, and produce more micronematous conidiophores than any other species in the genus.
However, M. chiangmaiensis produces longer macronematous conidiophores and shorter conidia than
M. musigena, which produces conidiogenous cells directly from the apex of its macronematous
conidiophores, which is not the case in M. chiangmaiensis.
Metulocladosporiella malaysiana Y. Marín & Crous, sp. nov. MycoBank MB824032. Fig. 33.
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Etymology: Name refers to Malaysia, the country from where it was isolated.
Mycelium internal and external, superficial; hyphae 1–3.5 µm wide, branched, septate, occasionally
constricted at septa, with swellings, hyaline, subhyaline to pale brown, thin-walled, smooth, hyphae
occasionally aggregated, forming ropes; occasionally with intermixed micronematous conidiophores,
erect from vegetative mycelium, intercalary, straight to flexuous, unbranched, subhyaline, usually with
simple terminal conidial chains. Macronematous conidiophores 57–565 µm long, arising from superficial
hyphae, erect, solitary to loosely aggregated, composed of a subcylindrical stipe, 5–8 µm wide, (1–)2–11septate, swollen or lobed at base, 7–15 µm diam, with short, rhizoid, subhyaline to pale brown hyphae
growing from base, medium to brown in lower half, paler towards apex, tips pale brown or even
subhyaline, thick-walled below, thin-walled towards apex, smooth; apex persistently branched, composed
of fairly compact, closely arranged subcylindrical branchlets; primary branches (12–)16–44(–60) × 3.5–
5.5 µm, 0–1(–2)-septate, giving rise to 1–2(–3) secondary branches, or to conidiogenous cells; secondary
branches 11.5–26.5(–39) × 3–4.5 µm, 0(–1)-septate, giving rise to 1–2(–3) conidiogenous cells;
conidiogenous cells 10–23 × 2.5–4.5 µm, subcylindrical, terminal or occasionally intercalary, sympodial,
polyblastic, conidiogenous loci subconspicuous to conspicuous, subdenticulate, somewhat protuberant,
ACCEPTED MANUSCRIPT
truncate, wall not to slightly thickened, darkened-refractive, 1–2 µm wide. Conidia 5.5–13.5(–14.5) × (2.5–
)3–4.5(–5) µm, in simple and branched acropetal chains, ellipsoid-ovoid, fusoid, subcylindrical, aseptate,
subhyaline to pale brown, thin-walled, smooth, with 1–3 hila, 0.8–1.4 µm diam, up to 2 µm diam at base of
ramoconidia, truncate, not to slightly thickened, and somewhat darkened-refractive, secession schizolytic;
ramoconidia 7–13(–14) × 3–4.5(–5) µm.
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Culture characteristics: Colonies on PDA reaching 23–27 mm diam after 2 wk at 25 °C, moderate
aerial mycelium, giving cottony appearance, umbonate, margins fringed; surface olivaceous, mycelium
smoke grey; reverse grey olivaceous to dark grey olivaceous. Colonies on OA reaching 30–34 mm
diam after 2 wk at 25 °C, sparse aerial mycelium, u mbonate, margins fringed; surface olivaceous,
mycelium smoke grey to grey olivaceous; reverse dark grey olivaceous.
Material examined: Malaysia, from leaves of Musa sp. (Musaceae), 2010, M.H. Wong (holotype CBS H-23394, culture ex-type
CBS 143919 = CPC 18131).
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Notes: Metulocladosporiella malaysiana is related to M. samutensis. These can be easily distinguished
based on the length of the macronematous conidiophores (57–565 in M. malaysiana vs. 200–1120 µm
in M. samutensis), and the almost total absence of secondary branches in M. samutensis. Moreover,
M. malaysiana produces fewer micronematous condiophores than the other species of
Metulocladosporiella.
Metulocladosporiella musigena Y. Marín, Cheew. & Crous, sp. nov. MycoBank MB824033. Fig. 34.
Etymology: Name refers to Musa, the host from which it was isolated.
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Mycelium internal and external, superficial; hyphae 1–4 µm wide, branched, septate, occasionally slightly
constricted at septa, with small swellings, hyaline, subhyaline to pale olivaceous, thin-walled, smooth,
hyphae occasionally aggregated, forming ropes; with a great amount of intermixed micronematous
conidiophores, erect from vegetative mycelium, intercalary, straight to flexuous, unbranched, subhyaline,
usually with simple terminal conidial chains. Macronematous conidiophores (115–)170–780 µm long,
arising from superficial hyphae, erect, solitary to loosely aggregated, composed of a subcylindrical stipe,
4.5–7.5 µm wide, 5–14 septate, swollen or lobed at base, 7.5–14.5(–18.5) µm diam, with short, rhizoid,
subhyaline to pale brown hyphae growing from base, medium to brown in lower half, paler towards apex,
tips brown, pale brown or subhyaline, thick-walled below, thinner-walled towards apex, smooth; apex
usually persistently branched, branched part composed of usually fairly compact, closely arranged
subcylindrical branchlets, or sometimes giving rise directly to conidiogenous cells; primary branches 19–
32.5(–43.5) × 3–4.5 µm, 0(–1)-septate, giving rise to 1–3 secondary branches, or to conidiogenous cells;
secondary branches 15.5–31(–38) × 3–4.5 µm, 0(–1)-septate, giving rise to 1–3 conidiogenous cells;
conidiogenous cells 13.5–28(–39.5) × 2.5–5 µm, subcylindrical, terminal or occasionally intercalary,
sympodial, polyblastic, conidiogenous loci subconspicuous to conspicuous, subdenticulate, somewhat
protuberant, truncate, wall unthickened to somewhat so, darkened-refractive, 1–2 µm wide. Conidia 5–
15.5(–19.5) × 2–3.5 µm, in simple and branched acropetal chains, ellipsoid-ovoid, fusoid, subcylindrical,
aseptate, subhyaline to pale brown, thin-walled, smooth, with 1–3(–4) hila, 0.8–1.2 µm diam, up to 1.7 µm
diam at base of ramoconidia, truncate, unthickened or almost so, and somewhat darkened-refractive,
secession schizolytic; ramoconidia 8–20.5 × 2.5–4 µm.
Culture characteristics: Colonies on PDA reaching 21–24 mm diam after 2 wk at 25 °C, moderate
aerial mycelium, powdery, umbonate; surface smoke grey to grey olivaceous; reverse grey olivaceous
to olivaceous black. Colonies on OA reaching 31–34 mm diam after 2 wk at 25 °C, moderate aerial
mycelium, powdery, umbonate, slightly lobate; surface dark olivaceous, mycelium smoke grey to grey
olivaceous; reverse dark slate blue.
Material examined: Thailand, Chiang Mai Province, Mae Rim District, Queen Sirikit Botanic Garden, from Musa sp. (Musaceae),
19 Jul. 2008, P.W. Crous & R. Cheewangkoon (holotype CBS H-23395, culture ex-type CBS 143920 = CPC 31490).
Notes: Metulocladosporiella musigena produces macronematous conidiophores directly producing
conidiogenous cells at the apex, or composed of fairly compact branches, which are not observed in
ACCEPTED MANUSCRIPT
the other species of the genus. Moreover, it is characterised by the production of an excessive number
of micronematous conidiophores.
Metulocladosporiella samutensis Y. Marín, Luangsa-ard & Crous, sp. nov. MycoBank MB824034.
Fig. 35.
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Etymology: Name from Samut Songkhram, the province in Thailand from where it was isolated.
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Mycelium internal and external, superficial; hyphae 1.5–4 µm wide, branched, septate, occasionally
slightly constricted at septa, with small swellings, hyaline, subhyaline to pale brown, thin-walled, smooth,
hyphae occasionally aggregated, forming ropes; with intermixed micronematous conidiophores, erect
from vegetative mycelium, intercalary, straight to flexuous, unbranched, subhyaline to pale brown, usually
with simple terminal conidial chains. Macronematous conidiophores (200–)425–1000(–1120) µm long,
arising from superficial hyphae, erect, solitary to loosely aggregated, composed of a subcylindrical stipe,
5.5–7.5(–8) µm wide, (4–)8–21(–29)-septate, swollen or lobed at base, 7–14.5(–20.5) µm diam, with short
rhizoid subhyaline to pale brown hyphae growing from base, medium to brown in lower half, paler towards
apex, tips pale brown or occasionally subhyaline, thick-walled below, thinner towards apex, smooth; apex
giving rise directly to 1–2(–3) conidiogenous cells, or branched, branched part composed of loosely
arranged subcylindrical branchlets; primary branches 24.5–39(–44) × 3–4(–5) µm, 0–1-septate, giving
rise to 1–2(–3) conidiogenous cells, or rarely 1–2 secondary branches; conidiogenous cells (12–)14–25(–
32) × 3.5–5 µm, subcylindrical, terminal or occasionally intercalary, sympodial, polyblastic, conidiogenous
loci subconspicuous to conspicuous, subdenticulate, somewhat protuberant, truncate, wall unthickened to
somewhat so, darkened-refractive, 1–2 µm wide. Conidia 4.5–12.5(–13.5) × 3–4 µm, in simple and
branched acropetal chains, ellipsoid–ovoid, fusiform, subcylindrical, aseptate, subhyaline to pale brown,
thin-walled, smooth, with 1–3 hila, 1–1.5(–2) µm diam, truncate, unthickened or almost so, and somewhat
darkened-refractive, secession schizolytic; ramoconidia 8–13.5(–15.5) × 3–5 µm.
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Culture characteristics: Colonies on PDA reaching a diameter of 34–36 mm after 2 wk at 25 °C,
moderate aerial mycelium, powdery because of macroconidia, margins fringed; surface smoke grey to
grey olivaceous, margins olivaceous; reverse olivaceous grey. Colonies on OA reaching a diameter of
39–40 mm after 2 wk at 25 °C, moderate aerial mycel ium, powdery because of macroconidia, margins
fringed; surface smoke grey to grey olivaceous, margins grey olivaceous; reverse olivaceous grey.
Material examined: Thailand, Samut Songkhram Province, from Musa sp. (Musaceae), 8 Jun. 2008, P.W. Crous (holotype CBS
H-23396, culture ex-type CBS 143921 = CPC 33939).
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Notes: Metulocladosporiella samutensis can easily be distinguished from other species of
Metulocladosporiella by the production of conidiogenous cells directly from the apex, or loosely
arranged primary branches, being almost totally absent of secondary branches.
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Authors: Y. Marin-Felix, R. Cheewangkoon, J. Luangsa-ard & P.W. Crous
Microdochium Syd. & P. Syd., Ann. Mycol. 22: 267. 1924. Fig 36.
Synonyms: Monographella Petr., Ann. Mycol. 22: 144. 1924.
Griphosphaerella Petr., Ann. Mycol. 25: 209. 1927.
Gloeocercospora D.C. Bain & Edgerton, Phytopathology 33: 225. 1943. (nom. inval., Art. 39.1,
Melbourne).
Gloeocercospora D.C. Bain & Edgerton ex Deighton, Trans. Brit. Mycol. Soc. 57: 358. 1971.
Gerlachia W. Gams & E. Müll., Neth. J. Pl. Path. 86: 49. 1980.
Classification: Sordariomycetes, Xylariomycetidae, Xylariales, Microdochiaceae.
Type species: Microdochium phragmitis Syd. Holotype: K-IMI 193888. Epitype and ex-epitype strain
designated by Hernández-Restrepo et al. (2016a): CBS H-22135, CBS 285.71.
DNA barcode (genus): LSU.
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DNA barcodes (species): ITS, rpb2, tub2. Table 11. Fig. 37.
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Ascomata perithecial, immersed, subepidermal, solitary or in groups, pale brown to black, globose,
subglobose to oval; ostiole central, neck papillate and often acute, usually more distinctly pigmented than
ascomatal body, filled with slightly clavate periphyses; ascomatal wall brown, thin-walled, thickened and
darker around ostiole, in face view textura angularis-epidermoidea. Hamathecium comprising septate,
filamentous, apically free, thin-walled paraphyses. Asci unitunicate, oblong to clavate with 8 bi- to
multiseriate ascospores, apex with an amyloid, refractive, flat, funnel-shaped ring. Ascospores clavate,
fusoid or oblong, hyaline to brownish, straight or curved, smooth, septate. Conidiomata absent or
present, sporodochial, epidermal or subepidermal, erumpent through stomata, or rupture of outer
epidermal wall and cuticle, or by specialised egression hyphae through outer epidermal wall, hyaline,
pseudoparenchymatic, spreading after egress. Conidiophores more or less verticillate, often slightly
differentiated, reduced to conidiogenous cells, hyaline, smooth. Conidiogenous cells holoblastic,
discrete, hyaline, smooth, solitary or aggregated in small sporodochia. Two kinds: with sympodial
proliferation, cylindrical or slightly tapering to clavate, denticulate with one or more apical denticles; or
with percurrent proliferation (annellidic), subcylindrical, obpyriform, ampulliform to lageniform. Conidia
dry or in slimy mass, unicellular or multiseptate, hyaline, smooth, lunate, falcate, fusiform, filiform,
obovoid or subpyriform, straight or curved, apex rounded, base flattened. Sometimes conidia originate
directly from hyphae. Chlamydospores terminal or intercalary, solitary, in chains or grouped in clusters,
brown (adapted from Hernández-Restrepo et al. 2016a).
Culture characteristics: Colonies on OA saffron, salmon, peach or white when young, some species grey
or dark grey when mature, glabrous or with moderate amount of mycelium, cottony to floccose, margin
effuse.
Optimal media and cultivation conditions: OA at 25 °C under dark conditions.
Distribution: Worldwide.
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Hosts: Mainly pathogens of grasses and cereals, but some also occur on non-grass hosts as Opuntia
(Cactaceae) and Lycopodium (Lycopodiaceae), may cause losses to crops including rice, maize, wheat,
barley and sorghum. Other species can be found in harvested grains.
Disease symptoms: Microdochium patch or pink snow patch, leaf scald disease, tar spot disease, root
necrosis and decay of grasses, leaf spots, among others.
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Notes: Microdochium includes plant pathogenic as well as saprobic and soil fungi (Sydow 1924, de
Hoog & Hermanides-Nijhof 1977, Parkinson et al. 1981, Jaklitsch & Voglmayr 2012, Zhang et al. 2015,
Hernández-Restrepo et al. 2016a, Crous et al. 2018). For many years, species of Microdochium were
recognised as fusarium-like fungi; however, morphological and molecular data separate these genera.
Conidiogenesis in Microdochium is not phialidic as in true Fusarium species and the conidia have
truncate basal cells rather than “foot-cells”. The sexual morphs of Microdochium are monographellalike, and it belongs in the Microdochiaceae (Xylariales) phylogenetically distant from true Fusarium in
Nectriaceae (Hypocreales).
For an accurate species identification of Microdochium species, DNA sequence analyses are
required. Among the four loci studied (i.e. LSU, ITS, rpb2 and tub2), LSU is useful only for generic
placement. Phylogenies based on individual gene regions of ITS, rpb2 and tub2, can be used to
distinguish 14 species in Microdochium; those phylogenies generated from tub2 show longer
distances between species and higher support values. This is more informative than ITS and rpb2
(Hernández-Restrepo et al. 2016a).
References: von Arx 1981, 1984, Braun 1995 (taxonomy); Parkinson et al. 1981, Müller & Samuels
1984, Zhang et al. 2015 (morphology and pathogenicity); Hong et al. 2008 (pathogenicity); HernándezRestrepo et al. 2016a (morphology and phylogeny).
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Microdochium novae-zelandiae Hern.-Restr., Thangavel & Crous, sp. nov. MycoBank MB824606. Fig.
38.
Etymology: Name is derived from New Zealand, the country where this fungus was collected.
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Mycelium superficial and immersed, composed of septate, branched, hyaline, smooth, 1–2.5 wide
hyphae. Conidiomata sporodochium-like, formed in aerial mycelium or on agar surface, hyaline to pink.
Conidiophores often reduced to conidiogenous cells. Conidiogenous cells 4–10 × 2–3 µm, integrated,
terminal, polyblastic, proliferation sympodial, cylindrical to lageniform, hyaline, smooth; sometimes conidia
formed directly on mycelium. Conidia 5.5–10 × 2–2.5 µm, solitary, fusoid, allantoid, lunate or slightly
sigmoid, straight or curved, hyaline, smooth, 0(–1)-septate, base truncate. Chlamydospores not
observed.
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Culture characteristics: Colonies on OA reaching 45 mm diam after 1 wk at 25 °C, centre flat and rosy
buff, periphery cottony and white, margins effuse; reverse rosy buff in centre.
Materials examined: New Zealand, Christchurch, from turf leaves (Poaceae), 2015, R. Thangavel (holotype CBS H-23384, culture
ex-type CBS 143847 = CPC 29376 = ICMP 21872 = MPI T15_05208H); ibid., Richmond, Nelson, on spruce (Pinaceae), 2014, R.
Thangavel (CPC 29693 = MPI T14_00277D).
Notes: Microdochium novae-zelandiae is known from two isolates, both of which were collected in New
Zealand from different hosts belonging to the families Pinaceae and Poaceae. Based on a four-gene
analysis, the new species was placed in a clade distinct from M. bolleyi, M. colombiense, M. majus and
M. nivale (Fig. 37). Morphologically, M. novae-zelandiae has conidia similar in size to M. bolleyi and M.
colombiense. However, subtle morphological differences exist in the conidial shapes of these taxa with M.
novae-zelandiae having sigmoidal conidia. Compared with M. phragmitis, conidia of M. novae-zelandiae
are smaller, mainly aseptate with variable shape [5.5–10 × 2–2.5 µm, 0(–1)-septate, fusiform, allantoid,
lunate or slightly sigmoid in M. novae-zelandiae vs. 10–14.5 × 2–3 µm, 0–1-sepate, fusiform to navicular
in M. phragmitis].
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Authors: M. Hernández-Restrepo, R. Thangavel & P.W. Crous
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Oculimacula Crous & W. Gams, Eur. J. Pl. Path. 109: 845. 2003. Fig. 39.
Synonym: Helgardia Crous & W. Gams, Eur. J. Pl. Path. 109: 845. 2003.
Classification: Leotiomycetes, Leotiomycetidae, Helotiales, Incertae sedis.
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Type species: Oculimacula yallundae (Wallwork & Spooner) Crous & W. Gams = Helgardia
herpotrichoides (Fron) Crous & W. Gams, basionym Cercosporella herpotrichoides Fron. Holotype: K(M)
233697. Neotype and ex-neotype strain of Helgardia herpotrichoides designated by Crous et al. (2003):
CBS H-23003, CBS 110665.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, tef1. Table 12. Fig. 40.
Ascomata 0.5–2.5 mm diam, apothecial, sessile, gregarious, circular to lobate, on a subiculum of white to
dark brown hyphae, attached to substrate via a superficial mat of pale brown, thin hyphae. Disk smooth,
grey with a pale grey margin, becoming emarginate and flattened to convex at maturity. Receptacle pale
brown to grey-brown, cup-shaped. Medullary excipulum of multiseptate, hyaline hyphae. Ectal excipulum
of thin-walled, dark brown, angular cells, becoming more elongated towards margin. Paraphyses filiform
with obtuse ends, similar in length to asci. Asci 8-spored, unitunicate, clavate to subcylindrical or fusoid,
with a short stalk, and an apical pore staining blue in Melzer’s reagent. Ascospores bi- to multiseriate,
hyaline, smooth, aseptate, fusoid to subcylindrical or clavate with rounded ends, mostly straight.
Conidiophores fasciculate or solitary on superficial mycelium, or arising from pale brown stromata,
subcylindrical to geniculate-sinuous, rarely branching, hyaline to pale olivaceous, smooth, consisting of
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conidiogenous cells only, or slightly differentiated with up to 2 septa. Conidiogenous cells integrated,
proliferating sympodially at apex, with inconspicuous, dense geniculations; loci unthickened,
inconspicuous, not darkened. Conidia solitary, hyaline, smooth, arranged in slimy packets, acicular,
filiform, straight to curved, one- to multiseptate, forming smaller, secondary conidia via microcyclic
conidiation (adapted from Crous et al. 2003).
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Culture characteristics: Colonies with moderate aerial mycelium giving a cottony appearance. On PDA
surface grey to olive grey, brownish-grey, pinkish-grey or greenish; reverse grey to greenish-black,
greenish, brownish or creamy pink.
Optimal media and cultivation conditions: SNA under continuous near-ultraviolet light at 25 °C.
Distribution: Africa, Australasia, Europe, New Zealand and North America.
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Hosts: Poaceae, including Triticum and Hordeum, among others.
Disease symptoms: Eyespot lesions that girdle the stem and soften the stem-base.
Notes: Oculimacula encompasses four species associated with eyespot disease symptoms of cereals in
the temperate regions of the world. Eyespot is an important disease of stem bases in which the infection
occurs at or near the soil line, attacking chiefly the basal leaf sheaths and internodal tissues of the culms
(Sprague & Fellows 1934, Lucas et al. 2000). The fungus sporulates in the fall and spring producing the
disease, and survives the winter on diseased stubble standing or lying in the field (Sprague & Fellows
1934). Disease control relies on the use of fungicides, delayed seeding in the fall and by planting resistant
cultivars (Murray 1996, Douhan et al. 2002).
Oculimacula was introduced by Crous et al. (2003) to accommodate sexual morphs previously
classified in Tapesia, while Helgardia was introduced for the asexual morphs linked to Oculimacula.
Johnston et al. (2014) synonymised these generic names and conserved the name Oculimacula because
it is most commonly used by plant pathologists for the eyespot diseases of wheat and barley.
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References: Sprague & Fellows 1934 (morphology and pathogenicity); Sprague 1936, Lucas et al.
2000 (pathogenicity); Nirenberg 1981 (morphology and pathogenicity); Crous et al. 2003 (morphology
and phylogeny).
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Oculimacula acuformis (Nirenberg) Y. Marín & Crous, comb. et stat. nov. MycoBank MB824638.
Basionym: Pseudocercosporella herpotrichoides var. acuformis Nirenberg, Z. PflKrankh. PflSchutz 88:
244. 1981.
Synonyms: Ramulispora herpotrichoides var. acuformis (Nirenberg) Boerema, et al., Netherlands Journal
of Plant Pathology, Supplement 1 98: 22. 1992.
Tapesia yallundae var. acuformis Boerema et al., Netherlands J. of Pl. Path., Supplement 1 98: 22. 1992.
(nom. inval., Art. 40.3, Melbourne).
Ramulispora acuformis (Nirenberg) Crous, S. Afr. J. Bot. 61: 46. 1995.
Tapesia acuformis (Boerema et al.) Crous, S. Afr. J. Bot. 61: 46. 1995. (nom. inval., Art. 40.3, Melbourne).
Helgardia acuformis (Nirenberg) Crous & W. Gams, Eur. J. Pl. Path. 109: 846. 2003.
Oculimacula acuformis (Boerema et al.) Crous & W. Gams, Eur. J. Pl. Path. 109: 846. 2003. (nom. inval.,
Art. 40.3, Melbourne).
Material examined: Germany, Tübingen, from Secale cereale (Poaceae) culm base, 1978, H. Nirenberg (culture ex-type CBS
495.80).
Notes: Oculimacula acuformis was introduced to accommodate Tapesia yallundae var. acuformis (Crous
et al. 2003). However, the combination is invalid because the basionym lacks details for the ex-type strain
(Art 40.3). We have consequently proposed the new combination based on its asexual morph
Pseudocercosporella herpotrichoides var. acuformis. This latter species was transferred to Helgardia in
the same publication where Oculimacula acuformis was proposed and when both genera were first
introduced (Crous et al. 2003). Helgardia acuformis is a synonym of Oculimacula acuformis.
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Authors: Y. Marin-Felix, J.Z. Groenewald & P.W. Crous
Paraphoma Morgan-Jones & J.F. White, Mycotaxon 18: 58. 1983. Fig. 41.
Synonym: Phoma section Paraphoma (Morgan-Jones & J.F. White) Boerema, Stud. Mycol. 32: 7. 1990.
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Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
Type species: Paraphoma radicina (McAlpine) Morgan-Jones & J.F. White, basionym: Pyrenochaeta
radicina McAlpine. Holotype: in VPRI (Australia, Shepparton, Victoria, on roots of Prunus cerasus
(Rosaceae), 21 Oct 1901, Piscott, 2064.3). Epitype and ex-epitype strain designated by de Gruyter et al.
(2010): CBS H-16560, CBS 111.79.
DNA barcodes (genus): LSU, SSU.
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DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 13. Fig. 42.
Conidiomata pycnidial, globose to subglobose, papillate, thick-walled, pseudoparenchymatous, ostiolate,
uniloculate; conidiomatal matrix white or buff, cream, yellow, brown or hyaline; setae abundant, straight or
flexuous, septate, pale brown to brown, short or relatively long, stiff or hyphal-like, scattered on surface of
conidiomata, or abundant around ostioles. Micropycnidia fertile or sterile, produced abundantly in some
species of Paraphoma, submerged in medium. Conidiophores ampulliform, hyaline, mostly reduced to
phialidic conidiogenous cells. Conidiogenous cells lageniform, monophialidic, hyaline to subhyaline. Conidia
ellipsoidal to subglobose, hyaline, guttulate, aseptate in vivo and in vitro. Chlamydospores absent or
present, solitary, in short or long chains or aggregated, uni- or multicellular; multicellular chlamydospores
alternarioid, pseudosclerotioid, epicoccoid and botryoid depending on species. Sexual morph unknown.
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Culture characteristics: Colony colour, growth and pigmentation greatly dependant on media and
incubation conditions. Colonies black, brown, olivaceous, yellow, red to pink, or grey and white; slow
growing; aerial mycelium flat to effuse, aerial mycelium sparsely formed, floccose to tufted, felty, woolly or
compact; margins regular, smooth and sharp, or irregular, crenate and lobate.
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Optimal media and cultivation conditions: CHA for colony growth and pigmentation, MEA mostly for colony
pigmentation and acidified OA for both colony pigmentation and morphological identification, incubated for 1
wk in dark and 1 wk under near-ultraviolet light (13 h light, 11 h dark) at 20–22 °C to simulate colon y
pigmentation and sporulation.
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Distribution: Temperate areas of Australia, Eurasia and North America.
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Hosts: Mostly foliar pathogens of herbaceous plants, chiefly soil-borne, with wide host range including
monocotyledonous plants, Asteraceae, Cupressaceae, Rosaceae and Solanaceae, occasionally
saprobic.
Disease symptoms: Crown discolouration, root rot and necrotic leaf spots.
Notes: The type species of Paraphoma, Pa. radicina, clustered in a separate group outside
Didymellaceae and hence was excluded from Phoma (de Gruyter et al. 2013). In a phylogenetic analysis
based on LSU and SSU, Paraphoma radicina clustered in the Phaeosphaeriaceae, although other
species belonged to the Cucurbitariaceae and Coniothyriaceae. Setose pycnidial conidiomata and
dictyochlamydospores, which are characteristics of species of Paraphoma and Peyronellaea, can be
observed in species of other phoma-like genera, such as Pyrenochaeta and Pleurophoma. Therefore,
these morphological characters are not specific to these genera. In order to delineate Paraphoma,
phylogenetic studies based on ITS, LSU, rpb2, tef1 and tub2 have been performed (Aveskamp et al.
2010, Moslemi et al. 2016, 2018, Crous et al. 2017a). Using ITS and LSU in combination with protein
coding genes rpb2, tef1 and tub2 for precise identification of species of Paraphoma is necessary, as LSU
alone is too conservative.
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References: de Gruyter & Boerema 2002, Zhang et al. 2009, 2012, de Gruyter et al. 2010, 2013
(pathogenicity, phylogeny and distribution); Boerema et al. 2004 (morphology, pathogenicity, media and
incubation conditions); Aveskamp et al. 2009, 2010 (morphology, phylogeny and key of all Paraphoma
spp.); Hay et al. 2015 (hosts).
Phaeoacremonium W. Gams et al., Mycologia 88: 789. 1996. Fig 43.
Synonym: Togninia Berl., Icon. fung. (Abellini) 3: 9. 1900.
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Authors: A. Moslemi, P.W.J. Taylor & P.W. Crous
Classification: Sordariomycetes, Sordariomycetidae, Togniniales, Togniniaceae.
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Type species: Phaeoacremonium parasiticum (Ajello et al.) W. Gams et al., basionym: Phialophora
parasitica Ajello, et al. Holotype and ex-type strain: CBS H-17463, CBS 860.73.
DNA barcodes (genus): SSU, LSU.
DNA barcodes (species): act, tub2. Table 14. Fig. 44.
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Ascomata perithecial, aggregated or solitary, superficial to immersed, non-stromatic, globose to
subglobose, dark, opaque, long-necked; necks straight or flexuous; ascomatal wall fragile to leathery,
comprising two layers of textura angularis: outer layer brown to dark brown, with cells smaller and
more rounded than those of inner layer; inner layer hyaline to pale brown, cells flattened. Paraphyses
abundant, broadly cellular, slightly constricted at septa, branching, hyaline, slightly tapering apically or
thread-like towards apex. Ascogenous hyphae hyaline, sometimes branched in basal region,
elongating during ascal formation with remnant tissue from which single asci arise. Asci arising in
acropetal succession, appearing spicate when mature, unitunicate, 8-spored, ascal apex thickened
with a nonamyloid apical ring, basally bluntly obtuse, sessile. Ascospores hyaline, aseptate, allantoid,
reniform, cylindrical or oblong-ellipsoidal, mostly biseriate or in a single row. Conidiophores branched
in basal region or unbranched, arising from aerial or submerged hyphae, erect, nearly cylindrical when
unbranched, slightly tapering, straight or flexuous, variable in length, up to 7-septate, mostly pale brown,
paler towards tip, percurrent rejuvenation observed, small warts or verruculose ornamentation mostly at
base, usually with one integrated terminal phialide and one or two additional, discrete phialides at
uppermost septum. Conidiogenous cells phialidic, discrete or integrated, terminal or lateral, mostly
monophialidic, sometimes polyphialidic, sparsely warted, verruculose or smooth, pale brown to hyaline,
with an inconspicuous funnel-shaped collarette. Three distinct classes of phialides (Types I–III) can be
observed. Conidia aggregated into round, slimy heads at apices of phialides, hyaline, aseptate, smoothwalled, oblong-ellipsoidal to obovate, cylindrical, allantoid or reniform, uncommonly fusiform-ellipsoidal or
globose, becoming biguttulate with age.
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Culture characteristics: Colonies on MEA flat with entire margins, mostly moderately dense,
predominantly felty, and sometimes woolly; brown, olive-grey, pale yellow to beige or pink to dark pink.
Optimal media and cultivation conditions: 2 % MEA to induce sporulation of asexual morph. Cultural
characters that are useful to distinguish Phaeoacremonium species include colour of colonies on MEA,
and yellow pigment production on PDA and OA. For the sexual morph 2 % WA is used with twiceautoclaved pieces of 3–4 cm of grapevine cane at 22 °C (GWA).
Distribution: Worldwide.
Hosts: Frequently isolated from both diseased woody plants with brown wood streaking, and humans
with phaeohyphomycotic infections. Other hosts include larvae of bark beetles, arthropods, and soil.
Because of the involvement of members of this genus in Petri disease and esca of grapevines (Vitis
spp.), isolates from this host have been intensively studied (Mostert et al. 2006, Gramaje et al. 2015,
Spies et al. 2018). Even though Phaeoacremonium species can infect a wide range of woody hosts
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(more than 40 host plants), recent publications have shown the importance of Phaeoacremonium
species in causing brown wood streaking of Olea europaea (Oleaceae) and Prunus spp (Rosaceae)
(Damm et al. 2008, Carlucci et al. 2015).
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Notes: Species delimitation based on morphology alone has little value since many species have
overlapping characters. Moreover, the morphology of the sexual morph cannot be used because only
15 taxa are known. The two gene regions used most frequently for phylogenetic analyses are actin
(act) and partial beta-tubulin (tub2) genes (Mostert et al. 2006). Phylogenetic analyses combining
these two regions allow for the resolution of almost all currently known Phaeoacremonium species
with good support (≥ 0.97 PP, ≥ 96% BS) (Fig. 44). The three exceptions to this are Pha. griseorubrum
(paraphyletic), Pha. roseum (0.72 PP, 100 % BS) and Pha. viticola (0.87 PP, 62 % BS) (Fig. 44, also
see Gramaje et al. 2015 and Spies et al. 2018). Other gene regions that have been used include the
ITS, tef1 and cal (Groenewald et al. 2001, Mostert et al. 2005, Úrbez-Torres et al. 2014). Úrbez-Torres
et al. (2014) included ITS and tef1 data along with act and tub in their phylogeny, which resolved all
included species with more than 97 % or 96 % bootstrap support in maximum parsimony and
neighbour joining analyses respectively. The ITS region is considered insufficiently variable to
distinguish between several of the species and is not recommended as a barcode (Mostert et al.
2005); however, considering the resolution and support in the phylogeny of Úrbez-Torres et al. (2014),
the tef1 region is valuable in resolving issues with support and resolution in the act-tub2 phylogeny.
The cal region was sequenced for a limited number of species by Mostert et al. (2005) to resolve taxa
related to Pha. rubrigenum. Unfortunately, sequence data for this region are available for a limited
number of species and its usefulness in distinguishing between Phaeoacremonium species remains
uncertain.
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References: Crous et al. 1996 (taxonomy); Eskalen et al. 2005, Rooney-Latham et al. 2005 (sexual
morph); Mostert et al. 2006, Gramaje et al. 2015 (taxonomy, distribution, host range, detection,
identification, pathogenesis and epidemiology); Aroca & Raposo 2007, Pouzoulet et al. 2013, ÚrbezTorres et al. 2015 (detection and identification); Halleen et al. 2007, Damm et al. 2008; Aroca &
Raposo 2009, Gramaje et al. 2010 (pathogenicity); Blanco-Ulate et al. 2013 (genome sequence);
Moyo et al. 2014, Agustí-Brisach et al. 2015 (epidemiology); Réblová et al. 2015 (systematics).
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Phaeoacremonium pravum C.F.J. Spies, L. Mostert & Halleen, sp. nov. MycoBank MB821019. Fig.
45.
Etymology: Latin, pravum meaning crooked, in reference to the crooked shape of some phialides.
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Mycelium of branched, prominently septate, hyaline to pale brown, smooth to finely verruculose (1–
)1.5–2.5 (av. 2) µm diam hyphae, forming bundles of up to 5 strands, individual strands in bundles
often forming direct hyphal connections. Conidiophores (14.5–)16–61(–77) × 1.5–2.5 (av. 28.5 × 2)
µm, smooth to finely verruculose, usually branched, hyaline, up to 9 septa. Phialides terminal or
lateral, monophialidic, sometimes proliferating vegetatively behind collarette, types I and II dominant,
collarettes funnel-shaped, 0.5–1.5 × 0.5–2 (av. 1 × 1.5) µm, smooth, hyaline; type I mainly
subcylindrical, sometimes elongate ampulliform, (2–)2.5–10.5(–11) × 1–2 (av. 6 × 1.5) µm; type II
subcylindrical with tapering apex to elongate ampulliform, sometimes curved or bent especially at
apex, (8–)8.5–14(–14.5) × 1.5–2(–2.5) (av. 11.5 × 2) µm; type III subcylindrical with tapering apex to
subulate, sometimes slender navicular, (14–)14.5–26.5(–31.5) × 1.5–2 (av. 19 × 1.5) µm. Conidia 3–
4(–4.5) × 1.5(–2) (av. 3.5 × 1.5) µm, borne in slimy heads, oblong-ovoid to ellipsoidal to allantoid.
Culture characteristics: Colonies reaching a radius of 8–10 mm after 8 d at 25 °C. Minimum
temperature for growth 10 °C, optimum 20 °C, maximu m 35 °C. Colonies on MEA smooth, submerged
with entire edge, after 16 d white to pale buff above and in reverse. Colonies on PDA smooth,
submerged, with central folds, with entire margin, after 16 d white to pale buff above and in reverse.
Colonies on OA felty, folded, with submerged margins, with entire edge, after 16 d white to pale smoke
grey with darker margins.
Materials examined: South Africa, from wood of Vitis berlandieri × rupestris (rootstock cv. Richter 110) (Vitaceae), 18 Sep.
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2014, A. Vermeulen (holotype CBS-H 23158, culture ex-type CBS 142686 = STE-U 8363 = CSN3); ibid., on Vitis vinifera cv.
Early Sweet cordon (Vitaceae), 18 Sep. 2014, A. Vermeulen, CBS 142687 = STE-U 8364 = CSN11.
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Notes: There are several differences between the ex-type strain (CBS 142686) and strain CBS
142687. Strain CBS 142687 had a higher optimum and maximum temperatures for growth (25 °C and
37 °C, respectively) than strain CBS 142686 and rea ched a radius of 11–12 mm after 8 d at 25 °C.
After 16 d, colonies of strain CBS 142687 also had pronounced pigmentation on MEA i.e. rosy
vinaceous with dark purple patches with central white tufts of aerial mycelium, and on PDA i.e. livid red
to dark vinaceous with white to smoke grey woolly aerial mycelium, and on OA i.e. mouse grey to
olivaceous grey with white margins. The act sequence of strain CBS 142687 differs from that of the
ex-type (CBS 142686) at six positions over a length of 210 bases, resulting in paraphyly of this
species in an act-only phylogeny (Spies et al. 2018). Considering the high similarity of tub2 sequences
(598/599 identical bases), strong support for the monophyly of Pha. pravum in the combined act-tub2
phylogeny, and the fact that both strains produced curved phialides, CBS 142687 is regarded as Pha.
pravum until additional strains and data become available to indicate differently.
Authors: D. Gramaje, L. Mostert, C.F.J. Spies & F. Halleen
Phyllosticta Pers., Traité sur les Champignons Comestibles (Paris): 55. 147. 1818. Fig. 46.
Synonym: Guignardia Viala & Ravaz, Bull. Soc. mycol. Fr. 8: 63. 1892.
Classification: Dothideomycetes, Dothideomycetidae, Botryosphaeriales, Phyllostictaceae.
Type species: Phyllosticta convallariae Pers., nom. inval. (= Phyllosticta cruenta (Fr.) J. Kickx f.).
Reference strain: CBS 858.71.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, act, gapdh, tef1. Table 15. Fig. 47.
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Ascomata pseudothecial, separate to gregarious, globose to subglobose, brown to black, unilocular
with a central ostiole. Pseudoparaphyses mostly absent at maturity, filamentous, branched, septate
when present. Asci bitunicate, fissitunicate, clavate to subcylindrical, 8-spored, fasciculate, stipitate,
with an ocular chamber. Ascospores bi- to triseriate, hyaline, guttulate to granular, aseptate, ellipsoid,
ellipsoid-fusoid to limoniform, smooth-walled, usually with mucilaginous caps at ends, or surrounded
by a mucilaginous sheath. Conidiomata and spermatogonia pycnidial, immersed, subepidermal to
erumpent, unilocular, rarely multilocular, glabrous, ostiolate, dark brown to black; ostiole circular to
oval; conidiomatal wall thick-walled, dark brown, textura angularis, with inner layers of hyaline to pale
brown, thin-walled, textura prismatica to angularis. Conidiophores lining cavity of conidioma, reduced
to conidiogenous cells, invested in mucus. Conidiogenous cells discrete, producing macroconidia and
spermatia, also produced in separate spermatogonia, ampulliform, lageniform, doliiform to
subcylindrical, hyaline, smooth, proliferating percurrently near apex, invested in a mucoid layer.
Conidia ellipsoid-fusoid to obovoid or ovoid, rarely subcylindrical, aseptate, broadly rounded at apex,
often tapering strongly toward base, unicellular, hyaline, smooth-walled, guttulate to granular, often
enclosed in a persistent mucilaginous sheath, and bearing an unbranched, tapering, straight to curved,
mucoid apical appendage. Spermatogenous cells ampulliform to lageniform or subcylindrical, hyaline,
smooth, phialidic. Spermatia hyaline, smooth, granular, subcylindrical or dumbbell-shaped, with
rounded or blunt ends (adapted from Wikee et al. 2013b).
Culture characteristics: Colonies on MEA, OA and PDA after 2 wk in dark at 27 °C erumpent or flat,
spreading with sparse or moderate aerial mycelium; on MEA, OA and PDA surface frequently iron-grey
or olivaceous grey, less frequently greenish to dark green; reverse iron-grey, olivaceous grey or black.
Optimal media and cultivation conditions: PNA, OA, PDA and SNA under near-ultraviolet light at 27 °C to
induce sporulation.
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Distribution: Worldwide.
Hosts: Wide range of hosts from trees to ornamentals.
Disease symptoms: Leaf spots and various fruit diseases.
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Notes: Phyllosticta was introduced by Persoon (1818), with Phy. convallariae designated as type
species (Donk 1968). However, this species was invalid because it lacked a description. Therefore,
Phy. cruenta, which is a synonym of Phy. convallariae, was designated as type of the genus (van der
Aa & Vanev 2002). There is no available type material for this species, which was described from
Polygonatum multiflorum collected in Germany. A strain deposited in CBS previously identified as
Guignardia reticulata, which is the sexual morph of Phy. cruenta, was isolated from Polygonatum
odoratum in the Czech Republic, being a potential neotype for Phy. cruenta. However, this strain is
sterile and we have chosen to consider it as a reference strain since we could not confirm its
identification based on morphology.
Phyllosticta includes plant pathogenic species that cause diseases of significant economic
importance. For example, Phy. citricarpa is the responsible for citrus black spot, which is considered a
quarantine pest in Europe and the USA (Baayen et al. 2002, Glienke et al. 2011, Guarnaccia et al.
2017). Other examples include the Phy. ampelicida species complex that causes black rot disease on
grapevines (Wicht et al. 2012, Carstens et al. 2017), and the Phy. musarum species complex that is
responsible for banana freckle disease (Pu et al. 2008, Wong et al. 2012).
Phoma and Phyllosticta have been difficult to distinguish since both genera were recognised as
pycnidial fungi producing unicellular, hyaline conidia. Subsequent molecular data enabled the
discrimination of both genera, as well as the fact that Phyllosticta was linked to its sexual morph,
Guignardia (Glienke et al. 2011, Wikee et al. 2011, 2013b, Wong et al. 2012, Zhou et al. 2015,
Guarnaccia et al. 2017).
Phyllosticta was formerly placed in the Botryosphaeriaceae, together with Botryosphaeria (Schoch
et al. 2006). However, Wikee et al. (2013b) showed that it represents a different phylogenetic lineage,
for which the family name Phyllostictaceae (Fries 1849) was resurrected.
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References: van der Aa 1973 (morphology and pathogenicity); van der Aa & Vanev 2002 (synonyms,
collection information and notes); Wikee et al. 2011 (pathogenicity and phylogeny); Glienke et al. 2011,
Wong et al. 2012, Wikee et al. 2013b, Zhou et al. 2015, Guarnaccia et al. 2017 (ecology, morphology
and phylogeny).
Phyllosticta iridigena Y. Marín & Crous, sp. nov. MycoBank MB823971. Fig. 48.
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Etymology: Name reflects the host it was isolated from, Iris.
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Conidiomata 90–200 µm diam, pycnidial, solitary, globose, dark brown, with central ostiole; conidiomatal
wall of 3–8 layers of brown textura angularis. Conidiophores lining cavity, reduced to conidiogenous cells.
Conidiogenous cells 4–7 × 4–6 µm, doliiform, hyaline, smooth, proliferating percurrently at apex. Conidia
(10–)12–13(–15) × (7–)8(–9) µm, solitary, ellipsoid to obovoid, aseptate, smooth, hyaline, guttulate,
granular; conidia encased in a mucoid sheath 2–3 µm diam, and a single apical mucoid appendage, 7–15
× 2 µm, tapering to acutely rounded apex.
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, feathery
margins, reaching 45 mm diam after 2 wk at 25 °C. On MEA surface pale olivaceous grey, reverse irongrey. On PDA surface and reverse olivaceous grey. On OA surface pale olivaceous grey with diffuse
yellow pigment in agar.
Material examined: South Africa, on Iris sp. (Iridaceae), 16 Jan. 2010, P.W. Crous (holotype CBS H-23385, culture ex-type CBS
143410 = CPC 32669).
Notes: This species clusters in a well-supported clade (95 % BS / 1 PP) with Phy. hypoglossi and Phy.
cussoniae. Phyllosticta hypoglossi produces longer conidiogenous cells (10–15 µm) and wider conidia
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[(9–)10(–11) µm] than Phy. iridigena. Moreover, these three species are isolated from different hosts, i.e.
Phy. hypoglossi from Ruscus (Ruscaceae), Phy. cussoniae on Cussonia (Araliaceae) and Phy. iridigena
on Iris (Iridaceae). Phyllosticta cussonia and Phy. iridigena have been found in the same country, South
Africa, while Phy. hypoglossi is an European species.
Phyllosticta persooniae Y. Marín & Crous, sp. nov. MycoBank MB823972. Fig. 49.
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Etymology: Name reflects the host genus Persoonia from which it was isolated.
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Conidiomata 200–300 µm diam, pycnidial, solitary, globose, dark brown, with central ostiole; conidiomatal
wall of 3–8 layers of brown textura angularis. Conidiophores 10–18 × 6–7 µm, lining cavity, 0–1-septate,
subcylindrical, hyaline, smooth, rarely branched. Conidiogenous cells 9–17 × 4–5 µm, terminal,
subcylindrical, hyaline, smooth, proliferating percurrently at apex. Conidia (9–)10–11(–12) × (7–)8(–9) µm,
solitary, ellipsoid to obovoid, aseptate, smooth, hyaline, guttulate, granular; conidia encased in a mucoid
sheath that is inconspicuous and dissolves at maturity, but with a single apical mucoid appendage, 7–15
× 2–3 µm, tapering to acutely rounded apex.
Culture characteristics: Colonies flat to erumpent, spreading, with sparse to moderate aerial mycelium
and feathery, lobate margins, reaching 30 mm diam after 2 wk at 25 °C. On MEA surface smoke grey,
reverse olivaceous grey. On PDA surface and reverse olivaceous grey. On OA surface pale mouse grey.
Material examined: Australia, New South Wales, South East Forests National Park, Nunnock Swamp, on Persoonia sp.
(Proteaceae), 28 Nov. 2016, P.W. Crous (holotype CBS H-23386, culture ex-type CBS 143409 = CPC 32603).
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Notes: Phyllosticta persooniae is phylogenetically distant from all other species of Phyllosticta, the most
closely related species being Phy. foliorum. Morphologically, these can be distinguished by the size of
their conidia. Phyllostica foliorum is characterised by its larger conidia i.e. (12–)13–14(–15) × (9–)10(–11)
m vs. (9–)10–11(–12) × (7–)8(–9) µm in Phy. persooniae. Moreover, Phy. foliorum has never been found
on Persoonia (Proteaceae) or in Australia, which is the host and distribution of Phy. persooniae (Farr &
Rossman 2017). Most species of Phyllosticta are host-specific.
Proxipyricularia Klaubauf et al., Stud. Mycol. 79: 109. 2014. Fig 50.
Classification: Sordariomycetes, Sordariomycetidae, Magnaporthales, Pyriculariaceae.
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Type species: Proxipyricularia zingiberis (Y. Nisik.) Klaubauf, et al., basionym: Pyricularia zingiberis Y.
Nishik. Lectotype designated here: plate 4, fig. 3–8 in Nishikado Y. 1917. Ber. Ohara Inst. Landwirt.
Forsch. 1: 222. Epitype and ex-epitype strain designated here: CBS H-23356, CBS 133594.
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DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, act, cal, rpb1. Table 16. Fig. 28.
Conidiophores solitary or in fascicles, subcylindrical, erect, olivaceous to medium brown, smooth,
septate. Conidiogenous cells terminal and intercalary, pale brown, with denticulate conidiogenous loci
and rhexolytic secession. Conidia solitary, formed sympodially, pyriform to obclavate, narrowed toward
apex, rounded at base, 2-septate, subhyaline to pale brown, with a distinct protruding basal hilum,
frequently with minute marginal frill (adapted from Klaubauf et al. 2014).
Culture characteristics: Colonies reaching 43–50 mm in 1 wk at 25 °C, with out or with moderate aerial
mycelium. On CMA surface and reverse transparent. On OA surface salmon to ochreous; reverse pale
luteous to luteous. On PDA surface olivaceous to grey olivaceous with margins transparent; reverse
olivaceous to grey olivaceous with margins buff.
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Optimal media and cultivation conditions: On CMA and OA at 25 °C in dark, or autoclaved bar ley seeds
placed on SNA at 25 °C under near-ultraviolet light (12 h light, 12 h dark).
Distribution: Japan.
Hosts: Zingiber mioga and Z. officinale (Zingiberaceae).
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Disease symptoms: Leaf spots.
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Notes: In a phylogenetic study of the genus Pyricularia based on LSU, ITS, act, cal and rpb1,
Pyricularia zingiberis clustered in an independent clade distant from the type species of Pyricularia
(Klaubauf et al. 2014). Therefore, the genus Proxipyricularia was introduced to accommodate this
species, which is pathogen of Zingiber in Japan. Morphologically, both genera are similar, being
characterised by medium brown conidiophores and a terminal and intercalary denticulate rachis, and
subhyaline, 2-septate, obclavate conidia (Klaubauf et al. 2014).
References: Nishikado 1917 (morphology and pathogenicity); Klaubauf et al. 2014 (morphology and
phylogeny).
Proxipyricularia zingiberis (Y. Nisik.) Klaubauf et al., Stud. Mycol. 79: 109. 2014. Fig. 50.
Basionym: Pyricularia zingiberis Y. Nishik. (as “Piricularia zingiberi”), Ber. Ohara Inst. Landwirt.
Forsch. 1: 216. 1917.
Description: Klaubauf et al. (2014).
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Culture characteristics: Colonies on CMA reaching 43–50 mm after 1 wk at 25 °C, without aerial
mycelium; surface and reverse transparent. On OA reaching 48–50 mm after 1 wk at 25 °C, with
moderate aerial mycelium appearing slightly cottony, margins arachnoid; surface salmon to ochreous;
reverse pale luteous to luteous. On PDA reaching 47–48 mm after 1 wk at 25 °C, with sparse aerial
mycelium, margins fringed; surface olivaceous to grey olivaceous with margins transparent; reverse
olivaceous to grey olivaceous with margins buff.
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Materials examined: Japan, on leaves of Zingiber officinale (Zingiberaceae) (lectotype of Pyricularia zingiberis designated
here, MBT379808, plate 4, fig. 3–8 in Nishikado Y. 1917. Ber. Ohara Inst. Landwirt. Forsch. 1: 222). Japan, Hyogo, on Zingiber
mioga (Zingiberaceae), 2002, H. Kato [epitype of Pyricularia zingiberis designated here CBS H-23356, MBT379809, culture
ex-epitype CBS 133594 = MAFF 240222 = HYZiM201-0-1(Z-2J)].
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Notes: Type material was not designated when Py. zingiberis was introduced (Nishikado 1917).
Therefore, we selected the drawings of Nishikado in the original description as lectotype (Ber. Ohara
Inst. Landwirt. Forsch. 1: 222, plate 4, fig. 3–8). To fix the application of the generic name, an epitype
for this species is designated here from the same country (Japan) and host (Zingiber) as that of the
original specimen.
Pyriculariomyces Y. Marín, M.J. Wingf. & Crous, gen. nov. MycoBank MB823760. Fig. 51. Table 17.
Etymology: Named after the genus Pyricularia, which it resembles morphologically.
Ascomata separate, immersed, globose, brown, with central papillate neck and ostiole; ascomatal wall of
2–4 layers of brown textura angularis. Hamathecium dissolving upon maturity, with some cells remaining
among asci. Asci unitunicate, hyaline, smooth, 8-spored, subcylindrical, stipitate, apical mechanism
refractive, but not staining in Meltzer’s. Ascospores biseriate, fusoid-ellipsoid, widest in middle, tapering
towards subobtusely rounded ends, slightly curved to straight, 3-septate, pale brown, guttulate.
Conidiophores solitary, erect, straight to flexuous, unbranched, subcylindrical, brown, smooth, 1–8septate. Conidiogenous cells integrated, terminal, apex somewhat swollen with numerous denticle-like
loci, slightly thickened and darkened. Conidia solitary, pyriform, brown, finely verruculose, guttulate,
granular, apex subobtusely rounded, with or without mucoid cap, base truncate, hilum darkened,
thickened, 2-septate.
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Culture characteristics: Colonies reaching 90 mm after 2 wk at 25 °C, with m oderate aerial mycelium and
smooth, even margins. On MEA surface pale mouse grey with patches of dirty white, reverse isabelline
with patches of pale luteous. On OA surface honey with patches of pale mouse grey. On PDA surface
honey, reverse isabelline to honey.
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Type species: Pyriculariomyces asari (Crous & M.J. Wingf.) Y. Marín, M.J. Wingf. & Crous. Holotype and
ex-type cultures: CBS H-22625, CBS 141328 = CPC 27444.
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Notes: Pyriculariomyces is phylogenetically closely related to Pyricularia. However, Pyriculariomyces can
easily be distinguished by production of integrated terminal conidiogenous cells, while Pyricularia
produces terminal and intercalary conidiogenous cells. In Pyriculariaceae, the only genera characterised
by only terminal conidiogenous cells are Barretomyces and Utrechtiana. However, these genera can
easily be distinguished from Pyriculariomyces by the septation of the conidia i.e. 4(–5)-septate in
Barretomyces and 1-septate in Utrechtiana. Moreover, Utrechtiana differs in the presence of
conidiogenous cells that proliferate percurrently. Moreover, Pyriculariomyces can be distinguished from
Pyricularia by the production of ascomata with papillate necks with ostioles, while Pyricularia is
characterised by ascomata with long necks.
Pyriculariomyces asari (Crous & M.J. Wingf.) Y. Marín, M.J. Wingf. & Crous, comb. nov. MycoBank
MB823761. Fig. 51.
Basionym: Proxipyricularia asari Crous & M.J. Wingf., Persoonia 36: 393. 2016.
Description: Crous et al. (2016b).
Materials examined: Malaysia, Sabah, on leaves and stems of Asarum sp. (Aristolochiaceae), May 2015, M.J. Wingfield
(holotype CBS H-22625, culture ex-type CPC 27444 = CBS 141328); ibid., CPC 27442.
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Notes: Pyriculariomyces asari was introduced as a species of Proxypiricularia to accommodate two
specimens collected from Asarum (Crous et al. 2016b). However, the authors at the time suggested that
this species could represent another genus in the Pyricularia complex. The phylogenetic analysis
generated here based on four different loci (Fig. 28), support this hypothesis.
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Authors: Y. Marin-Felix, M.J. Wingfield & P.W. Crous
Pyricularia Sacc. Michelia 2: 20. 1880. Fig 52.
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Classification: Sordariomycetes, Sordariomycetidae, Magnaporthales, Pyriculariaceae.
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Type species: Pyricularia grisea Sacc. Lectotype designated by Rossman et al. (1990): BPI undistributed
set. Epitype and ex-epitype strain designated by Crous et al. (2015a): CBS H-22280, CBS 138707.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, act, cal, rpb1. Table 18. Fig. 28.
Ascomata ostiolate, solitary to gregarious, subspherical, brown to black, base immersed in host tissue,
with long neck protruding above plant tissue; ascomatal wall consisting of several layers of brown textura
angularis. Paraphyses intermingled among asci, unbranched, septate. Asci 8-spored, hyaline,
subcylindrical to clavate, unitunicate, short-stipitate, with prominent apical ring. Ascospores bi- to
multiseriate in asci, hyaline, guttulate, smooth-walled, fusiform, curved with rounded ends, transversely
3-septate, slightly constricted at septa. Conidiophores solitary or in fascicles, subcylindrical, erect, brown,
smooth, rarely branched, with sympodial proliferation. Conidiogenous cells terminal and intercalary, pale
brown, with denticulate conidiogenous loci and rhexolytic secession. Conidia solitary, pyriform to
obclavate, narrowed toward tip, rounded at base, 2-septate, hyaline to pale brown, with a distinct basal
hilum, sometimes with marginal frill (adapted from Klaubauf et al. 2014).
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Culture characteristics: Colonies on MEA white to vinaceous, pale olivaceous grey, smoke grey, or grey,
cottony. Colonies on OA iron grey, transparent with greenish olivaceous parts, fuscous black with grey
centre or hazel with smokey grey tufts.
Distribution: Worldwide.
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Optimal media and cultivation conditions: On OA at 25 °C in dark, or autoclaved barley seed s placed on
SNA at 25 °C under near-ultraviolet light (12 h lig ht, 12 h dark).
Hosts: Wide range of monocot plants, including important crops of the Poaceae such as rice, barley,
millet, oat and wheat.
Disease symptoms: Leaf spot and blast diseases.
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Notes: Pyricularia was recently re-evaluated in a phylogenetic study based on five loci (Klaubauf et al.
2014). In this study, the polyphyletic nature of the genus was resolved introducing eight new genera to
accommodate the species of Pyricularia that were not grouped with the type species Py. grisea and
Py. oryzae in Pyricularia s.str. Moreover, the family Pyriculariaceae was introduced to accommodate
Pyricularia, which was previously considered a member of Magnaporthaceae. Pyriculariaceae, as well
as Magnaporthaceae, accommodate mainly plant pathogenic species, some of which are of major
importance in agriculture.
Pyricularia oryzae is the causal agent of rice blast disease, which can result in up to a 30 % yield loss
worldwide (Skamnioti & Gurr 2009). In a phylogenetic study based on 10 loci and 128 isolates of this
species, three major clades were identified (Castroagudín et al. 2016). The first clade grouped the
isolates associated only with rice and corresponds to the previously described rice blast pathogen Py.
oryzae pathotype Oryza. The second clade accommodated isolates associated almost exclusively with
wheat and corresponds to the previously described wheat blast pathogen Py. oryzae pathotype
Triticum. A third clade accommodated isolates obtained from wheat as well as other Poaceae. This
clade was distinct from Py. oryzae and represented a new species, Pyricularia graminis-tritici. This
new species could not be morphologically distinguished from Py. oryzae, but a distinctive
pathogenicity spectrum was observed (Castroagudín et al. 2016). However, the “oryzae” clade consists
of various populations specific to different grass hosts that appear to be in the process of speciation.
Consequently, the species in the “oryzae” clade are not commonly accepted, and some authors refer
to them as lineages of Py. oryzae (Castroagudín et al. 2017). Further research is needed to resolve
speciation within what is presently circumscribed as Py. oryzae.
The sexual morph has been reported only in Py. grisea and Py. oryzae, both being heterothallic
species. These are indistinguishable in conidium, perithecium and ascospore morphology. However, Py.
oryzae was described as a new species distinct from Py. grisea based on DNA sequence differences in
three different loci (act, cal and tub) and host range, since Py. grisea infects only crab grass (Couch &
Kohn 2002).
References: Klaubauf et al. 2014 (morphology and phylogeny); Castroagudín et al. 2016, 2017
(morphology, pathogenicity and phylogeny); Reges et al. 2016 (pathogenicity and phylogeny).
Authors: Y. Marin-Felix & P.W. Crous
Stenocarpella Syd. & P. Syd., Ann. Mycol. 15: 258. 1917. Fig. 53.
Synonyms: Hendersoniopsis Woron., Fungal and Bacterial Diseases of Agricultural Plants: 255. 1922.
Phaeostagonosporopsis Woron., La Defense des Plantes, Leningrad 2: 333. 1925.
Classification: Sordariomycetes, Diaporthomycetidae, Diaporthales, Diaporthaceae.
Type species: Stenocarpella macrospora (Earle) B. Sutton, basionym: Diplodia macrospora Earle.
Isotype: IMI 12790. Ex-epitype strain designated by Crous et al. (2006b): CBS 117560 = MRC 8615.
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DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, tef1. Table 19.
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Mycelium immersed, brown, branched, septate. Conidiomata pycnidial, solitary or sometimes confluent,
globose or elongated, dark brown, subepidermal, unilocular, conidiomatal wall composed of dark brown,
thick-walled cells of textura angularis; neck single, circular, papillate, protruding. Conidiophores usually
reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, determinate, discrete, rarely
integrated on 1-septate conidiophores, cylindrical, collarette with minute channel, periclinal wall
thickened, formed from inner cells of pycnidial wall. Conidia pale brown, 0−3-septate, continuous or
constricted, cylindrical to fusiform, straight or curved, apex obtuse, base tapered and truncate, thick and
smooth-walled, eguttulate. Beta conidia absent or present, hyaline, scolecosporous, curved. Sexual
morph unknown (adapted from Sutton 1980).
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Culture characteristics: Colonies flat, with abundant aerial mycelium giving a cottony appearance; under
continuous near-ultraviolet light abundant sporulation in 1 wk. On OA surface white to rosy buff to
vinaceous buff, centre isabelline; reverse vinaceous buff, centre isabelline.
Optimal media and cultivation conditions: OA and PNA at 25 °C under continuous near-ultravi olet light to
promote sporulation.
Distribution: Africa, America and Asia.
Host: Zea mays (Poaceae).
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Notes: The genus Stenocarpella was introduced by Sydow & Sydow (1917), with S. zeae designated as
type species. Sutton (1977) synonymised S. zeae with Diplodia macrospora, transferring this latter
species to Stenocarpella, recognising S. macrospora as the correct name for the type species. Two
species are included in this genus i.e. S. macrospora and S. maydis, which cause Diplodia ear rot of
maize (Crous et al. 2006b).
Stenocarpella was initially placed in the Botryosphaeriaceae (Botryosphaeriales) because of the
similarity with Diplodia. In a phylogenetic study based on LSU sequences, Crous et al. (2006b) showed
that Stenocarpella belongs to the Diaporthaceae (Diaporthales). Subsequently, Lamprecht et al. (2011)
confirmed this placement based on ITS and tef1 sequences.
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References: Sutton 1977, 1980 (morphology and pathogenicity); Crous et al. 2006b (morphology and
phylogeny); Lamprecht et al. 2011 (morphology, pathogenicity and phylogeny).
Authors: Y. Marin-Felix & P.W. Crous
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Utrechtiana Crous & Quaedvl., Persoonia 26: 153. 2011. Fig 54.
Classification: Sordariomycetes, Sordariomycetidae, Magnaporthales, Pyriculariaceae.
Type species: Utrechtiana roumeguerei (Cavara) Videira & Crous, basionym Scolicotrichum
roumeguerei Cavara = Utrechtiana cibiessia Crous & Quaedvlieg. Holotype and ex-type strain of
Utrechtiana cibiessia: CBS H-20594, CBS 128780.
DNA barcodes (genus): LSU, ITS.
DNA barcodes (species): ITS, act, cal, rpb1. Table 20. Fig. 28.
Mycelium internal, consisting of septate, smooth, hyaline, branched hyphae. Conidiophores
predominantly solitary, erect, straight to flexuous, unbranched, 1-septate, medium brown to dark
brown, subcylindrical with swollen basal cell. Conidiogenous cells integrated, terminal, cylindrical or
subcylindrical, smooth or finely verruculose, thick-walled with thin-walled, clavate, bluntly rounded
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apex, with truncate, flattened scar, holoblastic; conidiophores rejuvenating percurrently. Conidia
solitary, obpyriform or ellipsoid, pale brown, guttulate to granular, finely verruculose, 1–2-septate, thinwalled, apex bluntly to acutely rounded, base obtusely rounded with a flattened, darkened and thickened
hilum that has a central pore. Synasexual morph selenosporella-like present or absent.
Microconidiophores arranged in rosettes, branched, septate, pale brown, smooth, subcylindrical.
Microconidiogenous cells pale brown, smooth to finely roughened, phialidic, terminal and lateral,
fusoid-ellipsoid to ampulliform. Microconidia hyaline, smooth, aseptate, subcylindrical, straight to
curved, ends obtuse.
Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and even smooth
margins. On MEA surface dirty white, sometimes turning grey olivaceous when fertile; reverse luteous
or olivaceous grey in centre and luteous in outer region. On OA olivaceous grey to iron-grey or dirty
white.
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Optimal media and cultivation conditions: On OA at 25 °C under dark, or autoclaved barley s eeds placed
on SNA at 25 °C under near-ultraviolet light (12 h light, 12 h dark).
Distribution: America, Asia, Australia and Europe.
Hosts: Phragmites spp (Poaceae).
Disease symptoms: Leaf spot.
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Notes: The genus Utrechtiana was described by Crous et al. (2011a) to accommodate the type species
named Utrecthiana cibiessia, which is a foliar pathogen of Phragmites. However, this genus was
considered synonymous with Deightoniella by Seifert et al. (2011) because of the morphology of the
conidiophores (solitary, brown, with percurrent rejuvenation) and conidia (brown and septate). Moreover,
U. cibiessia was demonstrated to be a synonym of Deightoniella roumeguerei, which Klaubauf et al.
(2014) showed to belong to Pyriculariaceae, a family containing numerous cryptic fungal genera on
Poaceae.
However, Deightoniella has been shown to represent a polyphyletic genus. For example,
Deightoniella torulosa, which is a foliar pathogen of Musa, proved to be a species of Corynespora
(Crous et al. 2013), while a similar fungus occurring on leaf spots of Phragmites in South Africa was
placed in Neodeightoniella (Crous et al. 2013). In a recent study, Videira et al. (2017) considered
Utrechtiana and Deightoniella based on the type species Deightoniella africana to be different genera
based on morphological characteristics. Utrechtiana lacks torsive to flexuous conidiophores with
prominent conidiophore swellings, and its conidia are also pale brown, smooth to finely roughened,
with prominent thickened, darkened scars. In contrast, conidia in Deightoniella are medium brown,
verruculose, and obpyriform with prominent apical taper. In order to clarify the phylogenetic
relationships between both genera, fresh material of Deightoniella africana is needed.
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References: Constantinescu 1983 (morphology and pathogenicity); Mel’nik & Shabunin 2011
(morphology); Crous et al. 2011a, Klaubauf et al. 2014, Videira et al. 2017 (morphology and
phylogeny).
Utrechtiana arundinacea (Corda) Crous, Quaedvl. & Y. Marín, comb. nov. MycoBank MB824141. Fig.
54.
Basionym: Helminthosporium arundinaceum Corda, as “Helmisporium”, Icon. fung. (Prague) 3: 10, tab. 2,
fig. 25. 1839.
Synonyms: Napicladium arundinaceum (Corda) Sacc., Syll. fung. 4: 482. 1886.
Deightoniella arundinacea (Corda) S. Hughes, Mycol. Pap. 48: 29. 1952.
Causing blight-like amphigenous lesions along leaves of Phragmites, medium brown with red-purple
margins and yellow halo, extending across breadth of leaf, up to 7 mm diam, and along length, up to 20
cm long. Macroconidiophores 30–50 × 9–12 µm, amphigenous, predominantly solitary, but at times in
fascicles of up to three, straight to flexuous, unbranched, 1-septate, medium brown, smooth,
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subcylindrical with swollen basal cell, 10–15 µm diam. Macroconidiogenous cells 20–35 × 7–9 µm,
integrated, terminal, cylindrical, thick-walled with thin-walled apex, holoblastic; conidiophores proliferate
percurrently. Macroconidia (22–)37–42(–45) × (17–)19–20(–21) µm, solitary, obpyriform, pale brown,
guttulate, finely verruculose, (1–)2-septate, with distinct dark brown hilum, 3–4 µm. A selenosporella-like
synasexual morph develops in culture, with microconidiophores arranged in rosettes, 15–40 × 3–6 µm,
branched, 3–6-septate, pale brown, smooth, subcylindrical. Microconidiogenous cells 5–14 × 3–4 µm,
pale brown, smooth to finely roughened, phialidic, terminal and lateral, fusoid-ellipsoid to ampulliform.
Microconidia 7–10 × 1.5–2 µm, hyaline, smooth, aseptate, subcylindrical, straight to curved, ends obtuse.
Macroconidiophores in culture up to 6-septate, 100 µm tall. Macroconidia 23–50 × 11–15 µm, slender,
pyriform, prominently verrucose, medium brown.
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Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and even smooth
margins. On MEA surface dirty white, reverse olivaceous grey in centre, luteous in outer region. On OA
olivaceous grey to iron-grey.
Materials examined: Czech Republic, Prague, on living leaves of Phragmites sp. (Poaceae), 1838 (holotype in PRM missing, but
slide ex-holotype, DAOM 19793). The Netherlands, on leaves of Phragmites sp. (Poaceae), 2 Jun. 2017, A. Mulder (epitype of
Helminthosporium arundinaceum designated here CBS H-23402, MBT380884, culture ex-epitype CPC 33994).
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Notes: Utrechtiana arundinacea is a commonly encountered European taxon treated in Deightoniella in
previous studies (Constantinescu 1983, Mel’nik & Shabunin 2011, Ghosta & Abrinbana 2016).
Morphologically, U. arundinacea and U. constantinescui appear to be related.
Macroconidia of U. arundinaceum exhibit a strange phenomenon where a third septum develops 3–5
µm from the apex. The conidium body is prominently guttulate, except for this terminal chamber, which is
smooth, pale brown, and lacks any guttules. This strange conidial apex is also visible in conidia of U.
constantinescui (Mel’nik & Shabunin 2011), and apparently plays some role in infection/attachment,
probably exuding a mucoid droplet, as is also seen in some genera in the Pyriculariaceae (Klaubauf et al.
2014). Furthermore, Mel’nik & Shabunin (2011) illustrate a selenosporella-like synasexual morph in both
species, which has not been seen in U. roumeguerei, the type species of the genus.
Utrechtiana roumeguerei was considered conspecific with U. arundinacea (Ellis 1957) until
Constantinescu (1983) demonstrated that they are distinct species based on morphology and
pathogenicity. Utrechtiana arundinacea often produces percurrently proliferating conidiogenous cells and
obclavate 2-septate conidia, while U. roumeguerei is characterised by rarely percurrent conidiogenous
cells and ovate to broadly ellipsoidal, 1-septate conidia. Moreover, U. arundinacea produces systemic
infection in the host issues, whereas U. roumeguerei induces a local infection with limited development. In
the present study, the DNA data support the placement of both taxa in the same genus (Fig. 28).
The holotype specimen of Helminthosporium arundinaceum could not be located in PRM, and is
presumed missing. However, a slide from the original material was preserved in DAOM. Due to the lack of
living culture of that species, a specimen isolated from the same host and region is here designated as
epitype.
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Utrechtiana constantinescui (Melnik & Shabunin) Crous & Y. Marín, comb. nov. MycoBank MB824142
Basionym: Deightoniella constantinescui Melnik & Shabunin, Mikol. Fitopatol. 45: 257. 2011.
Notes: The new combination U. constantinescui is designated here based on the morphology of its
macro- and microconidial morphs. Fresh material should be recollected to verify this placement. As we
mentioned above, this species is morphologically related to U. arundinacea. Both species can be
distinguished based on the shape of their macroconidia (obpyriform in U. arundinacea vs. barrel-shaped
in U. constantinescui) and the position of the conidial septa in U. constantinescui, 7–13 µm apart.
Authors: Y. Marin-Felix, W. Quaedvlieg & P.W. Crous
Wojnowiciella Crous et al., Persoonia 34: 201. 2015. Fig. 55.
Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Phaeosphaeriaceae.
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Type species: Wojnowiciella eucalypti Crous et al. Holotype and ex-type strain: CBS H-22233, CBS
139904.
DNA barcode (genus): LSU.
DNA barcodes (species): ITS, rpb2, tef1. Table 21. Fig. 56.
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Conidiomata pycnidial, globose, brown, separate, non-papillate or papillate, with central ostiole;
conidiomata wall composed of 3–6 layers of brown cells, textura angularis. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells lining cavity, hyaline to pale brown, smooth, ampulliform to
subcylindrical, appearing phialidic. Macroconidia subcylindrical, straight to slightly curved, apex
subobtuse, base truncate, septate, at times with 1–2 oblique septa, thick-walled, verruculose, guttulate,
golden brown. Microconidia in same or different conidiomata as macroconidia. Microconidiophores
intermingled with macroconidiogenous cells, branched at base, septate, subcylindrical, hyaline, smooth.
Microconidiogenous cells terminal and intercalary, hyaline, smooth, ampulliform to subcylindrical,
phialidic with periclinal thickening. Microconidia solitary, hyaline, guttulate, smooth, subcylindrical to
ellipsoid, apex obtuse to subobtuse, base truncate.
Culture characteristics: Colonies on MEA, cottony, isabelline, greenish olivaceous, mouse grey to
greyish sepia, sometimes with luteous exudate; reverse greyish sepia, chestnut, fulvous. Colonies on
PDA pale mouse grey, brown vinaceous or greenish olivaceous, sometimes with luteous diffusible
pigment; reverse luteous and black, greyish sepia or brown vinaceous.
Optimal media and cultivation conditions: On autoclaved banana leaves placed on SNA at 25 °C under
near-ultraviolet light (12 h light, 12 h dark).
Distribution: Australia, China, Colombia, Italy, South Africa.
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Disease symptoms: Leaf spots.
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Hosts: Eucalyptus grandis (Myrtaceae), Cissampelos capensis (Menispermaceae), Dactylis glomerata
(Poaceae), Leptocarpus sp. (Restionaceae), Lonicera sp. and Viburnum utile (Caprifoliaceae), and
Spartium sp. (Fabaceae).
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Notes: Wojnowiciella was established with W. eucalypti as type species, which differs from Septoriella
hirta (syn. Wojnowicia hirta) by non-setous conidiomata, dark brown conidia and hyaline microconidia
(Crous et al. 2015d). Although both genera belong to Phaeosphaeriaceae, Wojnowicia has been
synonymised with Septoriella (Crous et al. 2015a). Currently Wojnowiciella comprises seven species
isolated from leaf spots and twigs of different hosts (Table 21). Although they were associated with
disease symptoms, their pathogenicity needs to be proven.
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References: Wijayawardene et al. 2013 (morphology, as Wojnowicia); Crous et al. 2015d, 2016b
(morphology); Li et al. 2015 (morphology and phylogeny, as Wojnowicia); Liu et al. 2015 (morphology
and phylogeny, as Wojnowicia); Hernández-Restrepo et al. 2016c (morphology and phylogeny).
Authors: M. Hernández-Restrepo & P.W. Crous
ACKNOWLEDGEMENTS
Yasmina Marin-Felix is grateful for the financial support received from the Vice- Chancellor's postdoctoral fellowship programme
from University of Pretoria, South Africa. Keith A. Seifert is thanked for making images of Helminthosporium arundinaceum
(slide ex-holotype, DAOM 19793) available for comparison. Didier Tharreau is thanked for digital images of Pyricularia oryzae,
and Susan Thompson for Diaporthe gulyae.
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REFERENCES
AC
C
EP
TE
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M
AN
US
C
RI
PT
Agustí-Brisach C, León M, García-Jiménez J, et al. (2015). Detection of grapevine fungal trunk pathogens on pruning shears and
evaluation of their potential for spread of infection. Plant Disease 99: 976–981.
Al-Subhi AM, Al-Adawi AO, Van Wyk M, et al. (2006). Ceratocystis omanensis, a new species from diseased mango trees in
Oman. Mycological Research 110: 237–245.
Andjic V, Hardy GE, Cortinas MN, et al. (2007). Multiple gene genealogies reveal important relationships between species of
Phaeophleospora infecting Eucalyptus leaves. FEMS Microbiology Letters 268: 22−33.
Ando Y, Masuya H, Aikawa T, et al. (2017). Diaporthe toxicodendri sp. nov., a causal fungus of the canker disease on
Toxicodendron vernicifluum in Japan. Mycosphere 8: 1157–1167.
Ando Y, Motohashi K, Yaguchi Y (2013). Taxonomic re-examination of Cryptomeria gall disease causing fungus. Japanese
Journal of Mycology 54: 15−26.
Ariyawansa HA, Hyde KD, Jayasiri SC, et al. (2015). Fungal diversity notes 111–252—taxonomic and phylogenetic contributions to
fungal taxa. Fungal Diversity 75: 27–274.
Aroca A, Raposo R (2007). PCR-based strategy to detect and identify species of Phaeoacremonium causing grapevine diseases.
Applied Environmental Microbiology 73: 2911–2918.
Aroca A, Raposo R (2009). Pathogenicity of Phaeoacremonium species on grapevines. Journal of Phytopathology 157: 413–419.
Asher MJC, Shipton PJ (1981). Biology and control of take-all. Academic Press, London, UK.
Augustin C, Ulrich K, Ward E, et al. (1999). RAPD-based inter- and intravarietal classification of fungi of the Gaeumannomyces–
Phialophora complex. Journal of Phytopathology 147: 109–117.
Aveskamp MM, Verkley GJ, de Gruyter J, et al. (2009). DNA phylogeny reveals polyphyly of Phoma section Peyronellaea and
multiple taxonomic novelties. Mycologia 101: 363–382.
Aveskamp MM, de Gruyter J, Woudenberg JHC, et al. (2010). Highlights of the Didymellaceae: a polyphasic approach to
characterise Phoma and related pleosporalean genera. Studies in Mycology 65: 1–60.
Baayen RP, Bonants PJ, Verkley G, et al. (2002). Nonpathogenic isolates of the citrus black spot fungus, Guignardia citricarpa,
identified as a cosmopolitan endophyte of woody plants, G. mangiferae (Phyllosticta capitalensis). Phytopathology 92:
464−477.
Bateman GL, Ward E, Antoniw JF (1992). Identification of Gaeumannomyces graminis var. tritici and G. graminis var. avenae
using a DNA probe and non-molecular methods. Mycological Research 96: 737–742.
Baudoin ABAM (1986). First report of Dichotomophthora indica on common Purslane in Virginia. Plant Disease 70: 352.
Blanco-Ulate B, Rolshausen P, Cantu D (2013). Draft genome sequence of the ascomycete Phaeoacremonium aleophilum
strain UCR-PA7, a causal agent of the esca disease complex in grapevines. Genome Announcements 1: e00390–13.
Boerema GH, de Gruyter J, Noordeloos ME, et al. (2004). Phoma identification manual: differentiation of specific and infraspecific taxa in culture. CABI Publishing, Wallingford, UK.
Boesewinkel HJ (1982). Cylindrocladiella, a new genus to accommodate Cylindrocladium parvum and other small-spored
species of Cylindrocladium. Canadian Journal of Botany 60: 2288–2294.
Braun U (1995). A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic hyphomycetes). Vol. 1. IHW
Verlag, Eching, Germany.
Bussaban B, Lumyong S, Lumyong P, et al. (2005). Molecular and morphological characterization of Pyricularia and allied
genera. Mycologia 97: 1002–1011.
Carlucci A, Lops F, Cibelli F, et al. (2015). Phaeoacremonium species associated with olive wilt and decline in southern Italy.
European Journal of Plant Pathology 141: 717–729.
Carstens E, Linde CC, Slabbert R, et al. (2017). A global perspective on the population structure and reproductive system of
Phyllosticta citricarpa. Phytopathology 107: 758–768.
Castroagudín VL, Danelli A, Moreira SI, et al. (2017). The wheat blast pathogen Pyricularia graminis-tritici has complex origins
and a disease cycle spanning multiple grass hosts. BioRxiv 203455; doi.org/10.1101/203455
Castroagudín VL, Moreira SI, Pereira DAS, et al. (2016). Pyricularia graminis-tritici, a new Pyricularia species causing wheat
blast. Persoonia 36: 199–216.
Chen SF, Van Wyk M, Roux J, et al. (2013). Taxonomy and pathogenicity of Ceratocystis species on Eucalyptus trees in South
China, including C. chinaeucensis sp. nov. Fungal Diversity 58: 267–279.
Constantinescu O (1983). Deightoniella on Phragmites. Proceedings van de Koninklijke Nederlandse Akademie van
Wetenschappen Section C 86: 137–141.
Couch BC, Kohn LM (2002). A multilocus gene genealogy concord-ant with host preference indicates segregation of a new
species, Magnaporthe oryzae, from M. grisea. Mycologia 94: 683–693.
Crous PW (2002). Taxonomy and pathology of Cylindrocladium (Calonectria) and allied genera. APS Press, St. Paul,
Minnesota, USA.
Crous PW, Carris LM, Giraldo A, et al. (2015a). The Genera of Fungi – fixing the application of the type species of generic
names – G 2: Allantophomopsis, Latorua, Macrodiplodiopsis, Macrohilum, Milospium, Protostegia, Pyricularia, Robillarda,
Rotula, Septoriella, Torula, and Wojnowicia. IMA Fungus 6: 163–198.
Crous PW, Decock C, Schoch CL (2001). Xenocylindrocladium guianense and X. subverticmatum, two new species of
hyphomycetes from plant debris in the tropics. Mycoscience 42: 559–566.
Crous PW, Gams W, Stalpers JA, et al. (2004a). MycoBank: an online initiative to launch mycology into the 21st century.
Studies in Mycology 50: 19–22.
Crous PW, Gams W, Wingfield MJ, et al. (1996). Phaeoacremonium gen. nov. associated with wilt and decline diseases of woody
hosts and human infections. Mycologia 88: 786–796.
Crous PW, Giraldo A, Hawksworth DL, et al. (2014a). The Genera of Fungi: fixing the application of the type species of generic
names. IMA Fungus 5: 141–160.
Crous PW, Groenewald JZ, Gams W (2003). Eyespot of cereals revisited: ITS phylogeny reveals new species relationships.
European Journal of Plant Pathology 109: 841–850.
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Crous PW, Groenewald JZ, Shivas RG, et al. (2011a). Fungal Planet Description Sheets: 69–91. Persoonia 26:108–156.
Crous PW, Hawksworth DL, Wingfield MJ (2015b). Identifying and naming plant-pathogenic fungi: past, present, and future.
Annual Review of Phytopathology 53: 247–267.
Crous PW, Mohammed C, Glen M, et al. (2007). Eucalyptus microfungi known from culture. 3. Eucasphaeria and
Sympoventuria genera nova, and new species of Furcaspora, Harknessia, Heteroconium and Phacidiella. Fungal Diversity
25: 19–36.
Crous PW, Phillips AJL, Wingfield MJ (1991). The genera Cylindrocladium and Cylindrocladiella in South Africa, with special
reference to forest nurseries. South African Journal of Forestry 157: 69– 85.
Crous PW, Quaedvlieg W, Hansen K, et al. (2014b). Phacidium and Ceuthospora (Phacidiaceae) are congeneric: taxonomic
and nomenclatural implications. IMA Fungus 5: 173–193.
Crous PW, Schroers H-J, Groenewald JZ, et al. (2006a). Metulocladosporiella gen. nov. for the causal organism of
Cladosporium speckle disease of banana. Mycological Research 110: 264–275.
Crous PW, Schumacher RK, Wingfield MJ, et al. (2015c). Fungal Systematics and Evolution, FUSE 1. Sydowia 67: 81–118.
Crous PW, Shivas RG, Quaedvlieg W, et al. (2014c). Fungal Planet description sheets: 214–280. Persoonia 32: 184–306.
Crous PW, Schumacher RK, Wingfield MJ, et al. (2018). New and interesting fungi. 1. Fungal Systematics and Evolution 1:
169–215.
Crous PW, Shivas RG, Wingfield MJ, et al. (2012a). Fungal Planet description sheets: 128–153. Persoonia 29: 146–201.
Crous PW, Slippers B, Wingfield MJ, et al. (2006b). Phylogenetic lineages in the Botryosphaeriaceae. Studies in Mycology 55:
235–253.
Crous PW, Summerell BA, Shivas RG, et al. (2011b). Fungal Planet description sheets: 92–106. Persoonia 27: 130–162.
Crous PW, Summerell BA, Shivas RG, et al. (2012b). Fungal Planet description sheets: 107–127. Persoonia 28: 138–182.
Crous PW, Summerell BA, Shivas RG, et al. (2012c). A re-appraisal of Harknessia (Diaporthales), and the introduction of
Harknessiaceae fam. nov. Persoonia 28: 49–65.
Crous PW, Summerell BA, Swart L, et al. (2011c). Fungal pathogens of Proteaceae. Persoonia 27: 20–45.
Crous PW, Verkley GJM, Groenewald JZ, et al. (2009). Fungal biodiversity. CBS Laboratory Manual Series 1. CBS-KNAW
Fungal Biodiversity Centre, Utrecht, The Netherlands.
Crous PW, Wingfield MJ (1993). A re-evaluation of Cylindrocladiella, and a comparison with morphologically similar genera.
Mycological Research 97: 433–448.
Crous PW, Wingfield MJ, Burgess TI, et al. (2016a). Fungal Planet description sheets: 469–557. Persoonia 37: 218–403.
Crous PW, Wingfield MJ, Burgess TI, et al. (2017a). Fungal Planet description sheets: 558–624. Persoonia 38: 240–384.
Crous PW, Wingfield MJ, Burgess TI, et al. (2017b). Fungal Planet description sheets: 625–715. Persoonia 39: 270–467.
Crous PW, Wingfield MJ, Guarro J, et al. (2013). Fungal Planet description sheets: 154–213. Persoonia 31: 188–296.
Crous PW, Wingfield MJ, Guarro J, et al. (2015d). Fungal Planet description sheets: 320–370. Persoonia 34: 167–266.
Crous PW, Wingfield MJ, Le Roux JJ, et al. (2015e). Fungal Planet description sheets: 371–399. Persoonia 35: 264–327.
Crous PW, Wingfield MJ, Park RF (1991). Mycosphaerella nubilosa a synonym of M. molleriana. Mycological Research 95:
628–632.
Crous PW, Wingfield MJ, Richardson DM, et al. (2016b). Fungal Planet description sheets: 400–468. Persoonia 36: 316–458.
Crous PW, Wingfield MJ, Schumacher RK, et al. (2014d). Fungal Planet description sheets: 281–319. Persoonia 33: 212–289.
da Silva MA, Correia KC, Barbosa MAG, et al. (2017). Characterization of Phaeoacremonium isolates associated with Petri disease
of table grape in Northeastern Brazil, with description of Phaeoacremonium nordesticola sp. nov. European Journal of Plant
Pathology 149: 695–709.
Damm U, Mostert L, Crous PW, et al. (2008). Novel Phaeoacremonium species associated with necrotic wood of Prunus trees.
Persoonia 20: 87–102.
De Beer ZW, Duong TA, Barnes I, et al. (2014). Redefining Ceratocystis and allied genera. Studies in Mycology 79: 187–219.
De Beer ZW, Seifert KA, Wingfield MJ (2013a). The ophiostomatoid fungi: their dual position in the Sordariomycetes. In: The
ophiostomatoid fungi: expanding frontiers. CBS Biodiversity Series 12 (Seifert KA, De Beer ZW, Wingfield MJ, eds). CBSKNAW Fungal Biodiversity Centre, Utrecht, The Netherlands: 1–19.
De Beer ZW, Seifert KA, Wingfield MJ (2013b). A nomenclator for ophiostomatoid genera and species in the Ophiostomatales
and Microascales. In: The ophiostomatoid fungi: expanding frontiers. CBS Biodiversity Series 12 (Seifert KA, De Beer ZW,
Wingfield MJ, eds). CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands: 245–322.
de Gruyter, Boerema GH (2002). Contributions towards a monograph of Phoma (Coelomycetes) VIII. Section Paraphoma: Taxa
with setose pycnidia. Persoonia 17: 541–561.
de Gruyter J, Woudenberg JH, Aveskamp MM, et al. (2010). Systematic reappraisal of species in Phoma section Paraphoma,
Pyrenochaeta and Pleurophoma. Mycologia 102: 1066–1081.
de Gruyter J, Woudenberg JHC, Aveskamp MM, et al. (2013). Redisposition of phoma-like anamorphs in Pleosporales reevaluation. Studies in Mycology 75: 1–36.
De Hoog GS, Guarro J, Gené J, et al. (2000). Atlas of clinical fungi. 2nd ed. Centraalbureau voor Schimmelcultures, Utrecht and
Universitat Rovira i Virgili, Reus.
De Hoog GS, Hermanides-Nijhof EJ (1977). Survey of the black yeasts and allied fungi. Studies in Mycology 15: 178–222.
De Hoog GS, van Oorschot CAN (1983). Taxonomy of the Dactylaria complex I: Notes on the genus Dichotomophthora.
Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen Section C 86: 55–61.
Deacon JW (1973). Phialophora radicicola and Gaeumannomyces graminis on roots of grasses and cereals. Transactions of the
British Mycological Society 61: 471–485.
Deacon JW (1974). Further studies on Phialophora radicicola and Gaeumannomyces graminis on roots and stem bases of
grasses and cereals. Transactions of the British Mycological Society 63: 307–327.
Dennis RWG (1960). British cup fungi and their allies: An introduction to the Ascomycetes. Ray Society, London, UK.
Dissanayake AJ, Camporesi E, Hyde KD, et al. (2017a). Molecular phylogenetic analysis reveals seven new Diaporthe species
from Italy. Mycosphere 8: 853–877.
Dissanayake AJ, Phillips AJL, Hyde KD, et al. (2017b). The current status of species in Diaporthe. Mycosphere 8: 1106–1156.
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Dissanayake AJ, Zhang W, Liu M, et al. (2017c). Diaporthe species associated with peach tree dieback in Hubei, China.
Mycosphere 8: 533–549.
Doilom M, Dissanayake AJ, Wanasinghe DN, et al. (2017). Microfungi on Tectona grandis (teak) in Northern Thailand. Fungal
Diversity 82: 107–182.
Donk MA (1968). Report of the committee for Fungi and Lichen 1964–1968. Taxon 17: 578–581.
Douhan GW, Murray TD, Dyer PS (2002). Species and mating-type distribution of Tapesia yallundae and T. acuformis and
occurrence of apothecia in the U.S. Pacific Northwest. Phytopathology 92: 703–709.
Du Z, Fan XL, Hyde KD, et al. (2016). Phylogeny and morphology reveal two new species of Diaporthe from Betula spp. in
China. Phytotaxa 269: 90–102.
Dupont J, Laloui W, Magnin S, et al. (2000). Phaeoacremonium viticola, a new species associated with esca disease of grapevine
in France. Mycologia 92: 499–504.
Eken C (2003). Dichotomophthora portulacae on Portulaca oleracea in Turkey. Mycotaxon 87: 153–156.
Elliott ML (1991). Determination of an etiological agent of Bermuda grass decline. Phytopathology 81: 1380–1384.
Elliott ML, Hagan AK, Mullen JM (1993). Association of Gaeumannomyces graminis var. graminis with a St. Augustine grass
root rot disease. Plant Disease 77: 206–209.
Ellis MB (1957). Some species of Deightoniella. Mycological Papers 66: 1–12.
Ellis MB (1971). Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, UK.
Eskalen A, Rooney-Latham S, Gubler WD (2005). Occurrence of Togninia fraxinopennsylvanica on esca-diseased grapevines
(Vitis vinifera) and declining ash trees (Fraxinus latifolia) in California. Plant Disease 89: 528.
Essakhi S, Mugnai L, Crous PW, et al. (2008). Molecular and phenotypic characterization of novel Phaeoacremonium species
associated with Petri disease and esca of grapevine. Persoonia 21: 119–134.
Fan XL, Hyde KD, Udayanga D, et al. (2015). Diaporthe rostrata, a novel ascomycete from Juglans mandshurica associated
with walnut dieback. Mycological Progress 14: 82.
Farr DF, Rossman AY (2017). Fungal Databases, U.S. National Fungus Collections, ARS, USDA. https://nt.arsgrin.gov/fungaldatabases/. Accessed on November 2017.
Freeman J, Ward E (2004). Gaeumannomyces graminis, the take-all fungus and its relatives. Molecular Plant Pathology 5: 235–
252.
Fries EM (1849). Summa vegetabilium Scandinaviae. Typographis Academica, Uppsala, Sweden.
Gao YH, Liu F, Cai L (2016). Unravelling Diaporthe species associated with Camellia. Systematics and Biodiversity 14: 102–
117.
Gao YH, Liu F, Duan W, et al. (2017). Diaporthe is paraphyletic. IMA Fungus 8: 153–187.
Gao YH, Su YY, Sun W (2015). Diaporthe species occurring on Lithocarpus glabra in China, with descriptions of five new
species. Fungal Biology 119: 295–309.
Gao YH, Sun W, Su YY (2014). Three new species of Phomopsis in Gutianshan Nature Reserve in China. Mycological Progress
13: 111–121.
Giraldo A, Crous PW, Schumacher RK, et al. (2017). The Genera of Fungi – G3: Aleurocystis, Blastacervulus,
Clypeophysalospora, Licrostroma, Neohendersonia and Spumatoria. Mycological Progress 16: 325–348.
Ghosta Y, Abrinbana M (2016). Deightoniella arundinacea, new to mycobiota of Iran. Rostaniha 17: 92–94.
Glienke C, Pereira OL, Stringari D, et al. (2011). Endophytic and pathogenic Phyllosticta species, with reference to those
associated with Citrus Black Spot. Persoonia 26: 47−56.
Gomes RR, Glienke C, Videira SIR, et al. (2013). Diaporthe: a genus of endophytic, saprobic and plant pathogenic fungi.
Persoonia 31: 1–41.
Graham AB, Johnston PR, Weir BS (2009). Three new Phaeoacremonium species on grapevines in New Zealand. Australasian
Plant Pathology 38: 505–513.
Gramaje D, Agustí-Brisach C, Pérez-Sierra A, et al. (2012). Fungal trunk pathogens associated with wood decay of almond trees on
Mallorca (Spain). Persoonia 28: 1–713.
Gramaje D, Armengol J, Mohammadi H, et al. (2009). Novel Phaeoacremonium species associated with Petri disease and esca of
grapevines in Iran and Spain. Mycologia 101: 920–929.
Gramaje D, García-Jiménez J, Armengol J (2010). Grapevine rootstock susceptibility to fungi associated with Petri disease and
esca under field conditions. American Journal of Enology and Viticulture 61: 512–7520.
Gramaje D, León M, Pérez-Sierra A, et al. (2014). New Phaeocremonium species isolated from sandalwood trees in Western
Australia. IMA Fungus 5: 67–77.
Gramaje D, Mostert L, Groenewald JZ, et al. (2015). Phaeoacremonium: From esca disease to phaeohyphomycosis. Fungal
Biology 119: 759–783.
Groenewald M, Kang J-C, Crous PW, et al. (2001). ITS and beta-tubulin phylogeny of Phaeoacremonium and Phaeomoniella
species. Mycological Research 105: 651–657.
Guarnaccia V, Crous PW (2017). Emerging citrus diseases in Europe caused by Diaporthe spp. IMA Fungus 8: 317–334.
Guarnaccia V, Groenewald JZ, Li H, et al. (2017). First report of Phyllosticta citricarpa and description of two new species, P.
paracapitalensis and P. paracitricarpa, from citrus in Europe. Studies in Mycology 87: 161–185.
Guarnaccia V, Groenewald JZ, Woodhall J, et al. (2018). Diaporthe diversity and pathogenicity revealed from a broad survey of
grapevine diseases in Europe. Persoonia 40: 135–153.
Guarnaccia V, Vitale A, Cirvilleri G, et al. (2016). Characterisation and pathogenicity of fungal species associated with branch
cankers and stem-end rot of avocado in Italy. European Journal of Plant Pathology 146: 963–976.
Halleen F, Mostert L, Crous PW (2007). Pathogenicity testing of lesser-known vascular fungi of grapevines. Australasian Plant
Pathology 36: 277–285.
Hay FS, Gent DH, Pilkington SJ, et al. (2015). Changes in distribution and frequency of fungi associated with foliar diseases
complex of pyrethrum in Australia. Plant Disease 9: 1227–1235.
Heath RN, Wingfield MJ, Wingfield BD, et al. (2009). Ceratocystis species on Acacia mearnsii and Eucalyptus spp. in eastern
and southern Africa including six new species. Fungal Diversity 34: 41–68.
Hernández-Restrepo M, Groenewald JZ, Crous PW (2016a). Taxonomic and phylogenetic re-evaluation of Microdochium,
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Monographella and Idriella. Persoonia 36: 57–82.
Hernández-Restrepo M, Groenewald JZ, Elliott ML, et al. (2016b). Take-all or nothing. Studies in Mycology 83: 19–48.
Hernández-Restrepo M, Schumacher RK, Wingfield MJ, et al. (2016c). Fungal Systematics and Evolution: FUSE 2. Sydowia 68:
193–230.
Hibbett DS, Ohman A, Glotzer D, et al. (2011). Progress in molecular and morphological taxon discovery in Fungi and options
for formal classification of environmental sequences. Fungal Biology Reviews 25: 38–47.
Hirata K, Kusaba M, Chuma I, et al. (2007). Speciation in Pyricularia inferred from multilocus phylogenetic analysis. Mycological
Research 111: 799–808.
Hong SK, Kim WG, Choi HW, et al. (2008). Identification of Microdochium bolleyi associated with basal rot of creeping bent
grass in Korea. Mycobiology 36: 77–80.
Hu DM, Cai L, Hyde KD (2012). Three new ascomycetes from freshwater in China. Mycologia 104: 1478–1489.
Huang F, Hou X, Dewdney MM, et al. (2013). Diaporthe species occurring on citrus in China. Fungal Diversity 61: 237–250.
Huang F, Udayanga D, Wang X, et al. (2015). Endophytic Diaporthe associated with Citrus, a phylogenetic reassessment with
seven new species from China. Fungal Biology 119: 331–347.
Hyde KD, Hongsanan S, Jeewon R, et al. (2016). Fungal diversity notes 367–492, taxonomic and phylogenetic contributions to
fungal taxa. Fungal Diversity 80: 1–270.
Inderbitzin P, Bostock RM, Subbarao KV (2012). Cylindrocladiella hahajimaensis, a new species of Cylindrocladiella transferred
from Verticillium. MycoKeys 4: 1–8.
Jaklitsch WM, Voglmayr H (2012). Phylogenetic relationships of five genera of Xylariales and Rosasphaeria gen. nov.
(Hypocreales). Fungal Diversity 52: 75–98.
Johnston PR, Seifert KA, Stone JK, et al. (2014). Recommendations on generic names competing for use in Leotiomycetes
(Ascomycota). IMA Fungus 5: 91–120.
Jones DR (2000). Fungal diseases of the foliage. In: Diseases of Banana, Abaca and Enset (Jones DR, ed). CABI Publishing,
Wallingford, UK: 108–111.
Kamgan NG, Jacobs K, De Beer ZW, et al. (2008). Ceratocystis and Ophiostoma species, including three new taxa, associated
with wounds on native South African trees. Fungal Diversity 29: 37–59.
Kamgan Nkuekam G, Wingfield MJ, Mohammed C, et al. (2012). Ceratocystis species, including two new species associated
with nitidulid beetles, on eucalypts in Australia. Antonie van Leeuwenhoek 101: 217–241.
Kamgan Nkuekam G, Wingfield MJ, Roux J (2013). Ceratocystis species, including two new taxa, from Eucalyptus trees in
South Africa. Australasian Plant Pathology 42: 283–311.
Kirk PM, Stalpers JA, Braun U, et al. (2013). A without-prejudice list of generic names of fungi for protection under the
International Code of Nomenclature for algae, fungi and plants. IMA Fungus 4: 381–443.
Klaubauf S, Tharreau D, Fournier E, et al. (2014). Resolving the polyphyletic nature of Pyricularia (Pyriculariaceae). Studies in
Mycology 79: 85–120.
Klisiewicz JM (1985). Growth and reproduction of Dichotomophthora portulacae and its biological activity on Purslane. Plant
Disease 69: 761–762.
Lamprecht SC, Crous PW, Groenewald JZ, et al. (2011). Diaporthaceae associated with root and crown rot of maize. IMA
Fungus 2: 13–24.
Lee S, Groenewald JZ, Crous PW (2004). Phylogenetic reassessment of the coelomycete genus Harknessia and its teleomorph
Wuestneia (Diaporthales), and the introduction of Apoharknessia gen. nov. Studies in mycology 50: 235–252.
Li WJ, Bhat DJ, Camporesi E, et al. (2015). New asexual morph taxa in Phaeosphaeriaceae. Mycosphere 6: 681–708.
Liu JK, Hyde KD, Jones EBG, et al. (2015). Fungal diversity notes 1–110: taxonomic and phylogenetic contributions to fungal
species. Fungal Diversity 72: 1–197.
Lombard L, Cheewangkoon R, Crous PW (2017). New Cylindrocladiella spp. from Thailand soils. Mycosphere 8: 1088–1104.
Lombard L, Shivas RG, To-Anun C, et al. (2012). Phylogeny and taxonomy of the genus Cylindrocladiella. Mycological Progress
11: 835–868.
Lombard L, van der Merwe NA, Groenewald JZ, et al. (2015). Generic concepts in Nectriaceae. Studies in Mycology 80: 189–
245.
Lombard L, van Leeuwen GCM, Guarnaccia V, et al. (2014). Diaporthe species associated with Vaccinium, with specific
reference to Europe. Phytopathologia Mediterranea 53: 287–299.
Lucas JA, Dyer PS, Murray TD (2000). Pathogenicity, hostspecificity and population biology of Tapesia spp., causal agents of
eyespot disease of cereals. Advances in Botanical Research 33: 226–258.
Luo J, Walsh E, Zhang N (2014). Four new species in Magnaporthaceae from grass roots in New Jersey Pine Barrens.
Mycologia 106: 580–588.
Luo J, Zhang N (2013). Magnaporthiopsis, a new genus in Magnaporthaceae. Mycologia 105: 1019–1029.
Machingambi NM, Dreyer LL, Oberlander KC, et al. (2015). Death of endemic Virgilia oroboides trees in South Africa caused by
Diaporthe virgiliae sp nov. Plant Pathology 64: 1149–1156.
Madrid H, da Cunha KC, Gené J, et al. (2014). Novel Curvularia species from clinical specimens. Persoonia 33: 48–60.
Manamgoda DS, Cai L, McKenzie EHC, et al. (2012). A phylogenetic and taxonomic re-evaluation of the Bipolaris –
Cochliobolus – Curvularia complex. Fungal Diversity 56: 131–144.
Manamgoda DS, Rossman AY, Castlebury LA, et al. (2014). The genus Bipolaris. Studies in Mycology 79: 221–288.
Marin-Felix Y, Groenewald JZ, Cai L, et al. (2017). Genera of phytopathogenic fungi: GOPHY 1. Studies in Mycology 86: 99–
216.
Mayers CG, Mcnew DL, Harrington TC, et al. (2015). Three genera in the Ceratocystidaceae are the respective symbionts of
three independent lineages of ambrosia beetles with large, complex mycangia. Fungal Biology 119: 1075–1092.
Mbenoun M, Wingfield MJ, Begoude Boyogueno AD, et al. (2014). Molecular phylogenetic analyses reveal three new
Ceratocystis species and provide evidence for geographic differentiation of the genus in Africa. Mycological Progress 13:
219–240.
McNeill J, Barrie FF, Buck WR, et al. (eds.) (2012). International Code of Nomenclature for algae, fungi and plants (Melbourne
Code). [Regnum Vegetabile no. 154]. A.R.G. Gantner Verlag KG.
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Mehrlich FP, Fitzpatrick HM (1935). Dichotomophthora portulacae, a pathogen of Portulaca oleracea. Mycologia 27: 543–550.
Mel'nik VA, Shabunin DA (2011). Deightoniella constantinescui sp. nov. on Phragmites australis. Mikologiya i Fitopatologiya 45:
257–259.
Moreno-Rico O, Groenewald JZ, Crous PW (2014). Foliicolous fungi from Arctostaphylos pungens in Mexico. IMA Fungus 5: 7–
15.
Moslemi A, Ades PK, Crous PW, et al. (2018). Paraphoma chlamydocopiosa sp. nov. and Paraphoma pye sp. nov., two new
species associated with leaf and crown infection of pyrethrum. Plant Pathology 67: 124–135.
Moslemi A, Ades PK, Groom T, et al. (2016). Paraphoma crown rot of pyrethrum (Tanacetum cinerariifolium). Plant Disease
100: 2363–2369.
Mostert L, Crous PW, Kang JC, et al. (2001). Species of Phomopsis and a Libertella sp. occurring on grapevines with specific
reference to South Africa: morphological, cultural, molecular and pathological characterization. Mycologia 93: 146–167.
Mostert L, Groenewald JZ, Summerbell RC, et al. (2005). Species of Phaeoacremonium associated with infections in humans
and environmental reservoirs in infected woody plants. Journal of Clinical Microbiology 43: 1752–1767.
Mostert L, Groenewald JZ, Summerbell RC, et al. (2006). Taxonomy and pathology of Togninia (Diaporthales) and its
Phaeoacremonium anamorphs. Studies in Mycology 54: 1–115.
Motohashi K, Inaba S, Anzai K, et al. (2009). Phylogenetic analyses of Japanese species of Phyllosticta sensu stricto.
Mycoscience 50: 291−302.
Moyo P, Allsopp E, Roets F, et al. (2014). Arthropods vector grapevine trunk disease pathogens. Phytopathology 104: 1063–1069.
Müller E, Samuels GJ (1984). Monographella maydis sp. nov. and its connection to the tar-spot disease of Zea mays. Nova
Hedwigia 40: 112–120.
Murray TD (1996). Resistance to benzimidazole fungicides in the cereal eyespot pathogen, Pseudocercosporella
herpotrichoides, in the Pacific Northwest 1984 to 1990. Plant Disease 80: 19–23.
Nag Raj TR (1993). Coelomycetous anamorphs with appendage-bearing conidia. Mycologue Publications, Waterloo, Canada.
Nag Raj TR, DiCosmo F (1981). A monograph of Harknessia and Mastigosporella with notes on associated teleomorphs.
Bibliotheca Mycologica 80: 1–62.
Nel WJ, Duong TA, Wingfield BD, et al. (2017). A new genus and species for the globally important, multi-host root pathogen
Thielaviopsis basicola. Plant Pathology 10.1111/ppa.12803.
Nirenberg HI (1981). Differenzierung der Erreger der Halmbruchkrankheit I. Morphologie. Zeitschrift für Pflanzenkrankheiten und
Pflanzenschutz 88: 241–248.
Nishikado Y (1917). Studies on the rice blast fungus. Berichte des Ohara Instituts für landwirtschaftliche Forschungen 1: 171–
218.
Ondřej M (1988). Pyricularia luzulae Ondřej sp. n. Ceská Mykologie 42: 81–83.
Parkinson VO, Sivanesan A, Booth C (1981). The perfect state of the rice leaf-scald fungus and the taxonomy of both the
perfect and imperfect states. Transactions of the British Mycological Society 76: 59–69.
Peerally A (1991) The classification and phytopathology of Cylindrocladium species. Mycotaxon 40: 323–366.
Persoon CH (1818). Traité sur les champignons comestibles, contenant l’indication des espèces nuisibles; a l’histoire des
champignons. Belin-Leprieur, Paris, France.
Pfeiffer CM, Wheeler JE, Gilbertson RL (1989). First report of Dichotomophthora indica as a pathogen of Myrtillocactus
geometrizans and Gymnocalycium mihanovichii var. friedrichii in Arizona. Plant Disease 73: 81.
Pouzoulet J, Mailhac N, Couderc C, et al. (2013). A method to detect and quantify Phaeomoniella chlamydospora and
Phaeoacremonium aleophilum DNA in grapevine-wood samples. Applied Microbiology Biotechnology 97: 10163–10175.
Pu J, Xie Y, Zhang X, et al. (2008). Preinfection behaviour of Phyllosticta musarum on banana leaves. Australasian Plant
Pathology 37: 60–64.
Quaedvlieg W, Verkley GJM, Shin HD, et al. (2013). Sizing up Septoria. Studies in Mycology 75: 307–390.
Rachdawong S, Cramer CL, Grabau EA, et al. (2002). Gaeumannomyces graminis vars. avenae, graminis, and tritici identified
using PCR amplification of avenacinase-like genes. Plant Disease 86: 652–660.
Raimondo ML, Lops F, Carlucci A (2014). Phaeoacremonium italicum sp nov., a new species associated with esca of grapevine in
southern Italy. Mycologia 106: 1119–1126.
Rao PN (1966). A new species of Dichotomophthora on Portulaca oleracea from Hyderabad, India. Mycopathologia et
Mycologia Applicata 28: 137–140.
Rayner RW (1970). A mycological colour chart. Commonwealth Mycological Institute, Kew, UK.
Réblová M (2011). New insights into the systematics and phylogeny of the genus Jattaea and similar fungi of the Calosphaeriales.
Fungal Diversity 49: 167–198.
Réblová M, Mostert L (2007). Romellia is congeneric with Togninia, and description of Conidiotheca gen. nov. for one species of
this genus with polysporous asci. Mycological Research 111: 299–307.
Réblová M, Jaklitsch WM, Réblová K, et al. (2015). Phylogenetic reconstruction of the Calosphaeriales and Togniniales using five
genes and predicted RNA secondary structures of ITS, and Flabellascus tenuirostris gen. et sp. nov. PloS ONE 10: e0144616
Reges JTA, Negrisoli MM, Dorigan AF, et al. (2016). Pyricularia pennisetigena and P. zingibericola from invasive grasses infect
signal grass, barley and wheat. Pesquisa Agropecuária Tropical, Goiânia 46: 206–214.
Reid J, Booth C (1989). On Cryptosporella and Wuestneia. Canadian Journal of Botany 67: 879–908.
Robert V, Vu D, Amor ABH, et al. (2013). MycoBank gearing up for new horizons. IMA Fungus 4: 371–379.
Roll-Hansen F (1992). Important pathogenic fungi on conifers in Iceland. Acta Botanica Islandica 11: 9–12.
Rooney-Latham S, Eskalen A, Gubler WD (2005). Teleomorph formation of Phaeoacremonium aleophilum, cause of esca and
grapevine decline in California. Plant Disease 89: 177–184.
Rossman AY, Howard RJ, Valent B (1990). Pyricularia grisea, the correct name for the rice blast disease fungus. Mycologia 82:
509–512.
Rossman AY, Seifert KA, Samuels GJ, et al. (2013). Genera in Bionectriaceae, Hypocreaceae, and Nectriaceae (Hypocreales)
proposed for acceptance or rejection. IMA Fungus 4: 41–51.
Routien JB (1957). Fungi isolated from soils. Mycologia 49: 188–196.
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Santos JM, Correia VG, Phillips AJL (2010). Primers for mating-type diagnosis in Diaporthe and Phomopsis, their use in
teleomorph induction in vitro and biological species definition. Fungal Biology 114: 255–270.
Santos L, Phillips AJL, Crous PW (2017). Diaporthe species on Rosaceae with descriptions of D. pyracanthae sp. nov. and D.
malorum sp. nov. Mycosphere 8: 485–511.
Scattolin L, Montecchio L (2007). First report of damping-off of common oak plantlets caused by Cylindrocladiella parva in Italy.
Plant Disease 91: 771.
Schoch CL, Crous PW, Wingfield MJ, et al. (2000). Phylogeny of Calonectria and selected hypocrealean genera with cylindrical
macroconidia. Studies in Mycology 45: 45–62.
Schoch CL, Seifert KA, Huhndorf S, et al. (2012). Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA
barcode marker for Fungi. Proceedings of the National Academy of Sciences, USA 109: 6241–6246.
Schoch CL, Shoemaker RA, Seifert KA, et al. (2006). A multigene phylogeny of the Dothideomycetes using four nuclear loci.
Mycologia 98: 1041–1052.
Seifert K, Morgan-Jones G, Gams W, et al. (2011). The genera of Hyphomycetes. CBS Biodiversity Series 9. CBS-KNAW
Fungal Biodiversity Centre, Utrecht, the Netherlands.
Senanayake IC, Crous PW, Groenewald JZ, et al. (2017). Families of Diaporthales based on morphological and phylogenetic
evidence. Studies in Mycology 86: 217–296.
Sivanesan A (1987). Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs.
Mycological Papers 158: 1–261.
Skamnioti P, Gurr SJ (2009). Against the grain: safeguarding rice from rice blast disease. Trends in Biotechnology 27: 141–150.
Smith H, Wingfield MJ, Crous PW, et al. (1996). Sphaeropsis sapinea and Botryosphaeria dothidea endophytic in Pinus spp.
and Eucalyptus spp. in South Africa. South African Journal of Botany 62: 86–88.
Soares DJ, Nechet KL (2017). Dichotomophthora sp. causing leaf spot and foliar abscission on Anredera cordifolia in Brazil.
Australasian Plant Disease Notes 12: 51.
Speakman JB (1982). A simple, reliable method of producing perithecia of Gaeumannomyces gramnis var. tritici and its
application to isolates of Phialophora spp. Transactions of the British Mycological Society 79: 350–353.
Speakman JB (1984). Perithecia of Gaeumannomyces graminis var. graminis and G. graminis var. tritici in pure culture.
Transactions of the British Mycological Society 82: 720–723.
Spies CFJ, Moyo P, Halleen F, et al. (2018). Phaeoacremonium species diversity on woody hosts in the Western Cape Province
of South Africa. Persoonia 40: 26–62.
Sprague R (1936). Relative susceptihility of certain species of gramineae to Cercosporella herpotrichoides. Journal of
Agricultural Research 53: 659–670.
Sprague R, Fellows H (1934). Cercosporella foot rot of winter cereals. Technical Bulletin 428: 1–24.
Su YY, Cai L (2012). Polyphasic characterisation of three new Phyllosticta spp. Persoonia 28: 76−84.
Sutton BC (1977). Coelomycetes VI. Nomenclature of generic names proposed for Coelomycetes. Mycological Papers 141: 1–
253.
Sutton BC (1980). The Coelomycetes. Fungi imperfecti with pycnidia, acervuli and stromata. Commonwealth Mycological
Institute, Kew, UK.
Sydow H, Sydow P (1917). Beitrag zur Kenntniss der Pilzflora der Philippinen-Inseln. Annales Mycologici 15: 165–268.
Sydow H (1924). Sydow, Mycotheca germanica. Fasc. XLII-XLV (No. 2051–2250). Annales Mycologici 22: 257–268.
Tan YP, Edwards J, Grice KRE, et al. (2013). Molecular phylogenetic analysis reveals six new Diaporthe species from Australia.
Fungal Diversity 61: 251–260.
Tanney JB, McMullin DR, Green BD, et al. (2016). Production of antifungal and antiinsectan metabolites by the Picea endophyte
Diaporthe maritima sp. nov. Fungal Biology 120: 1448–1457.
Tarigan M, Gryzenhout M, Roux J, et al. (2010). Three new Ceratocystis spp. in the Ceratocystis moniliformis complex from
wounds on Acacia mangium and A. crassicarpa. Mycoscience 51: 53–67.
Thompson SM, Tan YP, Shivas RG, et al. (2015). Green and brown bridges between weeds and crops reveal novel Diaporthe
species in Australia. Persoonia 35: 39–49.
Thompson SM, Tan YP, Young AJ, et al. (2011). Stem cankers on sunflower (Helianthus annuus) in Australia reveal a complex
of pathogenic Diaporthe (Phomopsis) species. Persoonia 27: 80–89.
Turner EM (1940). Ophiobolus graminis Sacc. var. avenae var. n. as the cause of take-all or white-heads in Wales. Transactions
of the British Mycological Society 24: 269–281.
Udayanga D, Castlebury LA, Rossman AY, et al. (2014a). Insights into the genus Diaporthe: phylogenetic species delimitation in
the D. eres species complex. Fungal Diversity 67: 203–229.
Udayanga D, Castlebury LA, Rossman AY, et al. (2014b). Species limits in Diaporthe: molecular re-assessment of D. citri, D.
cytosporella, D. foeniculina and D. rudis. Persoonia 32: 83–101.
Udayanga D, Castlebury LA, Rossman AY, et al. (2015). The Diaporthe sojae species complex, phylogenetic re-assessment of
pathogens associated with soybean, cucurbits and other field crops. Fungal Biology 119: 383–407.
Udayanga D, Liu X, McKenzie EHC, et al. (2011). The genus Phomopsis: biology, applications, species concepts and names of
common phytopathogens. Fungal Diversity 50: 189–225.
Udayanga D, Liu XZ, MCkenzie EHC, et al. (2012). Multi-locus phylogeny reveals three new species of Diaporthe from Thailand.
Cryptogamie Mycologie 33: 295–309.
Ulrich K, Augustin C, Werner A (2000). Identification and characterization of a new group of root-colonizing fungi within the
Gaeumannomyces–Phialophora complex. New Phytologist 145: 127–135.
Uotila A (1990). Infection of pruning wounds in Scots pine by Phacidium coniferarum and selection of pruning season. Acta
Forestalia Fennica 215: 1–36
Úrbez-Torres JR, Haag P, Bowen P, et al. (2014). Grapevine trunk diseases in British Columbia: incidence and characterization
of the fungal pathogens associated with esca and Petri diseases of grapevine. Plant Disease 98: 469–482.
Úrbez-Torres JR, Haag P, Bowen P, et al. (2015). Development of a DNA-macroarray for the detection and identification of fungal
pathogens causing decline of young grapevines. Phytopathology 105: 1373–1383.
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Van Coller GJ, Denman S, Groenewald JZ, et al. (2005). Characterisation and pathogenicity of Cylindrocladiella spp. associated
with root and cutting rot symptoms of grapevines in nurseries. Australasian Plant Pathology 34: 489–498.
van der Aa HA (1973). Studies in Phyllosticta I. Studies in Mycology 5: 1−110.
van der Aa HA, Vanev S (2002). A revision of the species described in Phyllosticta. Centraalbureau voor Schimmelcultures,
Utrecht, The Netherlands.
Van Niekerk JM, Groenewald JZ, Farr DF, et al. (2005). Reassessment of Phomopsis species on grapevines. Australasian Plant
Pathology 34: 27–39.
Van Rensburg JCJ, Lamprecht SC, Groenewald JZ, et al. (2006). Characterization of Phomopsis spp. associated with die-back
of rooibos (Aspalathus linearis) in South Africa. Studies in Mycology 55: 65–74.
Van Wyk M, Roux J, Barnes I, et al. (2004). Ceratocystis bhutanensis sp. nov., associated with the bark beetle Ips
schmutzenhoferi on Picea spinulosa in Bhutan. Studies in Mycology 50: 365–379.
Van Wyk M, Roux J, Barnes I, et al. (2006). Molecular phylogeny of the Ceratocystis moniliformis complex and description of C.
tribiliformis sp. nov. Fungal Diversity 21: 181–201.
Van Wyk M, Wingfield BD, Wingfield MJ (2011). Four new Ceratocystis spp. associated with wounds on Eucalyptus,
Schizolobium and Terminalia trees in Ecuador. Fungal Diversity 46: 111–131.
Videira SIR, Groenewald JZ, Nakashima C, et al. (2017). Mycosphaerellaceae – Chaos or clarity? Studies in Mycology 87: 257–
421.
von Arx JA (1981). Notes on Microdochium and Idriella. Sydowia 34: 30–38.
von Arx JA (1984). Notes on Monographella and Microdochium. Transactions of the British Mycological Society 83: 373–374.
von Arx JA, Olivier D (1952). The taxonomy of Ophiobolus graminis Sacc. Transactions of the British Mycological Society 35:
29–33.
Walker J (1972). Type studies on Gaeumannomyces graminis and related fungi. Transactions of the British Mycological Society
58: 427–457.
Walker J (1975). Take-all disease of Gramineae: A review of recent work. Review of plant pathology 54: 113–143.
Walker J (1980). Gaeumannomyces, Linocarpon, Ophiobolus and several other genera of scolecospored Ascomycetes and
Phialophora conidial states, with a note on hyphopodia. Mycotaxon 11: 1–129.
Walker J (1981). Taxonomy of take-all fungi and related genera and species. In: Biology and Control of Take-all (Asher MJC,
Shipton PJ, eds). Academic Press, London, UK: 15–84.
Wang X, Chen G, Huang F, et al. (2011). Phyllosticta species associated with citrus diseases in China. Fungal Diversity 52:
209–224.
Ward E, Bateman GL (1999). Comparison of Gaeumannomyces- and Phialophora-like fungal pathogens from maize and other
plants using DNA methods. New Phytologist 141: 323–331.
Wicht B, Petrini O, Jermini M, et al. (2012). Molecular, proteomic and morphological characterization of the ascomycete
Guignardia bidwellii, agent of grape black rot: a polyphasic approach to fungal identification. Mycologia 104: 1036–1045.
Wijayawardene N, Song Y, Bhat DJ, et al. (2013). Wojnowicia viburni, sp. nov., from China and its phylogenetic placement.
Sydowia 65: 129–138.
Wikee S, Lombard L, Crous PW, et al. (2013a). Phyllosticta capitalensis, a widespread endophyte of plants. Fungal Diversity 60:
91–105.
Wikee S, Lombard L, Nakashima C, et al. (2013b). A phylogenetic re-evaluation of Phyllosticta (Botryosphaeriales). Studies in
Mycology 76: 1–29.
Wikee S, Udayanga D, Crous PW, et al. (2011). Phyllosticta – an overview of current status of species recognition. Fungal
Diversity 51: 43–61.
Wilson M (1920). A new disease of the Douglas Fir in Scotland. Transactions of the Royal Scottish Arboricultural Society 34:
145–149.
Wilson AM, Godlonton T, van der Nest MA, et al. (2015). Unisexual reproduction in Huntiella moniliformis. Fungal Genetics and
Biology 80: 1–9.
Wingfield BD, Van Wyk M, Roos H, et al. (2013). Ceratocystis: emerging evidence for discrete generic boundaries. In: The
ophiostomatoid fungi: Expanding frontiers. CBS biodiversity series 12 (Seifert KA, De Beer ZW, Wingfield MJ, eds). CBSKNAW Fungal Biodiversity Centre, Utrecht, The Netherlands: 57–64.
Wong MH, Crous PW, Henderson J, et al. (2012). Phyllosticta species associated with freckle disease of banana. Fungal
Diversity 56: 173–187.
Wong PTW (2002). Gaeumannomyces wongoonoo sp. nov., the cause of a patch disease of buffalo grass (St Augustine grass).
Mycological Research 106: 857–862.
Wong PTW, Walker J (1975). Germinating phialidic conidia of Gaeumannomyces graminis and phialophora-like fungi from
Gramineae. Transactions of the British Mycological Society 65: 41–47.
Wulandari NF, To-anun C, Hyde KD, et al. (2009). Phyllosticta citriasiana sp. nov., the cause of Citrus tan spot of Citrus maxima
in Asia. Fungal Diversity 34: 23–39.
Yang Q, Du Z, Tian CM (2018). Phylogeny and morphology reveal two new species of Diaporthe from Traditional Chinese
Medicine in Northeast China. Phytotaxa 336: 159–170.
Yang Q, Fan XL, Du Z, et al. (2017a). Diaporthe juglandicola sp. nov. (Diaporthales, Ascomycetes) evidenced by morphological
characters and phylogenetic analysis. Mycosphere 8: 817–826.
Yang Q, Fan XL, Du Z, et al. (2017b). Diaporthe species occurring on Senna bicapsularis in southern China, with descriptions of
two new species. Phytotaxa 302: 145–155.
Yang Q, Fan XL, Du Z, et al. (2017c). Diaporthe camptothecicola sp. nov. on Camptotheca acuminata in China. Mycotaxon 132:
591–601.
Yao JM, Wang YC, Zhu YG (1992). A new variety of the pathogen of maize take-all. Acta Mycologica Sinica 11: 99–104.
Yuan Z-Q, Mohammed C (2002). Ceratocystis moniliformopsis sp. nov., an early coloniser of Eucalyptus obliqua logs in
Tasmania, Australia. Australian Systematic Botany 15: 125–133.
Zhang K, Su YY, Cai L (2013a). Morphological and phylogenetic characterization of two new species of Phyllosticta from China.
Mycological Progress 12: 547–556.
ACCEPTED MANUSCRIPT
RI
PT
Zhang K, Zhang N, Cai L (2013b). Typification and phylogenetic study of Phyllosticta ampelicida and P. vaccinii. Mycologia 105:
1030–1042.
Zhang W, Nan Z, Tian P, et al. (2015). Microdochium paspali, a new species causing seashore paspalum disease in southern
China. Mycologia 107: 80–89.
Zhang Y, Crous PW, Schoch CL, et al. (2012). Pleosporales. Fungal Diversity 53: 1–221.
Zhang Y, Schoch CL, Fournier J, et al. (2009). Multi-locus phylogeny of Pleosporales: a taxonomic, ecological and evolutionary
re-evaluation. Studies in Mycology 64: 85–102.
Zhang ZF, Liu F, Zhou X, et al. (2017). Culturable mycobiota from Karst caves in China, with descriptions of 20 new species.
Persoonia 39:1–31.
Zhou N, Chen Q, Carroll G, et al. (2015). Polyphasic characterization of four new plant pathogenic Phyllosticta species from
China, Japan and the United States. Fungal Biology 119: 433–446.
Legends to Figures
Fig. 1. Allantophomopsiella pseudotsugae (CBS 841.91). A. Conidiomata forming on autoclaved barley leaves. B–E. Conidiogenous
cells giving rise to conidia. F. Conidia. Scale bars: A = 300 m; others = 10 m (from Crous et al. 2014b).
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Fig. 2. Apoharknessia spp. A. Disease symptoms of Apoharknessia eucalyptorum on Eucalyptus pellita. B, C. Conidiomata. B.
Apoharknessia eucalyptorum (ex-type CBS 142519). C. Apoharknessia insueta (ex-type CBS 111377). D–G. Conidiogenous cells
and conidia. D, E. Apoharknessia eucalyptorum (ex-type CBS 142519). F, G. Apoharknessia insueta (ex-type CBS 111377). H, I.
Conidia. H. Apoharknessia eucalyptorum (ex-type CBS 142519). I. Apoharknessia insueta (ex-type CBS 111377). Scale bars: C =
25 µm; all others = 10 µm. Picture B taken from Crous et al. (2017a).
Fig. 3. Apoharknessia eucalypti (ex-type CBS 142518). A. Conidiomata sporulating on SNA. B–E. Conidiogenous cells and conidia.
F, G. Conidia. Scale bars: A = 250 m, B–G = 10 m.
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Fig. 4. Cylindrocladiella spp. and disease symptoms. A–D. Disease symptoms associated with Cylindrocladiella spp. A–B.
Cylindrocladiella spp. sporulating on the seed coat of Medicago sativa. C. Root rot of M. sativa seedling. D. Cylindrocladiella spp. on
carnation leaf. E–AA. Asexual morph. E–I. Penicillate conidiophores. E. Cylindrocladiella longistipitata (ex-type CBS 116075). F.
Cylindrocladiella kurandica (ex-type CBS 129577). G. Cylindrocladiella lanceolata (ex-type CBS 129566). H. Cylindrocladiella
pseudoparva (ex-type CBS 129560). I. Cylindrocladiella nederlandica (ex-type CBS 152.91). J–N. Penicillate conidiophores. J.
Cylindrocladiella hawaiiensis (ex-type CBS 129569). K. Cylindrocladiella australiensis (ex-type CBS 129567). L. Cylindrocladiella
natalensis (ex-type CBS 114943). M. Cylindrocladiella cymbiformis (ex-type CBS 129553). N. Cylindrocladiella ellipsoidea (ex-type
CBS 129573). O–S. Subverticillate conidiophores. O. Cylindrocladiella australiensis (ex-type CBS 129567). P. Cylindrocladiella
longiphialidica (ex-type CBS 129557). Q. Cylindrocladiella pseudohawaiiensis (ex-type CBS 210.94). R–S. Cylindrocladiella
natalensis (ex-type CBS 114943). T–Y. Terminal vesicles of stipe extensions. T. Cylindrocladiella hawaiiensis (ex-type CBS
129569). U. Cylindrocladiella stellenboschensis (ex-type CBS 110668). V. Cylindrocladiella cymbiformis (ex-type CBS 129553). W.
Cylindrocladiella variabilis (ex-type CBS 129561). X. Cylindrocladiella lanceolata (ex-type CBS 129566). Y. Cylindrocladiella
kurandica (ex-type CBS 129577). Z–AA. Conidia. Z. Cylindrocladiella natalensis (ex-type CBS 114943). AA. Cylindrocladiella
brevistipitata (ex-type CBS 142783). Scale bars E–I, O = 50 m; J–N, P–AA = 10 m.
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Fig. 5. The Maximum Likelihood (ML) consensus tree of Cylindrocladiella spp. inferred from the combined ITS (547 bp), tef1 (527
bp) and tub2 (502 bp) sequence alignment. Thickened lines indicate branches present in the ML, Maximum parsimony (MP) and
Bayesian consensus trees. Branches with ML-bootstrap (BS) & MP-BS = 100 % and posterior probabilities (PP) = 1.00 are in blue.
Branches with ML-BS & MP-BS ≥ 75 % and PP ≥ 0.95 are in green. The scale bar indicates 0.02 expected changes per site. The
tree is rooted to Gliocladiopsis sagariensis CBS 199.55. Ex-type strains are indicated in bold. GenBank accession numbers are
indicated in Table 3 and in Lombard et al. (2012, 2017). TreeBASE: S22340.
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Fig. 6. Cylindrocladiella nauliensis (ex-type CBS 143792). A–D. Penicillate conidiophores. E–H. Penicillate conidiogenous
apparatus. I–L. Terminal vesicles of stipe extensions. M–N. Subverticillate conidiophores. O. Conidia. Scale bars A–D = 50 m; E–
O = 10 m.
Fig. 7. Disease symptoms associated with Diaporthe spp. A, B. Helianthus annuus plants affected by Diaporthe gulyae (courtesy
Susan Thompson). C, D. Branch canker of Persea americana with associated Diaporthe foeniculina and Diaporthe sterilis. E, F.
Phomopsis cane (courtesy Alessandro Vitale) and cane bleaching on shoot of Vitis vinifera caused by Diaporthe spp. (courtesy José
Luis Ramos Sáez de Ojer). G, H. Decay of Vaccinium corymbosum caused by Diaporthe baccae and Diaporthe sterilis and artificial
infection caused by inoculation of Diaporthe sterilis. I. Trunk canker with gummosis of Citrus limon caused by Diaporthe limonicola
and Diaporthe melitensis. Pictures C, D taken from Guarnaccia et al. (2016); I from Guarnaccia & Crous et al. (2017).
Fig. 8. Diaporthe spp. A–D. Sexual morph. A, B. Ascomata. A. Diaporthe ambigua (ex-type CBS 114015). B. Diaporthe aspalathi
(ex-type CBS 117169). C, D. Asci with ascospores. C. Diaporthe ambigua (ex-type CBS 114015). D. Diaporthe aspalathi (ex-type
CBS 117169). E–M. Asexual morph. E, F. Conidiomata sporulating. E. Diaporthe limonicola (ex-type CBS 142549). F. Diaporthe
pseudomangiferae (ex-type CBS 101339). G–I. Conidiogenous cells and conidia. G. Diaporthe raonikayaporum (ex-type CBS
133182). H. Diaporthe tecomae (CBS 100547). I. Diaporthe limonicola (ex-type CBS 142549). J, K. Alpha conidia. J. Diaporthe
cinerascens (CBS 719.96). K. Diaporthe pseudomangiferae (ex-type CBS 101339). L. Beta conidia of Diaporthe eleagni (CBS
504.72). M. Alpha, beta and gamma conidia of Diaporthe limonicola (ex-type CBS 142549). Scale bars = 10 m. Pictures A–D taken
from Van Rensburg et al. (2006); F–H, J–L from Gomes et al. (2013); E, I, M from Guarnaccia & Crous et al. (2017).
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Fig. 9. Consensus phylogram of 2 052 trees resulting from a Bayesian analysis of the combined ITS (637 bp), tub2 (833 bp), his3
(592 bp), tef1 (496 bp) and cal (817 bp) sequence alignment of Diaporthe spp. Bootstrap support values and Bayesian posterior
probability values are indicated at the nodes. Substrate and country of origin are listed next to the strain numbers. The newly
recognised species are showed in bold. The tree was rooted to Diaporthella corylina CBS 121124. T indicates ex-type strain.
TreeBASE: S21865.
Fig. 10. Diaporthe heterophyllae (ex-type CBS 143769). A–C. Colonies on MEA, PDA and OA, respectively. D. Conidiomata
sporulating on PNA. E. Conidiogenous cells and conidia. F. Alpha and beta conidia. Scale bars = 10 m.
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Fig. 11. Diaporthe racemosae (ex-type CBS 143770). A–C. Colonies on MEA, PDA and OA, respectively. D. Conidiomata
sporulating on PNA. E. Conidiogenous cells and conidia. F. Alpha conidia. Scale bars = 10 m.
Fig. 12. Dichotomophthora lutea. A–E. Colonies on PDA. F–J. Colonies on OA. K. Habit. L–N. Conidiophores and conidia. O–Q.
Conidiogenous cells. R, S. Conidiogenous cells and conidia. T, U. Conidia and microconidia. V–X. Sclerotia. Y. Anastomosing
conidia. (A, F = ex-type CBS 145.57; B, G, K, O–T, V, Y = CBS 584.71; C, H = CBS 585.71; D, I, U = CBS 132.81; E, J, W, X =
CBS 518.78). Scale bars: V–X = 100 m; L, P = 20 m; others = 10 m.
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Fig. 13. RAxML phylogram obtained from the combined ITS (759 bp), LSU (880 bp), gapdh (594 bp) and rpb2 (958 bp) sequence
alignment of all the accepted species of Dichotomophthora. The tree was rooted to Curvularia portulacae CBS 239.48 and BRIP
14541. The novel species described in this study are shown in bold. RAxML bootstrap support (BS) values above 70 % are shown
in the nodes. GenBank accession numbers are indicated in Table 5. T and PT indicate ex-type and ex-paratype strains, respectively.
TreeBASE: S21899.
Fig. 14. Dichotomophthora basella (ex-type CPC 33016). A–C. Disease symptoms caused by Di. basella in leaves of Basella rubra.
D. Colony on PDA. E. Colony on OA. F. Colony overview with sclerotia and conidiophores. G–I. Conidiophores and conidia. J, K.
Conidiogenous cells. L. Conidia. M. Microconidia. N, O. Sclerotia. Scale bars: N, O = 100 m; H = 50 m; others = 10 m.
Fig. 15. Dichotomophthora brunnea (ex-type CBS 149.49). A. Colony on PDA. B. Colony on OA. C. Colony overview with
conidiophores. D–J. Conidiophores, conidiogenous cells and conidia. K. Microconidia. L–Q. Conidia. Scale bars: D–G = 50 m; H =
20 m; others = 10 m.
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Fig. 16. Gaeumannomyces spp. A–F. Sexual morph. A. Perithecia of Gaeumannomyces oryzicola (ex-type CBS 141390). B. Asci
and paraphyses of Gaeumannomyces oryzinus (CPC 26065). C–E. Asci. C, D. Gaeumannomyces oryzinus (CPC 26043). E.
Gaeumannomyces oryzicola (ex-type CBS 141390). F. Ascospores of Gaeumannomyces oryzinus (CBS 235.32). G–AH. Asexual
morph. G–P. Conidiophores and conidiogenous cells. G. Gaeumannomyces californicus (ex-type CBS 141377). H.
Gaeumannomyces fusiformis (ex-type CBS 141379). I. Gaeumannomyces arxii (CBS 903.73). J, N. Gaeumannomyces walkeri
(ex-type CBS 141400). K. Gaeumannomyces graminis (CBS 141386). L. Gaeumannomyces graminicola (CBS 352.93). M.
Gaeumannomyces oryzicola (ex-type CBS 141390). O. Gaeumannomyces oryzinus (CPC 26032). P. Gaeumannomyces
radicicola (ex-type CBS 296.53). Q–Y. Conidia. Q. Gaeumannomyces radicicola (ex-type CBS 296.53). R. Gaeumannomyces
oryzicola (ex-type CBS 141390). S, T. Gaeumannomyces walkeri (ex-type CBS 141400). U. Gaeumannomyces oryzinus (CPC
26067). V. Gaeumannomyces ellisiorum (ex-type CBS 387.81). W. Gaeumannomyces floridanus (ex-type CBS 141378). X.
Gaeumannomyces graminicola (CPC 26036). Y. Gaeumannomyces arxii (CBS 903.73). Z–AH. Hyphopodia. Z.
Gaeumannomyces ellisiorum (ex-type CBS 387.81).
AA, AC. Gaeumannomyces glycinicola (CBS 141380). AB.
Gaeumannomyces floridanus (ex-type CBS 141378). AD. Gaeumannomyces graminicola (CPC 26025). AE. Gaeumannomyces
californicus (ex-type CBS 141377). AG. Gaeumannomyces oryzinus (CPC 26032). AF. Gaeumannomyces hyphopodioides (CPC
26267). AH. Gaeumannomyces walkeri (ex-type CBS 141400). Scale bars A, B= 50 m; C–F = 20 m; other = 10 m. Pictures
taken from Hernández-Restrepo et al. (2016b).
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Fig. 17. RAxML phylogram obtained from the combined ITS (715 bp), LSU (881 bp), rpb1 (617 bp) and tef1 (427 bp) sequence
alignment of all the accepted species of Gaeumannomyces. The tree was rooted to Pseudophialophora eragrostis CM12m9 and
Falciphora oryzae CBS 125863. RAxML bootstrap support (BS) values above 70 % are shown in the nodes. GenBank accession
numbers are indicated in Table 6. T, ET and A indicate ex-type, ex-epitype and authentic strains, respectively. TreeBASE: S21899.
Fig. 18. Harknessia spp. A–E. Disease symptoms on Eucalyptus. A. Harknessia fusiformis (CPC 13649). B. Harknessia
hawaiiensis (CPC 15003). C. Harknessia ravenstreetina (ex-type CBS 132125). D. Harknessia rhabdosphaera (CPC 13593). E.
Harknessia globispora (CPC 14924). F–L. Sexual morph of Harknessia eucalyptorum (CPC 12697). F. Ascoma with short neck,
oozing ascospores. G, H. Paraphyses and asci. I, J. Asci. K. Paraphyses and ascal tip. L. ascospores. M–AA. Asexual morphs. M.
Sporulating colony on OA of Harknessia ellipsoidea (ex-type CBS 132121). N–R. Conidiogenous cells giving rise to conidia. N, O.
Harknessia gibbosa (ex-type CBS 120033). P. Harknessia pseudohawaiiensis (CPC 17380). Q. Harknessia ravenstreetina (CBS
132125ET). R. Harknessia renispora (CPC 17163). S–X. Conidia. S, T. Harknessia australiensis (ex-type CBS 132119). U.
Harknessia kleinzeeina (ex-type CPC 16277). V. Harknessia eucalyptorum (CPC 12697). W. Harknessia ravenstreetina (ex-type
CBS 132125). X. Harknessia renispora (CPC 17163). Y. Microconidiogenous cells giving rise to microconidia of Harknessia
renispora (CPC 17163). Z, AA. Microconidia. Z. Harknessia renispora (CPC 17163). AA. Harknessia pseudohawaiiensis (CPC
17380). Scale bars: 10 m. Pictures taken from Crous et al. (2012c).
Fig. 19. RAxML phylogram obtained from the combined ITS (643 bp), cal (499 bp) and tub2 (838 bp) sequence alignment of all
accepted species of Harknessia. The tree was rooted to Cryphonectria parasitica. 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 were listed in Table 7 or in Crous et al. (2012c). T, ET and IsoT indicate ex-type, ex-epitype and
ex-isotype strains, respectively. TreeBASE: S21899.
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Fig. 20. Harknessia bourbonica (ex-type CBS 143913). A. Conidioma on OA. B, C. Conidiogenous cells giving rise to conidia. D.
Conidia. Scale bars: A = 150 m, B–D = 10 m.
Fig. 21. Harknessia corymbiae (ex-type CPC 33289). A. Conidiomata on OA. B. Conidiogenous cells giving rise to conidia. C, D.
Conidia. Scale bars: A = 250 m, B–D = 10 m.
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Fig. 22. Harknessia cupressi (ex-type CBS 143914). A. Conidiomata on OA. B, C. Conidiogenous cells giving rise to conidia. D, E.
Conidia. Scale bars: A = 250 m, B–E = 10 m.
Fig. 23. Harknessia pilularis (ex-type CPC 33218). A. Conidiomata on OA. B, C. Conidiogenous cells giving rise to conidia. D.
Conidia. Scale bars: A = 250 m, B–D = 10 m.
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Fig. 24. Huntiella spp. A–D. Disease symptoms. A. Eucalyptus trees artificially wounded to trap Huntiella spp. B. Lesion associated
with inoculation with Huntiella sumatrana on an Acacia mangium stem. C. Discolouration of wood associated with artificially induced
wound on the stem of a Eucalyptus tree from which Huntiella spp. were isolated. D. Blue-stained Eucalyptus wood associated with
Huntiella infection. E–H, L. Sexual morphs. E. Ascomata on 2 % MEA in various developmental stage from young (paler) to mature
(darker). F. Mature ascomata with fresh ascospore droplets at the tip of necks and ostiolar neck with a disc-like base (arrow). G, H.
Young ascoma showing developing of conical spines (G) and ascomatal hyphae (H). L. Ascospores. I–K. Asexual morphs. I.
Tubular-form conidiogenous cell producing barrel-shaped conidia. J. Flask-shaped conidiogenous cells producing rectangularshaped conidia. K. Rectangular-shaped conidia. E, F, H, K. Huntiella omanensis (CMW 11056). G, L. Huntiella moniliformis (CMW
36908). I, J. Huntiella abstrusa (CMW 21092). Scale bars: E = 500 µm; F = 250 µm; G, H = 50 µm; I–L = 10 µm.
Fig. 25. A maximum likelihood (ML) phylogram constructed using the combined dataset of the 60S (370 bp), LSU (813 bp), ITS (393
bp), mcm7 (589 bp) and tub2 (433 bp) gene regions of Huntiella spp. ML analyses were performed using PhyML with Smart Model
Selection Online, selecting the GTR substitution model. Bootstrap values of 70 % or more are indicated above the branches. The
newly described species, Hu. abstrusa, is indicated in bold face. Trees and alignments were submitted to TreeBASE: S20860.
Fig. 26. Huntiella abstrusa (ex-type CBS 142243 = CMW 21092). A. Ascomata on host substrate. B. Ascospore droplets of cream to
yellow-coloured at the tip of ascomatal neck and ascomata embedded in aerial hyphae producing asexual structures. C. Colony on
2 % MEA grown in the dark at 30 °C for 3 d. D. Mature ascoma with a disk-like base (arrow). E. Base of young ascoma (1 wk) with
conical spines (arrow) (image taken in bright field). F. Ascospores. G. Two types of conidia: rectangular-shaped and barrel-shaped
conidia. H. Flask-shaped conidiogenous cells and rectangular-shaped conidia. I. Tubular-shaped conidiogenous cell and barrelshaped conidia. Scale bars: B = 500 µm; D = 100 µm; E = 50 µm; F–I = 10 µm.
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Fig. 27. A–H. Macgarvieomyces juncicola (CBS 610.82). A. Colony sporulating on OA. B–G. Conidiophores and conidia forming on
SNA. H. Conidia. Scale bars = 10 m. All pictures taken from Klaubauf et al. (2014).
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Fig. 28. RAxML phylogram obtained from the combined ITS (548 bp), act (375 bp), cal (579 bp) and rpb1 (1011 bp) sequence
alignment of members of Pyriculariaceae. The tree was rooted to Bussabanomyces longisporus CBS 125232. 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 were listed in Klaubauf et al. (2014). T, ET and NT indicate
ex-type, ex-epitype and ex-neotype strains, respectively. TreeBASE: S21899.
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Fig. 29. A–H. Macgarvieomyces luzulae (ex-epitype CBS 143401). A. Disease symptoms on leaves of Luzula sylvatica. B. Colony
sporulating on OA. C–H. Conidiophores and conidia forming on SNA. I. Conidia. Scale bars = 5 m.
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Fig. 30. Metulocladosporiella spp. A. Disease symptoms on Musa sp (indicated by the arrows). B–K. Asexual morph. B–D.
Macronematous conidiophores. B. Metulocladosporiella musicola (CBS 121396). C. Metulocladosporiella musicola (ex-type CBS
110960). D. Metulocladosporiella musae (CPC 33937). E–G. Conidiogenous apparatus. E, F. Metulocladosporiella musae (CPC
33937). G. Metulocladosporiella musicola (ex-type CBS 110960). H. Lobed bases of macronematous conidiophore of
Metulocladosporiella musae (CPC 33937). I–K. Micronematous conidiophores. I, J. Metulocladosporiella musae (CPC 33937). K.
Metulocladosporiella musicola (CBS 121396). Scale bars: 10 m.
Fig. 31. RAxML phylogram obtained from the combined ITS (667 bp), cal (524 bp) and tef1 (454 bp) sequence alignment of taxa
belonging to Metulocladosporiella. The tree was rooted to Cladosporium tenuissimum CBS 125995ET. 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 indicated in Table 10. T and ET indicate ex-type and ex-epitype
strains, respectively. TreeBASE: S21899.
Fig. 32. Metulocladosporiella chiangmaiensis (ex-type CBS 143918). A. Colony on PDA. B. Colony on OA. C–E. Macronematous
conidiophores. F. Conidiogenous apparatus. G. Lobed bases of macronematous conidiophore. H–K. Micronematous conidiophores.
Scale bars: C= 50 m; D = 20 m; all others 10 m.
Fig. 33. Metulocladosporiella malaysiana (ex-type CBS 143919). A. Colony on PDA. B. Colony on OA. C–F. Macronematous
conidiophores. G. Conidiogenous apparatus. H. Lobed bases of macronematous conidiophore. I–J. Micronematous conidiophores.
Scale bars: C, D = 20 m; all others 10 m.
Fig. 34. Metulocladosporiella musigena (ex-type CBS 143920). A. Colony on PDA. B. Colony on OA. C–F. Macronematous
conidiophores. G–I. Conidiogenous apparatus. J, K. Lobed bases of macronematous conidiophore. L, M. Micronematous
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conidiophores. Scale bars: C–F = 20 m; G–M = 10 m; K applies to J, K.
Fig. 35. Metulocladosporiella songkhramensis (ex-type CBS 143921). A. Colony on PDA. B. Colony on OA. C. Macronematous
conidiophores. D–F. Conidiogenous apparatus. G. Lobed bases of macronematous conidiophore. H–J. Micronematous
conidiophores. Scale bars: C = 20 m; all others 10 m.
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Fig. 36. Microdochium spp. A–F. Sexual morph of Microdochium seminicola (ex-type CBS 139951). A. Colony overview. B.
Ascomata. C–E. Asci. F. Ascospores. G–U. Asexual morphs. G, H. Sporodochium G. Microdochium phragmites (CBS 423.78). H.
Microdochium lycopodinum (CBS 109399). I–N. Conidiophores and conidiogenous cells. I. Microdochium neoqueenslandicum (extype CBS 108926). J. Microdochium citrinidiscum (ex-type CBS 109067). K. Microdochium seminicola (ex-type CBS 139951). L.
Microdochium phragmites (ex-epitype CBS 285.71). M. Microdochium phragmites (CBS 423.78). N. Microdochium fisheri (ex-type
CBS 242.91). O–U. Conidia. O. Microdochium seminicola (ex-type CBS 139951). P. Microdochium lycopodinum (CBS 109399). Q.
Microdochium fisheri (ex-type CBS 242.91). R. Microdochium neoqueenslandicum (ex-type CBS 108926). S. Microdochium
phragmites (CBS 423.78). T. Microdochium phragmites (ex-epitype CBS 285.71). U. Microdochium citrinidiscum (ex-type CBS
109067). V, W. Chlamydospores. V. Microdochium bolleyi (CPC 29379). W. Microdochium trichocladiopsis (ex-type CBS 623.77).
Scale bars H = 100 µm; G = 50 µm; others = 10 µm.
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Fig. 37. RAxML phylogram obtained from the combined ITS (618 bp), LSU (838 bp), tub2 (689 bp) and rpb2 (858 bp) sequence
alignment of all the accepted species of Microdochium. The tree was rooted to Thamnomyces dendroidea CBS 123578 and Xylaria
polymorpha MUCL 49884. The novel species described in this study is shown in bold. RAxML bootstrap support (BS) values above
70 % are shown in the nodes. GenBank accession numbers are indicated in Table 11. T and ET, ex-type and ex-syntype strains,
respectively. TreeBASE: S21899.
Fig. 38. Microdochium novae-zelandiae (ex-type CPC 29376) A. Colony overview. B, C. Sporodochium overview. B. From aerial
mycelium. C. From agar surface. D–G. Conidiogenous cells and conidia. H, I. Hyphae and conidia. J. Conidia. Scale bars D–J = 10
µm.
Fig. 39. Oculimacula spp. A–G. Disease symptoms. A, B. Eyespot lodging. C–E. Eyespots. F. Whiteheads of wheat. G. Apothecia
of Oculimacula gamsii on wheat stubble. H–K. Sexual morphs of Oculimacula yallundae. H, I. Ascomata. J. Section through the
ascoma showing ascal layer. K. Ascus and ascospores. L–O. Asexual morphs. L. Conidia and conidiogenous cells of Oculimacula
gamsii. M. Conidial hila and conidiogenous cell of Oculimacula yallundae. N. Conidial hila and conidiogenous cell of Oculimacula
gamsii. O. Conidial hila and conidiogenous cell of Oculimacula anguoides. Scale bars: I = 100 µm; J, K = 10 µm; L = 5 µm; M–O = 1
µm.
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Fig. 40. RAxML phylogram obtained from the combined ITS (631 bp) and tef1 (575 bp) sequence alignment of taxa belonging to
Oculimacula. The tree was rooted to Cadophora melinii. The new combination 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 are indicated in Table 12. T and NT indicate ex-type and ex-neotype strains, respectively. TreeBASE: S21899.
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Fig. 41. Paraphoma spp. A–E. Disease symptoms. A–B. Crown discolouration caused by Paraphoma vinacea (ex-type BRIP
63684). C. Water-soaked and necrotic leaf lesions caused by Paraphoma chlamydocopiosa (ex-type BRIP 65168). D. Marginal leaf
chlorosis caused by Paraphoma pye on pyrethrum leaf (ex-type BRIP 65169). E–O. Asexual morphs. E. Conidiomata on SNA of
Paraphoma fimeti (ex-neotype CBS 170.70). F. Conidiomata of Paraphoma vinacea (ex-type BRIP 63684). G. Conidiomatal wall of
Paraphoma vinacea (ex-type BRIP 63684). H. Ostiolar zone of Paraphoma vinacea (ex-type BRIP 63684). I–L. Condiogenous cells.
I, J. Paraphoma dioscoreae (ex-type CBS 135100). K, L. Paraphoma fimeti (ex-neotype CBS 170.70). M, N. Conidia. M.
Paraphoma dioscoreae (ex-type CBS 135100). N. Paraphoma fimeti (ex-neotype CBS 170.70). O. Chlamydospores of Paraphoma
vinacea (ex-type BRIP 63684). Scale bars: F = 100 µm; G, H = 20 µm; I, K, M–O = 10 µm; I applies to I, J; K applies to K, L.
Pictures B, F, G taken from Moslemi et al. (2016); C, D from Moslemi et al. (2017); I, J, M from Quaedvlieg et al. (2013).
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Fig. 42. Maximum likelihood PhyML combined phylogenetic tree of Paraphoma spp. inferred from ITS (680 bp), tef1 (550 bp) and
tub2 (350 bp) using a GTR substitution model. Highest log likelihood -3812.4179. Bootstrap support values less than 65 % were
removed. Scale bar indicated expected changes per site. The tree was rooted to Neosetophoma samarorum CBS 138.96. GenBank
accession numbers are indicated in Table 13. TreeBASE: S22303.
Fig. 43. Morphological structures of Phaeoacremonium spp. A–K. Asexual morphs. A. Conidiophores branched. B. Conidiophores
unbranched. C. Percurrent rejuvenation of conidiophore. D. Mycelium showing prominent exudate droplets observed as warts. E.
Conidiophore with polyphialides. F. Type III phialides. G. Type II phialide. H. Type I phialide. I. Conidia. J. Conidiophore. K. Type II
phialide. L–T. Sexual morph. L, M. Ascomata on canes of Vitis vinifera. N. Longitudinal section through ascoma. O. One
paraphyses. P, Q. Asci attached to ascogenous hyphae. R. Asci. S. Ascospores. A, B, D, F, G, I. Phaeoacremonium parasiticum
(ex-type CBS 860.73). C. Phaeoacremonium hispanicum (ex-type CBS 123910). E. Phaeoacremonium amygdalinum (ex-type CBS
128570). H. Phaeoacremonium minimum (ex-type CBS 246.91). L–S. Phaeoacremonium minimum (holotype CBS 17463). Scale
bars: A–I = 10 µm; A applies to A, B, D, F–I; C applies to C, E; L, M = 200 µm; N = 100 µm; O, P = 10 µm; P applies to P–S.
Fig. 44. Bayesian consensus tree of the genus Phaeoacremonium as estimated from the combined act (~260 bp) and tub2 (~680
bp) regions. Bayesian posterior probability values and bootstrap support percentages are shown at the nodes. Support values of
less than 0.7 posterior probability and 70 % bootstrap are not shown. Jattaea algeriensis, Calosphearia africana and Pleurostoma
richardsiae were used as outgroups. GenBank accession numbers are listed in Spies et al. (2018). T indicates ex-type strains.
TreeBASE: S22407.
ACCEPTED MANUSCRIPT
Fig. 45. Phaeoacremonium pravum (ex-type CBS 142686). A–C. Eight-d-old colonies on MEA (A), PDA (B) and OA (C). D.
Subcylindrical type I phialides with funnel-shaped collarettes (arrows). E, F. Type III phialides. G, J. Branched conidiophores with
crooked type II phialides. H. Elongate ampulliform type I phialide. I. Type I phialide with conidia borne in a slimy head. K. Crooked
elongate ampulliform type I phialide with a funnel-shaped collarette showing lateral vegetative proliferation (arrow). Scale bar: K =
10 µm; K applies to D–K.
RI
PT
Fig. 46. Phyllosticta spp. A−E. Disease symptoms. A. Aloe with dead leaf tips that harbour Phyllosticta aloeicola. B. Symtoms on
Citrus maxima caused by Phyllosticta citrimaxima. D. Symptomatic leaf of Cussonia sp. caused by Phyllosticta cussoniae. E.
Symptoms on lemon leaf caused by Phyllosticta sp. F−H. Sexual morphs. F, G. Asci and ascospores of Phyllosticta abieticola (extype CBS 112067). H. Ascospores of Phyllosticta capitalensis (ex-epitype CBS 128856). I−U. Asexual morphs. I. Conidiomata
sporulating on OA of Phyllosticta cussoniae (ex-epitype CPC 14873). J. Vertical section through conidioma of Phyllosticta
rhaphiolepidis (ex-type MUCC 432). K. Conidiomatal wall of textura angularis of Phyllosticta rhaphiolepidis (ex-type MUCC 432). L.
Conidioma with ostiole (arrowed) of Phyllosticta cordylinophila (ex-neotype CPC 20261). M, N. Conidiogenous cells giving rise to
conidia. M. Phyllosticta foliorum (ex-neotype CBS 447.68). N. Phyllosticta capitalensis (ex-epitype CBS 128856). O−Q. Conidia. O.
Phyllosticta aloeicola (CPC 20677). P. Phyllosticta podocarpicola (ex-type CBS 728.79). Q. Phyllosticta capitalensis (ex-epitype
CBS 128856). R, S. Appressoria of Phyllosticta mangifera-indica (ex-type CPC 20274). T, U. Spermatia. T. Phyllosticta cussoniae
(ex-epitype CPC 14873). U. Phyllosticta leucothoicola (ex-type MUCC 553). Scale bars: I = 25 µm; all others = 10 µm. Pictures N, Q
taken from Glienke et al. (2011); all the other pictures taken from Wikee et al. (2013b).
M
AN
US
C
Fig. 47. RAxML phylogram obtained from the combined ITS (492 bp), act (291 bp), gapdh (629 bp) and tef1 (341 bp) sequence
alignment of all the accepted species of Phyllosticta. The tree was rooted to Peyronellaea obtusa CMW8232. 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 indicated in Table 15. T, ET, IsoT, IsoET and NT indicate extype, ex-epitype, ex-isotype, ex-isoepitype and ex-neotype strains, respectively. TreeBASE: S21899.
Fig. 48. Phyllosticta iridigena (ex-type CBS 143410). A. Conidiomata sporulating on SNA. B–E. Conidiogenous cells giving rise
to conidia. F. Conidia. Scale bars A= 200 m, B–F= 10 m.
Fig. 49. Phyllosticta persooniae (ex-type CBS 143409). A. Conidiomata sporulating on OA. B, C. Conidiogenous cells giving
rise to conidia. D. Conidia. A= 200 m, B–D= 10 m.
Fig. 50. Proxipyricularia zingiberis (ex-epitype CBS 133594). A–C. Conidiophores and conidia. D. Conidia. Scale bars = 5 m.
Fig. 51. Pyriculariomyces asari (ex-type CBS 141328). A. Ascomata on host tissue. B. Section of ascoma. C–E. Asci and
ascospores. F. Conidiophores on SNA. G, H. Conidiophores and conidia. I. Conidia. Scale bars: B = 100 m; all others = 10 m; C
applies to C, D; G applies to G, I. Pictures A, B, D, E, G, H taken from Crous et al. (2016b).
EP
TE
D
Fig. 52. Pyricularia spp. A–C. Leaf spots of rice caused by Pyricularia oryzae. D–H. Sexual morph of Pyricularia oryzae. D, E.
Crossing of different strains of Pyricularia oryzae to produce the sexual morph. F. Ascoma. G. Asci. H. Germinating ascospore. I–U.
Asexual morph. I, J. Sporulation on sterile barley seed on SNA. I. Pyricularia grisea (BR0029). J. Pyricularia graminis-tritici (ex-type
URM7380). K–O. Conidiophores and conidia. K, L. Pyricularia ctenantheicola (GR0002). M, N. Pyricularia graminis-tritici (ex-type
URM7380). O. Pyricularia oryzae (URM7369). P. Pyricularia oryzae (BF0028). Q–S. Conidia. Q. Pyricularia ctenantheicola
(GR0002). R. Pyricularia oryzae (URM7369). S. Pyricularia graminis-tritici (ex-type URM7380). T. Macroconidia of Pyricularia grisea
(BR0029) (arrows indicate apical marginal frill, which is a remnant of the apical mucoid cap). U. Microconidia of Pyricularia grisea
(BR0029). Scale bars: F = 50 m; others = 10 m. Pictures D, E, G taken by Dounia Saleh, CIRAD; F, H by Didier Tharreau,
CIRAD; I, K, L, P, Q, T from Klaubauf et al. (2014); J, M–O, R, S from Castroagudín et al. (2016).
AC
C
Fig. 53. Stenocarpella spp. A. Zea mays infected with Stenocarpella maydis. B–K. Asexual morphs. B. Conidiomata with exuding
conidial mass on pine needle agar of Stenocarpella maydis (ex-epitype CBS 117559). C. Hyaline layer of conidiogenous cells giving
rise to brown conidial mass of Stenocarpella macrospora (CPC 11863). D–F. Conidiogenous cells giving rise to conidia. D, E.
Stenocarpella macrospora (CPC 11863). F. Stenocarpella maydis (ex-epitype CBS 117559). G–I. Conidia. G, H. Stenocarpella
macrospora (CPC 11863). I. Stenocarpella maydis (ex-epitype CBS 117559). J. Conidiogenous cells giving rise to beta conidia of
Stenocarpella macrospora (CPC 11863). K. Beta conidia of Stenocarpella macrospora (CPC 11863). Scale bars: 10 m. All
pictures except for A taken from Lamprecht et al. (2011).
Fig. 54. A–I. Utrechtiana arundinacea (ex-epitype CPC 33994). A. Leaf spot on Phragmites sp. B–E. Macroconidiophores bearing
macroconidia. G–H. Microconidiophores bearing microconidia. I. Microconidia. J–S. Utrechtiana roumeguerei (ex-type CBS
128780). J. Leaf spot on Phragmites australis. K. Close-up of conidiophores on leaf surface. L–P. Conidiophores bearing conidia.
Q. Germinating conidium. R, S. Conidia. Scale bars = 10 m. Pictures J–S taken from Klaubauf et al. (2014).
Fig. 55. Wojnowiciella spp. A–F. Conidiomata overview. A. Wojnowiciella leptocarpi (ex-type CBS 116584). B, E. Wojnowiciella
dactylidis (CPC 30353). C. Wojnowiciella cissampeli (ex-type CBS 141297). D. Wojnowiciella eucalypti (ex-type CBS 139904). F.
Wojnowiciella dactylidis (CPC 32741). G, H. Hand section of the conidiomata with hyaline conidiogenous cells and dark brown
conidia of Wojnowiciella leptocarpi (ex-type CBS 116584). I–L. Conidiogenous cells. I, J. Wojnowiciella cissampeli (ex-type CBS
141297). K. Wojnowiciella eucalypti (ex-type CBS 139904). L. Wojnowiciella leptocarpi (CBS 116585). M–Q. Macroconidia. M, N.
Wojnowiciella leptocarpi (ex-type CBS 116584). O. Wojnowiciella cissampeli (ex-type CBS 141297). P. Wojnowiciella dactylidis
(CPC 32741). Q. Wojnowiciella dactylidis (CPC 30353). R. Microconidia of Wojnowiciella eucalypti (ex-type CBS 139904). Scale
bars: E–G = 50 m; H = 20 m; others = 10 m. Photos taken from Crous et al. (2015d, 2016b).
Fig. 56. RAxML phylogram obtained from the combined ITS (742 bp), LSU (885 bp), rpb2 (1029 bp) and tef1 (998 bp) sequence
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
US
C
RI
PT
alignment of all the accepted species of Wojnowiciella. The tree was rooted to Phaeosphaeria caricis CBS 120249 and Septoriella
hirta CBS 536.77. RAxML bootstrap support (BS) values above 70 % are shown in the nodes. GenBank accession numbers are
indicated in Table 21. T indicates ex-type strains. TreeBASE: S21899.
ACCEPTED MANUSCRIPT
Table 1. DNA barcodes of accepted Allantophomopsiella sp.
Species
Isolate1
GenBank accession
numbers2
ITS
rpb2
KJ663825
KJ663905
Reference
AC
C
EP
TE
D
M
AN
US
C
RI
PT
Allantophomopsiella CBS 320.53ET
Crous
et
al.
pseudotsugae
(2014b)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. ET indicates exepitype strain.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial RNA polymerase II
largest subunit gene.
ACCEPTED MANUSCRIPT
Table 2. DNA barcodes of accepted Apoharknessia spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
References
GenBank accession numbers2
ITS
cal
tub2
Apoharknessia eucalypti CBS 142518T
MG934432 MG934510 MG934505 Present study
T
Ap. eucalyptorum
CBS 142519
KY979752 KY979867 KY979919 Crous et al. (2017a)
MG934511 MG934506 Crous et al. (2012c), present study
Ap. insueta
CBS 111377ET JQ706083
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T and ET indicate ex-type and ex-epitype strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; cal: partial calmodulin gene; tub2: partial β-tubulin gene.
Species
ACCEPTED MANUSCRIPT
Table 3. DNA barcodes of accepted Cylindrocladiella spp.
2
GenBank accession numbers
LSU
his3
tef1
MH111393
-
CBS 143794
ITS
MH111383
CBS 143793
MH111385
-
-
MH111395
CBS 143795
MH111384
-
-
MH111394
T
tub2
MH111388
RI
PT
Cylindrocladiella
addiensis
1
Isolates
MH111389
Present study
T
JN100624
AF220952
JN099222
JN099249
JN098932
AY793509
JN099060
MF444940
JN099087
JN098747
MF444926
AY793471
C. clavata
C. cymbiformis
C. elegans
CBS 129564
T
CBS 129553
T
CBS 338.92
T
JN099095
JN099103
AY793444
JN099135
JN099143
JN099201
JN098858
JN098866
AY793512
JN098974
JN098988
JN099039
JN098752
JN098753
AY793474
C. ellipsoidea
C. hahajimaensis
C. hawaiiensis
C. horticola
C. humicola
C. infestans
CBS 129573
T
MAFF 238172
T
CBS 129569
T
CBS 142784
T
CBS 142779
T
CBS 111795
T
JN099094
JN687561
JN100621
MF444911
MF444906
AF220955
JN099134
JN099219
JN099199
JN098857
JN098929
AY793513
JN098973
JN687562
JN099057
MF444938
MF444933
JN099037
JN098757
JN098761
MF444924
MF444919
AF320190
C. kurandica
C. lageniformis
CBS 129577
T
CBS 340.92
T
JN100646
AF220959
JN099245
JN099165
JN098953
AY793520
JN099083
JN099003
JN098765
AY793481
C. lanceolata
C. lateralis
C. longiphialidica
C. longistipitata
CBS 129566
T
CBS 142788
T
CBS 129557
T
CBS 116075
T
JN099099
MF444914
JN100585
AF220958
JN099139
JN099264
JN099155
JN098862
JN098851
AY793546
JN098978
MF444942
JN098966
JN098993
JN098789
MF444928
JN098790
AY793506
C. microcylindrica
CBS 111794
T
AY793452
JN099203
AY793523
JN099041
AY793483
TE
D
M
AN
U
CBS 129567
T
CBS 142786
IMI 346845
EP
Present study
Present study
C. australiensis
C. brevistipitata
C. camelliae
AC
C
References
MH111390
SC
Species
Lombard et al. (2012)
Lombard et al. (2017)
Van Coller et al. (2005), Lombard et al.
(2012)
Lombard et al. (2012)
Lombard et al. (2012)
van Coller et al. (2005), Lombard et al.
(2012)
Lombard et al. (2012)
Inderbitzin et al. (2012)
Lombard et al. (2012)
Lombard et al. (2017)
Lombard et al. (2017)
Schoch et al. (2000), Crous et al. (2001),
van Coller et al. (2005), Lombard et al.
(2012)
Lombard et al. (2012)
Schoch et al. (2000), van Coller et al.
(2005), Lombard et al. (2012)
Lombard et al. (2012)
Lombard et al. (2017)
Lombard et al. (2012)
Schoch et al. (2000), van Coller et al.
(2005), Lombard et al. (2012)
van Coller et al. (2005), Lombard et al.
ACCEPTED MANUSCRIPT
JN100588
JN100603
AF220963
JN099178
JN099195
JN099212
JN098895
JN098910
AY793525
JN099016
JN099033
JN099050
CBS 143792
T
MH111387
-
-
MH111397
CBS 143791
MH111386
-
-
MH111396
CBS 114943
T
CBS 152.91
T
CBS 486.77
C. nauliensis
MH111392
(2012)
Lombard et al. (2012)
Lombard et al. (2012)
Schoch et al. (2000), van Coller et al.
(2005), Lombard et al. (2012)
Present study
MH111391
Present study
JN098794
JN098800
AY793485
RI
PT
T
C. natalensis
C. nederlandica
C. novazelandica
Schoch et al. (2000), van Coller et al.
(2005), Lombard et al. (2012)
T
C. peruviana
IMUR 1843
AF220966 JN099266 AY793540 JN098968 AY793500 Schoch et al. (2000), van Coller et al.
(2005), Lombard et al. (2012)
T
C. pseudocamelliae
CBS 129555
JN100577 JN099256 JN098843 JN098958 JN098814 Lombard et al. (2012)
T
C. pseudohawaiiensis
JN099128 JN099174 JN098890 JN099012 JN098819 Lombard et al. (2012)
CBS 210.94
T
C. pseudoinfestans
AF220957 JN099166 AY793548 JN099004 AY793508 Schoch et al. (2000), van Coller et al.
CBS 114531
(2005), Lombard et al. (2012)
T
C. pseudoparva
CBS129560
JN100620 JN099218 JN098927 JN099056 JN098824 Lombard et al. (2012)
T
C. queenslandica
CBS 129574
JN099098 JN099098 JN098861 JN098977 JN098826 Lombard et al. (2012)
T
C. reginae
MF444909
MF444936 MF444922 Lombard et al. (2017)
CBS 142782
T
C. stellenboschensis
JN100615 JN099213 JN098922 JN099051 JN098829 Lombard et al. (2012)
CBS 110668
T
C. terrestris
CBS 142789
MF444915
MF444943 MF444929 Lombard et al. (2017)
T
C. thailandica
JN100582 JN099261 JN098848 JN098963 JN098834 Lombard et al. (2012)
CBS 129571
T
C. variabilis
CBS 129561
JN100643 JN099242 JN098950 JN099080 JN098719 Lombard et al. (2012)
T
C. viticola
AY793468 JN099226 AY793544 JN099064 AY793504 van Coller et al. (2005), Lombard et al.
CBS 112897
(2012)
T
C. vitis
CBS 142517
KY979751 KY979806
KY979891 KY979918 Crous et al. (2017a)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane,
U.K; IMUR: Institute of Mycology, University of Recife, Recife, Brazil; MAFF: Genetic Resources Centre, National Agriculture and Food Research
T
Organization (NARO), NARO Genebank, Ibaraki, Japan. indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; his3: partial histone H3 gene; tef1: partial translation elongation factor 1-alpha gene; tub2:
partial β-tubulin gene.
AY793526
JN099009
D
TE
AY793486
SC
JN099171
M
AN
U
AF220964
EP
CBS 114524
AC
C
C. parva
ACCEPTED MANUSCRIPT
Table 5. DNA barcodes of accepted Dichotomophthora spp.
Species
Isolates1
GenBank accession numbers2
gapdh
rpb2
References
RI
PT
ITS
CPC 33016T
Present study
CBS 149.94T
Present study
CBS 145.57T
Present study
CBS 584.71
Present study
CBS 585.71
Present study
CBS 518.78
Present study
CBS 132.81
Present study
Present study
Di. portulacae
CBS 174.35PT
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at the
Westerdijk Fungal Biodiversity Institute. T and PT indicate ex-type and paratype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; rpb2:
partial RNA polymerase II largest subunit gene.
AC
C
EP
TE
D
M
AN
U
SC
Dichotomophthora basellae
Di. brunnea
Di. lutea
ACCEPTED MANUSCRIPT
Table 6. DNA barcodes of accepted Gaeumannomyces spp.
Gaeumannomyces amomi
Isolates1
GenBank accession numbers2
CBS 109354T
ITS
rpb1
tef1
AY265318
-
KX306679
References
RI
PT
Species
Bussaban
et
al.
(2005),
Hernández-Restrepo et al. (2016b)
G. ellisiorum
G. floridanus
G. fusiformis
CPC 26258
KX306486
T
KX306490
CBS 141377
T
CBS 387.81
KM484835
CBS 141378
T
KX306491
CBS 141379
T
T
CPC 26057
G. graminicola
CBS 352.93T
Klaubauf
et
al.
(2014),
Hernández-Restrepo et al. (2016b)
KX306683
Hernández-Restrepo et al. (2016b)
KX306622
KX306688
Hernández-Restrepo et al. (2016b)
KX306625
KX306691
Hernández-Restrepo et al. (2016b)
KM485051
KX306692
Klaubauf
et
al.
(2014),
Hernández-Restrepo et al. (2016b)
KX306626
KX306693
Hernández-Restrepo et al. (2016b)
KX306492
KX306627
KX306694
Hernández-Restrepo et al. (2016b)
KX306493
KX306628
KX306695
Hernández-Restrepo et al. (2016b)
KM485050
KX306697
Klaubauf
KM484834
AC
C
G. glycinicola
KX306619
KX306681
SC
KX306480
ET
CBS 141387
M
AN
U
G. californicus
T
KM485053
D
G. avenae
KM484837
TE
G. australiensis
CBS 903.73T
EP
G. arxii
et
al.
(2014),
Hernández-Restrepo et al. (2016b)
G. graminis
CPC 26020
KX306498
KX306633
KX306701
Hernández-Restrepo et al. (2016b)
G. hyphopodioides
CBS 350.77T
KX306506
KM009192
KM009204
Hernández-Restrepo
et
al.
(2016b), Luo et al. (2014)
G. oryzicola
CBS 141390T
KX306516
KX306646
KX306717
Hernández-Restrepo et al. (2016b)
G. oryzinus
CBS 235.32
JX134669
KM485049
JX134695
Klaubauf et al. (2014), Luo &
ACCEPTED MANUSCRIPT
CBS 296.53T
KM484845
KM485061
KM009206
Zhang (2013)
Klaubauf et al. (2014)
G. setariicola
CBS 141394T
KX306524
KX306654
KX306725
Hernández-Restrepo et al. (2016b)
G. tritici
CBS 905.73
KM484841
KM485057
KX306731
RI
PT
G. radicicola
Klaubauf
et
al.
(2014),
Hernández-Restrepo et al. (2016b)
CBS 141400T
KX306543
KX306670
KX306746
Hernández-Restrepo et al. (2016b)
G. wongoonoo
BRIP 60376A
KP162137
-
-
Wong (2002)
1
SC
G. walkeri
AC
C
EP
TE
D
M
AN
U
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. T, ET and A indicate extype, ex-epitype and authentic strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb1: partial RNA polymerase II largest subunit gene; tef1: partial
elongation factor gene.
ACCEPTED MANUSCRIPT
Table 7. DNA barcodes of accepted Harknessia spp.
Ha. platyphyllae
Ha. proteae
References
RI
PT
SC
M
AN
U
Ha. ellipsoidea
Ha. eucalypti
Ha. eucalyptorum
Ha. fusiformis
Ha. gibbosa
Ha. globispora
Ha. hawaiiensis
Ha. ipereniae
Ha. karwarrae
Ha. kleinzeeina
Ha. leucospermi
Ha. malayensis
Ha. molokaiensis
Ha. pellitae
Ha. pilularis
D
CBS 137228ET
CBS 132119T
CBS 142539T
CBS 142541T
CBS 142540T
CBS 143913T
CBS 111829T
CBS 142538T
CPC 33289T
CBS 143914T
CPC 30174
CBS 132121T
CBS 342.97
CBS 111115T
CBS 110785T
CBS 120033T
CBS 111578T
CBS 114811
CBS 120030T
CBS 115648
CPC 16277T
CBS 775.97T
CBS 142544T
CBS 114877T
CBS 142543T
CPC 33218T
CPC 33356
CBS 142542T
CBS 136426T
TE
Harknessia arctostaphyli
Ha. australiensis
Ha. banksiae
Ha. banksiae-repens
Ha. banksiigena
Ha. bourbonica
Ha. capensis
Ha. communis
Ha. corymbiae
Ha. cupressi
GenBank accession number2
ITS
cal
tub2
KJ152781
KJ179923
JQ706085
JQ706171
JQ706130
KY979782 KY979872 KY979938
KY979785 KY979875 KY979940
KY979784 KY979874 MG934433 MG934512 AY720719 AY720782 AY720751
KY979778 KY979868 MG934434 MG934513 MG934507
MG934435 MG934514 MG934436 MG934515 JQ706087
JQ706173
JQ706132
AY720745 AY720808 AY720777
AY720747 AY720810 AY720779
AY720721 AY720784 AY720753
EF110615
JQ706182
JQ706142
AY720722 AY720785 AY720754
AY720723 AY720786 AY720755
EF110614
JQ706192
JQ706151
AY720748 AY720811 AY720780
JQ706108
JQ706193
JQ706152
AY720727 AY720790 AY720759
KY979789 KY979879 KY979941
AY720749 AY720812 AY579335
KY979788 KY979878 MG934438 MG934517 MG934508
MG934439 MG934518 MG934509
KY979787 KY979877 KF777162
-
EP
Isolates1
AC
C
Species
Moreno-Rico et al. (2014)
Crous et al. (2012c)
Crous et al. (2017a)
Crous et al. (2017a)
Crous et al. (2017a)
Present study
Lee et al. (2004)
Crous et al. (2017a)
Present study
Present study
Present study
Crous et al. (2012c)
Lee et al. (2004)
Lee et al. (2004)
Lee et al. (2004)
Crous et al. (2007), (2012c)
Lee et al. (2004)
Lee et al. (2004)
Crous et al. (2007), (2012c)
Lee et al. (2004)
Crous et al. (2012c)
Lee et al. (2004)
Crous et al. (2017a)
Lee et al. (2004), Mostert et al. (2005)
Crous et al. (2017a)
Present study
Present study
Crous et al. (2017a)
Crous et al. (2013)
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
Ha. protearum
CBS 112618T
AY720732 AY720795 AY720764 Lee et al. (2004)
JQ706111
JQ706196
JQ706155
Crous et al. (2012c)
Ha. pseudohawaiiensis
CBS 132124T
Ha. ravenstreetina
CBS 132125T
JQ706112
JQ706197
JQ706156
Crous et al. (2012c)
IsoT
Ha. renispora
CBS 153.71
AY720737 AY720800 AY720769 Lee et al. (2004)
Ha. rhabdosphaera
CBS 122373
JQ706118
JQ706201
JQ706161
Crous et al. (2012c)
Ha. spermatoidea
CBS 132127ET
JQ706120
JQ706203
JQ706163
Crous et al. (2012c)
Ha. syzygii
CBS 111124T
AY720738 AY720801 AY720770 Lee et al. (2004)
Ha. uromycoides
CBS 110729
AY720739 AY720802 AY720771 Lee et al. (2004)
T
Ha. viterboensis
CBS 115647
AY720740 AY720803 AY720772 Lee et al. (2004)
JQ706122
JQ706205
JQ706165
Crous et al. (2012c)
Ha. weresubiae
CBS 132128ET
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CPC: Culture collection of Pedro Crous, housed at the
Westerdijk Fungal Biodiversity Institute. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; cal: partial calmodulin gene; tub2: partial β-tubulin gene.
ACCEPTED MANUSCRIPT
Table 8. DNA barcodes of accepted Huntiella spp.
Isolates1
Huntiella abstrusa
Hu. bhutanensis
CBS 142243T
CBS 114289T
Hu. ceramica
Hu. chinaeucensis
Hu. cryptoformis
Hu. decipiens
Hu. inquinans
Hu. microbasis
Hu. moniliformis
Hu. moniliformopsis
CBS 122299T
CBS 127185T
CBS 131279T
CBS 129736T
CBS 124388T
CBS 124013T
CBS 118127
CBS 109441T
GenBank accession numbers2
ITS
tub2
mcm7
tef1
KY913291 KY913290 KY913289 AY528952 AY528962 KM495412 AY528962a
NR119506
EU245022 EU244994 KM495485 EU244926b
JQ862729
JQ862717
KM495416 JQ862741c
KC691464 KC691488 KC691512c
HQ203216 HQ203233 KM495422 HQ236435c
EU588587 EU588666 KM495436 EU588674a
EU588593 EU588672 KM495442 EU588680a
FJ151422
FJ151456
KM495443 FJ151478a
AY528998 AY528987 KM495444 AY529008a
References
Present study
Van Wyk et al. (2004), De Beer et al. (2014)
RI
PT
Species
AC
C
EP
TE
D
M
AN
U
SC
Heath et al. (2009), De Beer et al (2014)
Chen et al. (2013), De Beer et al. (2014)
Mbenoun et al. (2014)
Kamgan Nkuekam et al. (2013), De Beer et al.(2014)
Tarigan et al. (2010), De Beer et al (2014)
Tarigan et al. (2010); De Beer et al. (2014)
Van Wyk et al. (2011), De Beer et al. (2014)
Yuan & Mohammed (2002), Van Wyk et al. (2004),
De Beer et al. (2014)
Hu. oblonga
CBS 122291T
EU245019 EU244991 KM495447 EU244951b
Heath et al. (2009), De Beer et al. (2014)
Hu. omanensis
CBS 115787
DQ074742 DQ074732 KM495449 DQ074737a
Al-Subhi et al. (2006)
Hu. salinaria
CBS 129733T
HQ203213 HQ203230 KM495461 HQ236432c
Kamgan Nkuekam et al.(2013), De Beer et al. (2014)
Hu. savannae
CBS 121151T
EF408551
EF408565
KM495462 EF408572c
Kamgan et al. (2008), De Beer et al. (2014)
Hu. sublaevis
CBS 122517
FJ151431
FJ151465
KM495464 FJ151486b
Van Wyk et al. (2011), De Beer et al. (2014)
Hu. sumatrana
CBS 124011PT EU588589 EU588668 KM495465 EU588678a
Tarigan et al. (2010), De Beer et al. (2014)
Hu. tribiliformis
CBS 115866T
AY529003 AY529003 KM495468 AY529014a
Van Wyk et al. (2006), De Beer et al. (2014)
Hu. tyalla
CBS 128703T
HM071900 HM071913 KM495470 HQ236452c
Kamgan Nkuekam et al. (2012), De Beer et al. (2014)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T and PT indicate ex-type and ex-paratype strains, respectively.
2
ITS: internal transcribed spaces and intervening 5.8S nrDNA, tub2: partial β-tubulin gene, mcm7: mini-chromosome maintenance complex component 7, tef1:
partial translation elongation factor 1-alpha gene. a, b and c in tef1column indicate the primers used in sequencing: a: Ef1-728f, ef1-986r, b: EF1f, EF1r, c:EF1f,
EF2r.
ACCEPTED MANUSCRIPT
Table 9. DNA barcodes of accepted Macgarvieomyces spp.
1
2
SC
M
AN
U
D
TE
EP
AC
C
Isolates
RI
PT
GenBank accession numbers
References
ITS
act
cal
rpb1
Macgarvieomyces borealis
CBS 461.65T
KM484854
KM485170
KM485239
KM485070
Klaubauf et al. (2014)
M. juncicola
CBS 610.82
KM484855
KM485171
KM485240
KM485071
Klaubauf et al. (2014)
M. luzulae
CBS 143401ET
MG934440
MG934463
MG934519
MG934469
Present study
CPC 31555
MG934441
MG934464
MG934520
MG934470
Present study
CPC 31571
MG934442
MG934465
MG934521
MG934471
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.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; cal: partial calmodulin gene; rpb1: partial RNA polymerase
II largest subunit gene.
Species
ACCEPTED MANUSCRIPT
Table 10. DNA barcodes of accepted Metulocladosporiella spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
GenBank accession numbers2
References
ITS
cal
tef1
Metulocladosporiella chiangmaiensis CBS 143918T
MG934443
MG934522
MG934476
Present study
M. malaysiana
CBS 143919T
MG934444
MG934523
MG934477
Present study
ET
M. musae
CBS 161.74
DQ008137
MG934478
Crous et al. (2006a), present study
CBS 113863
DQ008138
MG934524
MG934479
Crous et al. (2006a), present study
CPC 33937
MG934445
MG934525
MG934480
Present study
T
M. musicola
CBS 110960
DQ008127
MG934526
MG934481
Crous et al. (2006a), present study
CBS 110962
MG934446
MG934527
MG934482
Crous et al. (2006a), present study
CBS 110964
MG934447
MG934528
MG934483
Crous et al. (2006a), present study
CBS 113860
DQ008130
MG934529
MG934484
Crous et al. (2006a), present study
CBS 113861
DQ008131
MG934530
MG934485
Crous et al. (2006a), present study
CBS 113862
DQ008132
MG934531
MG934486
Crous et al. (2006a), present study
CBS 113864
DQ008133
MG934532
MG934487
Crous et al. (2006a), present study
CBS 113865
DQ008134
MG934533
MG934488
Crous et al. (2006a), present study
CBS 113873
DQ008135
MG934534
MG934489
Crous et al. (2006a), present study
CPC 18124
MG934448
MG934535
MG934490
Present study
CPC 32807
MG934449
MG934536
MG934491
Present study
CPC 32849
MG934450
MG934537
MG934492
Present study
CPC 32970
MG934451
MG934538
MG934493
Present study
T
M. musigena
CBS 143920
MG934452
MG934539
MG934494
Present study
M. samutensis
CBS 143921T
MG934453
MG934540
MG934495
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.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; cal: partial calmodulin gene; tef1: partial translation elongation factor 1-alpha gene.
Species
ACCEPTED MANUSCRIPT
Table 12. DNA barcodes of accepted Oculimacula spp.
Isolates1
GenBank accession numbers2
References
RI
PT
Species
ITS
tef1
CBS 495.80T
MG934455
MG934497
Present study
CBS 114730
MG934454
MG934496
Present study
T
CBS 496.80
LT990662
LT990618
Present study
MG934456
MG934498
Present study
CBS 110665NT
CBS 128.31
MG934457
MG934499
Present study
CBS 494.80
JF412009
MG934500
Tsang (unpubl. data), present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T and NT indicate ex-type and ex-neotype strains,
respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA, tef1: partial translation elongation factor 1-alpha gene.
AC
C
EP
TE
D
M
AN
U
SC
Oculimacula acuformis
O. aestiva
O. anguioides
O. yallundae
ACCEPTED MANUSCRIPT
Table 13. DNA barcodes of accepted Paraphoma spp.
Species
Isolates1
GenBank accession number2
tef1
tub2
rpb2
References
RI
PT
ITS
AC
C
EP
TE
D
M
AN
U
SC
Paraphoma
BRIP 65168T
KU999072 KU999080
KU999084 Moslemi et al. (2018)
chlamydocopiosa
NT
Pa. chrysanthemicola
CBS 522.66
KF251166 KF253124
KF252661 KF252174
Quaedvlieg et al. (2013)
Pa. dioscoreae
CBS 135100T
KF251167 KF253125
KF252662 KF252175
Quaedvlieg et al. (2013)
KF252665 KF252178
Quaedvlieg et al. (2013)
Pa. fimeti
CBS 170.70NT KF251170 KF253128
Pa. pye
BRIP 65169T
KU999073 KU999081
KU999085 Moslemi et al. (2018)
ET
KF251172 KF253130
KF252667 KF252180
Quaedvlieg et al. (2013)
Pa. radicina
CBS 111.79
Pa. rhaphiolepidis
CBS 142524T
KY979758 KY979896
KY979924 KY979851 Crous et al. (2017a)
Pa. vinacea
BRIP 63684T
KU176884 KU176896
KU176892 Moslemi et al. (2016)
1
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands. T,
ET
and NT indicate ex-type, ex-epitype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin
gene; act: partial actin gene; rpb2: partial RNA polymerase II second largest subunit gene.
ACCEPTED MANUSCRIPT
Table 14. DNA barcodes of accepted Phaeoacremonium spp.
CBS 120863T
CBS 142688T
CBS 110034T
CBS 110627T
CBS 128570T
CBS 114992T
IFRDCC 3035T
CBS 777.83T
ICMP 17421T
CBS 142691T
CBS 113589T
CBS 112949T
CBS 142694T
PARC 327T
CBS 123909T
CBS 123037T
CBS 101585T
STE-U 5969T
CBS 142712T
CBS 142713T
ICMP 16988T
STE-U 5966T
CBS 111657T
CBS 123910T
CBS 123036T
CBS 391.71T
CBS 101357T
CBS 137763T
CBS 142697T
CBS 109479T
CBS 110156*
CBS 142699T
CBS 137497T
CBS 142710T
CBS 246.91T
CMM 4312T
ICMP 17037T
CBS 142704T
CBS 860.73T
STE-U 6104T
CBS 142705T
CBS 142686T
CBS 142706T
STE-U 5967T
CBS 142101T
PARC 273T
CBS 142708T
CBS 498.94T
CBS 137498T
CBS 113597T
CBS 123034T
CBS 142711T
CBS 337.90T
CBS 113584T
CBS 110573T
MFLUCC 13-0707T
CBS 111586T
CBS 123033T
TE
EP
AC
C
References
Damm et al. (2008)
Spies et al. (2018)
Mostert et al. (2005)
Mostert et al. (2005)
Gramaje et al. (2012)
Mostert et al. (2006)
Hu et al. (2012)
Mostert et al. (2006)
Graham et al. (2009)
Spies et al. (2018)
Mostert et al. (2005)
Mostert et al. (2006)
Spies et al. (2018)
Úrbez-Torres et al. (2014)
Gramaje et al. (2009)
Essakhi et al. (2008)
Groenewald et al. (2001)
Damm et al. (2008)
Spies et al. (2018)
Spies et al. (2018)
Graham et al. (2009)
Damm et al. (2008)
Mostert et al. (2005)
Gramaje et al. (2009)
Essakhi et al. (2008)
Mostert et al. (2006)
Mostert et al. (2006)
Raimondo et al. (2014)
Spies et al. (2018)
Mostert et al. (2005)
Réblová (2011)
Spies et al. (2018)
Gramaje et al. (2014)
Spies et al. (2018)
Mostert et al. (2006)
da Silva et al. (2017)
Graham et al. (2009)
Spies et al. (2018)
Mostert et al. (2006)
Damm et al. (2008)
Spies et al. (2018)
Present study
Spies et al. (2018)
Damm et al. (2008)
Crous et al. (2016a)
Úrbez-Torres et al. (2014)
Spies et al. (2018)
Mostert et al. (2006)
Gramaje et al. (2014)
Mostert et al. (2005)
Essakhi et al. (2008)
Spies et al. (2018)
Mostert et al. (2006)
Mostert et al. (2005)
Mostert et al. (2005)
Ariyawansa et al. (2015)
Mostert et al. (2006)
Essakhi et al. (2008)
RI
PT
Phaeoacremonium africanum
Pha. album
Pha. alvesii
Pha. amstelodamense
Pha. amygdalinum
Pha. angustius
Pha. aquaticum
Pha. argentinense
Pha. armeniacum
Pha. aureum
Pha. australiense
Pha. austroafricanum
Pha. bibendum
Pha. canadense
Pha. cinereum
Pha. croatiense
Pha. fraxinopennsylvanicum
Pha. fuscum
Pha. gamsii
Pha. geminum
Pha. globosum
Pha. griseo-olivaceum
Pha. griseorubrum
Pha. hispanicum
Pha. hungaricum
Pha. inflatipes
Pha. iranianum
Pha. italicum
Pha. junior
Pha. krajdenii
Pha. leptorrhynchum
Pha. longicollarum
Pha. luteum
Pha. meliae
Pha. minimum
Pha. nordesticola
Pha. occidentale
Pha. oleae
Pha. parasiticum
Pha. pallidum
Pha. paululum
Pha. pravum
Pha. proliferatum
Pha. prunicola
Pha. pseudopanacis
Pha. roseum
Pha. rosicola
Pha. rubrigenum
Pha. santali
Pha. scolyti
Pha. sicilianum
Pha. spadicum
Pha. sphinctrophorum
Pha. subulatum
Pha. tardicrescens
Pha. tectonae
Pha. theobromatis
Pha. tuscanicum
GenBank accession numbers2
act
tub2
EU128142
EU128100
KY906884
KY906885
AY579234
AY579301
AY579228
AY579295
JN191303
JN191307
DQ173127
DQ173104
n/a3
n/a3
DQ173135
DQ173108
EU595463
EU596526
KY906656
KY906657
AY579229
AY579296
DQ173122
DQ173099
KY906758
KY906759
KF764499
KF764651
FJ517153
FJ517161
EU863514
EU863482
DQ173137
AF246809
EU128141
EU128098
KY906740
KY906741
KY906648
KY906649
EU595466
EU596525
EU128139
EU128097
AY579227
AY579294
FJ517156
FJ517164
EU863515
EU863483
AY579259
AF246805
DQ173120
DQ173096
KJ534046
KJ534074
KY906708
KY906709
AY579267
AY579330
DQ173139
DQ173110
KY906688
KY906689
KF835406
KF823800
KY906824
KY906825
AY735497
AF246811
KY030803
KY030807
EU595460
EU596524
KY906936
KY906937
AY579253
AF246803
EU128144
EU128103
KY906880
KY906881
KY084248
KY084246
KY906902
KY906903
EU128137
EU128095
KY173569
KY173609
KF764506
KF764658
KY906830
KY906831
AY579238
AF246802
KF835403
KF823797
AY579224
AF246800
EU863520
EU863488
KY906838
KY906839
DQ173142
DQ173113
AY579231
AY579298
AY579233
AY579300
KT285563
KT285555
DQ173132
DQ173106
EU863490
EU863458
M
AN
US
C
Isolates1
D
Species
ACCEPTED MANUSCRIPT
Pha. venezuelense
Pha. vibratile
Pha. viticola
CBS 651.85T
CBS 117115T
CBS 101738T
AY579256
DQ649063
DQ173131
AY579320
DQ649064
AF192391
Mostert et al. (2005)
Réblová & Mostert (2007)
Dupont et al. (2000)
EP
TE
D
M
AN
US
C
Not available. Only ITS sequence available: NR136032
AC
C
3
RI
PT
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; IFRDCC: International Fungal Research and Development Center,
Bailongsi, China; ICMP: International Collection of Microorganisms from Plants, Auckland, New Zealand; PARC: Pacific Agri-Food Research
Centre in Summerland, British Columbia, Canada; CMM: Culture Collection of Phytopathogenic Fungi “Prof. Maria Menezes”, Universidade
Federal Rural de Pernambuco, Recife, Brazil; STE-U: Department of Plant Pathology, Stellenbosch University, South Africa; MFLUCC: Mae
Fah Luang Culture Collection, Chiang Rai, Thailand. T indicates ex-type strains. *Ex-type of Pha. novae-zealandiae, synonymised with Pha.
leptorrhynchum by Réblova (2011).
2
act: partial actin gene; tub2: partial β-tubulin gene.
ACCEPTED MANUSCRIPT
Table 15. DNA barcodes of accepted Phyllosticta spp.
Isolates1
Phyllosticta abieticola CBS 112067T
Phy. alliacea
MUCC 0014T
GenBank accession number2
ITS
act
gapdh
tef1
KF170306
KF289238
AB454263
AB704207
-
RI
PT
Species
CBS 136058T
ATCC 200578NT
MUCC 0031T
KF154280
KC193586
AB454274
KF289311
KC193581
AB704216
KF289124
KC193584
-
KF289193
-
Phy. aristolochiicola
Phy. aspidistricola
BRIP 53316aT
MUCC 0010T
JX486129
AB454260
AB704204
-
-
Phy. beaumarisii
CBS 535.87T
AY042927
KF306232
KF289074
KF289170
Phy. bifrenariae
Phy. brazillianiae
Phy. capitalensis
Phy. carissicola
Phy. carochlae
Phy. catimbauensis
Phy. cavendishii
Phy. citriasiana
CBS 128855T
CBS 129060T
CBS 128856ET
CPC 25665T
CGMCC 3.17317T
URM 7672T
BRIP 55420IsoT
CBS 120486T
JF343565
JF343572
JF261465
KT950849
KJ847422
MF466160
JQ743562
FJ538360
JF343649
JF343656
JF343647
KT950872
KJ847430
MF466157
FJ538476
JF343744
JF343758
JF343776
KT950876
KJ847438
JF343686
JF343586
JF343593
JF261507
KT950879
KJ847444
MF466155
FJ538418
Phy. citribraziliensis
CBS 100098T
FJ538352
FJ538468
JF343691
FJ538410
Phy. citricarpa
Phy. citrichinaensis
Phy. citrimaxima
Phy. concentrica
CBS 127454ET
CBS 130529T
CBS 136059T
CBS 937.70ET
JF343583
JN791597
KF170304
FJ538350
JF343667
JN791526
KF289300
KF289257
JF343771
KF289157
JF411745
JF343604
JN791452
KF289222
FJ538408
M
AN
U
D
TE
EP
AC
C
SC
Phy. aloeicola
Phy. ampelicida
Phy. ardisiicola
References
Wikee et al. (2013b)
Motohashi et al. (2009),
Ando et al. (2013)
Wikee et al. (2013b)
Zhang et al. (2013b)
Motohashi et al. (2009),
Ando et al. (2013)
Crous et al. (2012a)
Motohashi et al. (2009),
Ando et al. (2013)
Baayen et al. (2002),
Wikee et al. (2013b)
Glienke et al. (2011)
Glienke et al. (2011)
Glienke et al. (2011)
Crous et al. (2015e)
Zhou et al. (2015)
Crous et al. (2017b)
Wong et al. (2012)
Wulandari et al. (2009),
Glienke et al. (2011)
Wulandari et al. (2009),
Glienke et al. (2011)
Glienke et al. (2011)
Wang et al. (2011)
Wikee et al. (2013b)
Wulandari et al. (2009),
Glienke et al. (2011),
Wikee et al. (2013b)
CBS 136244NT
CBS 111639
CBS 858.71
CBS 136060ET
CBS 126.22T
KF170287
KF170307
MG934458
JF343578
FJ538353
KF289295
KF289234
MG934465
JF343662
FJ538469
KF289076
MG934474
JF343764
KF289164
KF289172
MG934501
JF343599
FJ538411
Phy. ericarum
Phy. eugeniae
CBS 132534T
CBS 445.82
KF206170
AY042926
KF28291
KF289246
KF289162
KF289139
KF289227
KF289208
Phy. fallopiae
MUCC 0113T
AB454307
AB704228
-
-
Phy. foliorum
Phy. gaultheriae
Phy. hamamelidis
Phy. hostae
Phy. hubeiensis
Phy.
hymenocallidicola
Phy. hypoglossi
CBS 447.68NT
CBS 447.70T
MUCC 149
CGMCC 3.14355T
CGMCC 3.14986T
CBS 131309T
KF170309
JN692543
KF170289
JN692535
JX025037
JQ044423
KF289247
KF289248
KF289309
JN692511
JX025032
KF289242
KF289132
JN692508
JN692503
JX025027
KF289142
KF289201
JN692531
JN692523
JX025042
KF289211
CBS 434.92NT
FJ538367
FJ538483
JF343695
FJ538425
Phy. ilicis-aquifolii
Phy. iridigena
Phy. kerriae
CGMCC 3.14358T
CBS 143410T
MUCC 0017T
JN692538
MG934459
AB454266
JN692514
MG934466
AB704209
-
JN692526
MG934502
KC342576
Phy. leucothoicola
CBS 136073T
AB454370
KF289310
-
-
Phy. ligustricola
MUCC 0024T
AB454269
AB704212
-
-
Phy. maculata
Phy. mangiferaeindicae
CBS 132581ET
CBS 136061T
JQ743570
KF170305
KF289296
KF289121
KF289190
M
AN
U
D
TE
EP
AC
C
RI
PT
Phy. cordylinophila
Phy. cornicola
Phy. cruenta
Phy. cussonia
Phy. elongata
SC
ACCEPTED MANUSCRIPT
Wikee et al. (2013b)
Wikee et al. (2013b)
Present study
Glienke et al. (2011)
Wulandari et al. (2009),
Wikee et al. (2013b)
Wikee et al. (2013b)
Baayen et al. (2002),
Wikee et al. (2013b)
Motohashi et al. (2009),
Ando et al. (2013)
Wikee et al. (2013b)
Su & Cai (2012)
Wikee et al. (2013b)
Su & Cai (2012)
Zhang et al. (2013a)
Crous et al. (2011b),
Wikee et al. (2013b)
Wulandari et al. (2009),
Glienke et al. (2011),
Wikee et al. (2013b)
Su & Cai (2012)
Present study
Motohashi et al. (2009),
Ando et al. (2013),
Wikee et al. (2013a)
Motohashi et al. (2009),
Wikee et al. (2013b)
Motohashi et al. (2009),
Ando et al. (2013)
Wong et al. (2012)
Wikee et al. (2013b)
ACCEPTED MANUSCRIPT
CBS 138899T
CBS 585.84NT
BRIP 55434IsoET
CBS 134750T
KP004447
KF206176
JQ743584
AB454318
KF289249
AB704233
KF289135
-
KF289204
-
Phy. owaniana
CBS 776.97ET
FJ538368
KF289254
JF343767
FJ538426
Phy. pachysandricola
MUCC 124T
AB454317
AB704232
-
-
Phy. paracapitalensis
Phy. paracitricarpa
Phy. parthenocisii
Phy. partricuspidatae
Phy. paxistimae
Phy. persooniae
Phy. philoprina
Phy. podocarpi
CBS 141353T
CBS 141357T
CBS 111645
NBRC 9466T
CBS 112527T
CBS 143409T
CBS 587.69
CBS 111646
KY855622
KY855635
JN692542
KJ847424
KF206172
MG934460
KF154278
AF312013
KY855677
KY855690
JN692518
KJ847432
KF289239
MG934467
KF289250
KC357670
KY855735
KY855748
KJ847440
KF289140
MG934475
KF289137
KF289169
KY855951
KY855964
JN692530
KJ847446
KF289209
MG934503
KF289206
KC357671
Phy. podocarpicola
Phy. pseudotsugae
Phy. rhaphiolepidis
CBS 728.79T
CBS 111649
MUCC 432T
KF206173
KF154277
DQ632660
KF289252
KF289236
AB704242
KF289134
KF289167
-
KF289203
KF289231
DQ632724
Phy. rubella
Phy. schimae
Phy. schimicola
Phy. sphaeropsoidea
CBS 111635T
CGMCC 3.14354T
CGMCC 3.17319T
CBS 756.70
KF206171
JN692534
KJ847426
AY042934
KF289233
JN692510
KJ847434
KF289253
KF289129
JN692506
KJ854895
KF289133
KF289198
JN692522
KJ847448
KF289202
Phy. spinarum
Phy. styracicola
Phy. telopeae
Phy. vaccinii
CBS 292.90
CGMCC 3.14985T
CBS 777.97T
ATCC 46255ET
JF343585
JX052040
KF206205
KC193585
JF343669
JX025035
KF289255
KC193580
JF343773
JX025030
KF289141
KC193583
JF343606
JX025045
KF289210
KC193582
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
Phy. mimusopisicola
Phy. minima
Phy. musarum
Phy. neopyrolae
Crous et al. (2014d)
Wikee et al. (2013b)
Wong et al. (2012)
Motohashi et al. (2009),
Ando et al. (2013)
Wulandari et al. (2009),
Glienke et al. (2011),
Wikee et al. (2013b)
Motohashi et al. (2009),
Ando et al. (2013)
Guarnaccia et al. (2017)
Guarnaccia et al. (2017)
Su & Cai (2012)
Zhou et al. (2015)
Wikee et al. (2013b)
Present study
Wikee et al. (2013b)
Wikee et al. (2013b),
Carroll (unpubl. data),
Wikee (unpubl. data)
Wikee et al. (2013b)
Wikee et al. (2013b)
Andjic et al. (2007),
Ando et al. (2013)
Wikee et al. (2013b)
Su & Cai (2012)
Zhou et al. (2015)
Baayen et al. (2002),
Wikee et al. (2013b)
Glienke et al. (2011)
Zhang et al. (2013a)
Wikee et al. (2013b)
Zhang et al. (2013b)
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
Phy. vacciniicola
CBS 136062T
KF170312
KF289287
KF289165
KF289229
Wikee et al. (2013b)
KJ847436
KJ847442
KJ847450
Zhou et al. (2015)
Phy. vitisCGMCC 3.17322T KJ847428
rotundifoliae
Phy. yuccae
CBS 117136
JN692541
JN692517
JN692507
JN692529
Su & Cai (2012)
1
ATCC: American Type Culture Collection, Virginia, USA; BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia;
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CGMCC: Chinese General Microbiological Culture
Collection Center, Beijing, China; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute;
MUCC: Murdoch University, Perth, Western Australia; NBRC: Biological Resource Center, NITE, Chiva, Japan; URM: Culture
Collection Mycobank, Prof. Maria Auxiliadora Cavalcanti, Federal University of Pernambuco, Recife, Brazil. T, ET, IsoT, IsoET and NT
indicate ex-type, ex-epitype, ex-isotype, ex-isoepitype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; gapdh: partial glyceraldehyde-3-phosphate
dehydrogenase gene; tef1: partial translation elongation factor 1-alpha gene.
ACCEPTED MANUSCRIPT
Table 16. DNA barcodes of accepted Proxipyricularia sp.
Proxypiricularia zingiberis
Isolates
1
CBS 133594ET
ITS
AB274434
GenBank accession numbers
act
cal
AB274446
KM485246
2
References
rpb1
KM485091
RI
PT
Species
1
Hirata et al. (2007),
Klaubauf et al. (2014)
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. ET indicates ex-epitype strain.
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; cal: partial calmodulin gene; rpb1: partial RNA polymerase
II largest subunit gene.
AC
C
EP
TE
D
M
AN
U
SC
2
ACCEPTED MANUSCRIPT
Table 17. DNA barcodes of accepted Pyriculariomyces sp.
1
CBS 141328T
ITS
KX228291
GenBank accession numbers
act
cal
KX228361
MG934541
2
References
rpb1
KX228368
EP
TE
D
M
AN
U
SC
Crous et al. (2016b),
present study
CPC 27442
KX228290
KX228360
MG934472
Crous et al. (2016b),
present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal
Biodiversity Institute. T indicates ex-type strain.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; cal: partial calmodulin gene; rpb1: partial RNA polymerase
II largest subunit gene.
AC
C
Pyriculariomyces asari
Isolates
RI
PT
Species
ACCEPTED MANUSCRIPT
Table 18. DNA barcodes of accepted Pyricularia spp.
Isolates1
Pyricularia angulata
Py. ctenantheicola
Py. graminis-tritici
NBRC 9625
CBS 138601T
URM7380T
ITS
AY265322
KM484879
-
GenBank accession numbers2
act
cal
KM485183
KM485253
KU952138
KU952892
References
rpb1
KM485099
-
RI
PT
Species
AC
C
EP
TE
D
M
AN
U
SC
Bussaban et al. (2005)
Klaubauf et al. (2014)
Castroagudín et al.
(2016)
Py. grisea
CBS 128304
KM484881
KM485184
KM485255
KM485101
Klaubauf et al. (2014)
Py. oryzae
CBS 255.38
KM484889
KM485190
KM485261
KM485109
Klaubauf et al. (2014)
Py. penniseticola
CBS 138603T
KM484929
KM485220
KM485148
Klaubauf et al. (2014)
KM484935
KM485225
KM485294
KM485153
Klaubauf et al. (2014)
Py. pennisetigena
CBS 138604T
Py. urashimae
CBS 142117T
KY173437
KY173571
KY173578
Crous et al. (2016a)
KM484941
KM485229
KM485297
KM485157
Klaubauf et al. (2014)
Py. zingibericola
CBS 138605T
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; NBRC: Biological Resource Center, NITE, Chiva, Japan;
URM: Culture Collection Mycobank, Prof. Maria Auxiliadora Cavalcanti, Federal University of Pernambuco, Recife, Brazil. T
indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; cal: partial calmodulin gene; rpb1: partial RNA
polymerase II largest subunit gene.
ACCEPTED MANUSCRIPT
Table 19. DNA barcodes of accepted Stenocarpella spp.
Isolates1
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
GenBank accession number2 References
ITS
tef1
MG934504
Lamprecht et al. (2011), present study
Stenocarpella macrospora CBS 117560ET FR748048
S. maydis
CBS 117558ET FR748051
FR748080
Lamprecht et al. (2011)
1
ET
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands. indicates ex-epitype strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; tef1: partial translation elongation factor 1-alpha gene.
Species
ACCEPTED MANUSCRIPT
Table 20. DNA barcodes of accepted Utrechtiana spp.
1
CPC 33994ET
CBS 128780T
ITS
MG934461
JF951153
GenBank accession numbers
act
cal
MG934468
MG934542
KM485163
KM485232
2
References
rpb1
MG934473
KM485047
EP
TE
D
M
AN
U
SC
Present study
Crous et al. (2011a),
Klaubauf et al. (2014)
1
T
ET
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. and indicate ex-type and ex-epitype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; cal: partial calmodulin gene; rpb1: partial RNA polymerase
II largest subunit gene.
AC
C
Utrechtiana arundinacea
U. roumeguerei
Isolates
RI
PT
Species
ACCEPTED MANUSCRIPT
Table 21. DNA barcodes of accepted Wojnowiciella spp.
1
ITS
KX228272
CBS 141297T
GenBank accession numbers
LSU
rpb2
KX228323
-
2
References
tef1
LT990616
EP
TE
D
M
AN
U
SC
Crous et al. (2016b),
present study
W. dactylidis
MFLUCC 13-0735T
KP744470
KP684149
Liu et al. (2015)
CPC 27468
LT990658
LT990630
LT990644
LT990611
Present study
CPC 30353
LT990659
LT990631
LT990612
Present study
CPC 32741
LT990660
LT990632
LT990613
Present study
CPC 33929
LT990661
LT990633
LT990645
LT990614
Present study
W. eucalypti
CBS 139904T
KR476741
KR476774
LT990617
Crous et al. (2015d),
present study
W. leptocarpi
CBS 115684T
KX306775
KX306800
LT990646
LT990615
Hernández-Restrepo et
al. (2016c), present
study
W. lonicerae
MFLUCC 13-0737T
KP744471
KP684151
Liu et al. (2015)
W. spartii
MFLUCC 13-0402T
KU058719
KU058729
Li et al. (2015)
W. viburni
MFLUCC 12-0733T
KC594286
KC594287
Wijayawardene et al.
(2013)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Personal collection of Pedro Crous housed at the Westerdijk
Fungal Biodiversity Institute; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand. T indicates ex-type strain.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit RNA gene; rpb2: partial RNA polymerase II largest
subunit gene; tef1: partial translation elongation factor 1-alpha gene.
AC
C
Wojnowiciella cissampeli
Isolates
RI
PT
Species
ACCEPTED MANUSCRIPT
Table 4. DNA barcodes of accepted Diaporthe spp.
Isolates1
GenBank accession numbers
ITS
T
CBS 129521
D. acericola
MFLUCC 17-0956
D. acerina
CBS 137.27
D. acutispora
D. alleghaniensis
D. alnea
D. ambigua
tub2
KP004509
his3
KP004504
tef1
KC343005
KC343973
KC343489
KY964224
KY964074
-
KC343006
KC343974
KC343490
KC343732
cal
-
-
Crous et al. (2014b)
KC343731
KC343247
Gomes et al. (2013)
KY964180
KY964137
KC343248
Dissanayake et al.
(2017a)
Gomes et al. (2013)
KX986764
KX999195
KX999235
KX999155
KX999274
Gao et al. (2017)
CBS 495.72
T
FJ889444
KC843228
KC343491
GQ250298
KC343249
Gomes et al. (2013)
CBS 146.46
T
KC343008
KC343976
KC343492
KC343734
KC343250
Gomes et al. (2013)
T
KC343010
KC343978
KC343494
KC343736
KC343252
Gomes et al. (2013)
T
AF230751
JX275452
-
GQ250351
JX197443
Gomes et al. (2013)
T
CBS 114015
CBS 114016
D. amygdali
CBS 126679
KC343022
KC343990
KC343506
KC343748
KC343264
Gomes et al. (2013)
D. anacardii
CBS 720.97
T
KC343024
KC343992
KC343508
KC343750
KC343266
Gomes et al. (2013)
KC343026
D
D. ampelina
CGMCC 3.18285
T
T
KP004460
RI
PT
D. acaciigena
CBS 138862
References
2
SC
Diaporthe acaciarum
T
M
AN
U
Species
KC343511
KC343752
KC343268
Gomes et al. (2013)
LC 3418
T
IFRDCC 3051
D. arctii
CBS 136.25
D. arecae
CBS 161.64T
KP267896
KP293476
KP293550
KP267970
-
Gao et al. (2016)
JQ797437
-
-
-
-
Hu et al. (2012)
KC343031
KC343999
KC343515
KC343757
KC343273
Gomes et al. (2013)
KC343032
KC344000
KC343516
KC343758
KC343274
Gomes et al. (2013)
AC
C
D. aquatica
T
KC343994
TE
D. apiculatum
CBS 111592
EP
D. angelicae
T
D. arengae
T
CBS 114979
KC343034
KC344002
KC343518
KC343760
KC343276
Gomes et al. (2013)
D. aseana
MFLUCC 12T
0299a
T
CBS 136967
KT459414
KT459432
-
KT459448
KT459464
Hyde et al. (2016)
D. asheicola
KJ160562
KJ160518
-
KJ160594
KJ160542
Lombard et al. (2014)
T
KC343036
KC344004
KC343520
KC343762
KC343278
T
KC343038
KC344006
KC343522
KC343764
KC343280
Van Rensburg et al.
(2006)
Gomes et al. (2013)
D. aspalathi
CBS 117169
D. australafricana
CBS 111886
ACCEPTED MANUSCRIPT
D. baccae
T
CBS 136972
KJ160565
MF418509
MF418264
KJ160597
-
Lombard et al. (2014)
CBS 122.21
KC343040
KC344008
KC343524
KC343766
KC343282
Gomes et al. (2013)
CBS 138.27
T
BRIP 54792
KC343041
JX862529
KC344009
KF170921
KC343525
-
KC343767
JX862535
KC343283
-
Gomes et al. (2013)
Thompson et al. (2015)
D. benedicti
CFCC 50062
T
KP208847
KP208855
KP208851
KP208853
KP208849
Fan et al. (2015)
CFCC 50469
T
KT732950
KT733020
KT732999
KT733016
KT732997
Du et al. (2016)
CFCC 51128
T
KX024653
KX024657
KX024661
KX024655
KX024659
Du et al. (2016)
T
KC343134
KC344102
KC343618
KC343860
KC343376
Gomes et al. (2013)
KJ490597
KJ490418
KJ490539
KJ490476
-
Huang et al. (2015)
KJ490582
KJ490403
KJ490524
KJ490461
-
Huang et al. (2015)
D. bicincta
CBS 121004
D. biconispora
CGMCC 3.17252
D. biguttusis
T
ICMP20657
KF576282
KF576306
-
KF576257
-
Gao et al. (2015)
T
MG281015
MG281188
MG281361
MG281536
MG281710
T
KC343042
KC344010
KC343526
KC343768
KC343284
Guarnaccia et al.
(2018)
Gomes et al. (2013)
T
KY085927
KY115600
KY115605
KY115603
KY115597
Crous et al. (2016a)
T
KY203726
KY228893
KY228881
KY228887
KY228877
Yang et al. (2017c)
T
KC843230
-
KC843120
KC843174
Crous et al. (2012b)
CGMCC 3.17081
D. bohemiae
CBS 143347
D. brasiliensis
CBS 133183
D. caatingaensis
T
D
D. biguttulata
T
CBS 141542
CFCC 51632
D. canthii
CBS 132533
JX069864
D. carpini
CBS 114437
KC344012
KC343528
KC343770
KC343286
Gomes et al. (2013)
KF777155
-
-
KF777244
-
Crous et al. (2013)
T
CBS 127268
KC343045
KC344013
KC343529
KC343771
KC343287
Gomes et al. (2013)
T
KC343047
KC344015
KC343531
KC343773
KC343289
Gomes et al. (2013)
CBS 136440
CBS 139.27
EP
D. celastrina
KC343044
T
AC
C
D. caulivora
TE
D. camptothecicola
D. cassines
SC
D. betulicola
M
AN
U
D. betulae
RI
PT
D. batatas
D. beckhausii
D. beilharziae
T
MG281017
MG281190
MG281363
MG281538
MG281712
T
D. celeris
CBS 143349
D. ceratozamiae
CBS 131306
JQ044420
-
-
-
-
Guarnaccia et al.
(2018)
Crous et al. (2011b)
D. cf. heveae 1
D. cf. heveae 2
CBS 852.97
CBS 681.84
KC343116
KC343117
KC344084
KC344085
KC343600
KC343601
KC343842
KC343843
KC343358
KC343359
Gomes et al. (2013)
Gomes et al. (2013)
D. chamaeropis
CBS 454.81
KC343048
KC344016
KC343532
KC343774
KC343290
Gomes et al. (2013)
KJ197288
KJ197268
-
KJ197250
-
Thompson et al. (2015)
D. charlesworthii
T
BRIP 54884m
ACCEPTED MANUSCRIPT
D. cinerascens
CBS 719.96
T
KC343050
KC344018
KC343534
KC343776
KC343292
Dissanayake et al.
(2017a)
Gomes et al. (2013)
T
KX228273
KX228384
KX228366
-
-
Crous et al. (2016b)
D. citri
T
CBS 135422
KC843311
KC843187
MF418281
KC843071
KC843157
D. citriasiana
CBS 134240T
JQ954645
KC357459
MF418282
JQ954663
KC357491
Udayanga et al.
(2014b)
Huang et al. (2013)
D. citrichinensis
CBS 134242T
JQ954648
MF418524
KJ420880
JQ954666
KC357494
Huang et al. (2013)
KP267854
KP293434
KP293508
KP267928
-
Gao et al. (2016)
D. cissampeli
D. compacta
CBS 141331
T
LC3083
KY964220
KY964104
-
KY964176
KY964133
RI
PT
MFLUCC 17-1023
SC
D. cichorii
CBS 124654
KC343054
KC344022
KC343538
KC343780
KC343296
Gomes et al. (2013)
CBS 114435
KC343055
KC344023
KC343539
KC343781
KC343297
Gomes et al. (2013)
D. crotalariae
CBS 162.33
KC343056
KC344024
KC343540
KC343782
KC343298
Gomes et al. (2013)
DAOM 42078
T
KM453210
KP118848
KM453212
KM453211
-
Udayanga et al. (2015)
D. cuppatea
T
CBS 117499
AY339322
JX275420
KC343541
AY339354
JX197414
D. cynaroidis
D. cytosporella
CBS 122676
T
CBS 137020
KC343058
KC843307
KC344026
KC843221
KC343542
MF418283
KC343784
KC843116
KC343300
KC843141
D
D. cucurbitae
T
M
AN
U
D. convolvuli
D. crataegi
D. decedens
CBS 109772
KC343059
KC343543
KC343785
KC343301
Van Rensburg et al.
(2006)
Gomes et al. (2013)
Udayanga et al.
(2014b)
Gomes et al. (2013)
D. detrusa
CBS 109770
KC343061
KC344029
KC343545
KC343787
KC343303
Gomes et al. (2013)
KF777156
-
-
-
-
Crous et al. (2013)
KJ490624
KJ490445
KJ490566
KJ490503
-
Huang et al. (2015)
KY964215
KY964099
-
KY964171
-
KX986779
KX999212
KX999251
KX999171
KX999281
Dissanayake et al.
(2017a)
Gao et al. (2017)
TE
D. discoidispora
T
CBS 136552
T
ICMP20662
MFLUCC 17-1015
D. elaeagni-glabrae
CGMCC 3.18287T
AC
C
D. dorycnii
D. eleagni
CBS 504.72
D. ellipicola
CGMCC 3.17084
D. endophytica
T
EP
D. diospyricola
KC344027
KC343064
KC344032
KC343548
KC343790
KC343306
Gomes et al. (2013)
KF576270
KF576291
-
KF576245
-
Gao et al. (2015)
T
KC343065
KC344033
KC343549
KC343791
KC343307
Gomes et al. (2013)
T
KJ210529
KJ420799
KJ420850
KJ210550
KJ434999
T
JX069862
-
-
-
-
Udayanga et al.
(2014a)
Crous et al. (2012b)
CBS 133811
D. eres
CBS 138594
D. eucalyptorum
CBS 132525
T
ACCEPTED MANUSCRIPT
CBS 444.82
KC343098
KC344066
KC343582
KC343824
KC343340
Gomes et al. (2013)
D. fibrosa
D. foeniculina
CBS 109751
T
CBS 111553
KC343099
KC343101
KC344067
KC344069
KC343583
KC343585
KC343825
KC343827
KC343341
KC343343
Gomes et al. (2013)
Gomes et al. (2013)
D. fraxini-angustifoliae
BRIP 54781
JX862528
KF170920
-
JX852534
-
Tan et al. (2013)
KF576281
KF576305
-
KF576256
KF576233
Gao et al. (2015)
KC343112
KC344080
KC343596
KC343838
KC343354
Gomes et al. (2013)
KC343839
KC343355
Gomes et al. (2013)
KT459457
KT459470
Hyde et al. (2016)
KJ197252
-
Thompson et al. (2015)
D. fusicola
T
CGMCC 3.17087
T
T
RI
PT
D. eugeniae
CBS 180.91
D. gardeniae
CBS 288.56
KC343113
KC344081
KC343597
D. garethjonesii
KT459423
KT459441
-
D. goulteri
MFLUCC 120542aT
T
BRIP 55657a
KJ197290
KJ197270
-
D. gulyae
BRIP 54025
T
JF431299
KJ197271
-
JN645803
-
Thompson et al. (2015)
T
M
AN
U
CBS 592.81
KC343115
KC344083
KC343599
KC343841
JX197454
Gomes et al. (2013)
T
KJ210538
KJ420828
KJ420875
KJ210559
KJ435043
T
MG600222
MG600226
MG600220
MG600224
MG600218
Udayanga et al.
(2014a)
Present study
KC343118
KC344086
KC343602
KC343844
KC343360
Gomes et al. (2013)
T
MG281123
D
D. helianthi
SC
D. ganjae
MG281471
MG281644
MG281820
T
D. helicis
CBS 138596
D. heterophyllae
CBS 143769
T
D. hickoriae
CBS 145.26
D. hispaniae
CBS 143351
D. hongkongensis
CBS 115448
KC343119
KC344087
KC343603
KC343845
KC343361
Guarnaccia et al.
(2018)
Gomes et al. (2013)
D. hordei
CBS 481.92
KC343120
KC344088
KC343604
KC343846
KC343362
Gomes et al. (2013)
TE
T
MG281296
CBS 143353
MG281126
MG281299
MG281474
MG281647
MG281823
D. impulsa
D. incompleta
CBS 114434
T
CGMCC 3.18288
KC343121
KX986794
KC344089
KX999226
KC343605
KX999265
KC343847
KX999186
KC343363
KX999289
Guarnaccia et al.
(2018)
Gomes et al. (2013)
Gao et al. (2017)
D. inconspicua
CBS 133813
T
KC343123
KC344091
KC343607
KC343849
KC343365
Gomes et al. (2013)
T
CBS 133812
KC343126
KC344094
KC343610
KC343852
KC343368
Gomes et al. (2013)
T
KC343052
KC344020
KC343536
KC343778
KC343294
T
KJ869133
KJ869245
-
-
-
Guarnaccia & Crous
(2017)
Crous et al. (2014c)
T
KU985101
KX024634
-
KX024628
KX024616
Yang et al. (2017a)
AC
C
D. infecunda
D. infertilis
CBS 230.52
D. isoberliniae
CBS 137981
D. juglandicola
EP
D. hungariae
CFCC 51134
ACCEPTED MANUSCRIPT
-
-
JN645809
-
Thompson et al. (2011)
BRIP 54031
KJ197272
-
JN645797
-
Thompson et al. (2011)
JN712460
KY435673
KY435653
KY435632
KY435663
Crous et al. (2011c)
T
MF418422
MF418582
MF418342
MF418501
MF418256
JX862533
KF170925
-
JX862539
-
Guarnaccia & Crous
(2017)
Tan et al. (2013)
KC153104
KF576311
-
KC153095
-
Gao et al. (2014)
MF190139
-
-
-
-
KF576267
KF576291
-
KF576242
-
Senanayake et al.
(2017)
Gao et al. (2015)
KJ590728
KJ610883
KJ659188
KJ590767
KJ612124
Udayanga et al. (2015)
KC343135
KC344103
KC343619
KC343861
KC343377
Gomes et al. (2013)
KY964190
KY964073
-
KY964146
KY964116
KC343136
KC344104
KC343620
KC343862
KC343378
Dissanayake et al.
(2017a)
Gomes et al. (2013)
KJ197289
KJ197269
-
KJ197251
-
Thompson et al. (2015)
KC153096
-
-
KC153087
-
Gao et al. (2014)
KY435668
KY435648
KY435627
KY435658
Santos et al. (2017)
D. limonicola
CBS 142549
D. litchicola
BRIP 54900
D. lithocarpus
T
CGMCC 3.15175
T
D. litoricola
MFLUCC 16-1195
D. longicicola
CGMCC 3.17089
D. longicolla
D. longispora
T
T
FAU 599
CBS 194.36
T
D. lonicerae
MFLUCC 17-0963
D. lusitanicae
CBS 123212T
T
D. macintoshii
BRIP 55064a
D. mahothocarpus
CGMCC 3.15181
T
D. malorum
CBS142383
D. manihotia
CBS 505.76
D. maritima
D. masirevicii
T
KY435638
DAOMC 250563
BRIP 57892a
T
T
T
M
AN
U
CBS 111980
SC
JF431301
T
RI
PT
JF431295
T
D
D. leucospermi
T
TE
D. kongii
BRIP 54033
KC343138
KC344106
KC343622
KC343864
KC343380
Gomes et al. (2013)
KU552025
KU574615
-
KU552023
-
Tanney et al. (2016)
EP
D. kochmanii
KJ197277
KJ197257
-
KJ197239
-
Thompson et al. (2015)
T
KC343139
KC344107
KC343623
KC343865
KC343381
Gomes et al. (2013)
CBS 133185
D. maytenicola
CBS 136441
T
KF777157
KF777250
-
-
-
Crous et al. (2013)
D. megalospora
D. melitensis
CBS 143.27
CBS 142551T
KC343140
MF418424
KC344108
MF418584
KC343624
MF418344
KC343866
MF418503
KC343382
MF418258
D. melonis
CBS 507.78
KC343142
KC344110
KC343626
KC343868
KC343384
Gomes et al. (2013)
Guarnaccia & Crous
(2017)
Gomes et al. (2013)
D. middletonii
BRIP 54884e
KJ197286
KJ197266
-
KJ197248
-
Thompson et al. (2015)
KJ197283
KJ197263
-
KJ197245
-
Thompson et al. (2015)
D. miriciae
AC
C
D. mayteni
T
T
T
BRIP 54736j
ACCEPTED MANUSCRIPT
ICMP20656
D. musigena
KU557563
KU557587
-
KU557631
KU557611
KJ490633
KJ490454
KJ490575
KJ490512
-
Dissanayake et al.
(2017c)
Huang et al. (2015)
CBS 129519
KC343143
KC344111
KC343627
KC343869
KC343385
Gomes et al. (2013)
T
KC343144
KC344112
KC343628
KC343870
KC343386
Gomes et al. (2013)
KC343145
KC344113
KC343629
KC343871
KC343387
Gomes et al. (2013)
KU712449
KU743988
-
KU749369
KU749356
Doilom et al. (2017)
KC343154
JX862530
KC344122
KF170922
KC343638
-
KC343880
JX862536
KC343396
-
Gomes et al. (2013)
Tan et al. (2013)
T
KC343157
KC344125
KC343641
KC343883
KC343399
Gomes et al. (2013)
T
MG386072
-
MG386137
-
-
Crous et al. (2017b)
T
T
T
D. neilliae
CBS 144.27
D. neoarctii
CBS 109490
MFLUCC 14-1136
D. nomurai
D. nothofagi
CBS 157.29
T
BRIP 54801
D. novem
CBS 127271
D. obtusifoliae
CBS 143449
D. ocoteae
CBS 141330
KX228293
KX228388
-
-
-
Crous et al. (2016b)
D. oncostoma
CBS 589.78
KC343162
KC344130
KC343646
KC343888
KC343404
Gomes et al. (2013)
KP267863
KP293443
KP293517
KP267937
-
Gao et al. (2016)
KJ490628
KJ490449
KJ490570
KJ490507
-
Huang et al. (2015)
KF576264
D
D. neoraonikayaporum
T
RI
PT
D. multigutullata
T
SC
MFLUCC 16-0113
M
AN
U
D. momicola
KF576288
-
KF576239
KF576222
Gao et al. (2015)
KC344132
KC343648
KC343890
KC343406
Gomes et al. (2013)
D. oraccinii
D. ovalispora
D. ovoicicola
LC 3166
T
T
ICMP20659
CGMCC 3.17092
T
T
CBS 133186
KC343164
D. padi var. padi
D. paranensis
CBS 114200
CBS 133184
KC343169
KC343171
KC344137
KC344139
KC343653
KC343655
KC343895
KC343897
KC343411
KC343413
Gomes et al. (2013)
Gomes et al. (2013)
D. parapterocarpi
CBS 137986
KJ869138
KJ869248
-
-
-
Crous et al. (2014c)
T
JX862532
KF170924
-
JX862538
-
Tan et al. (2013)
D. passiflorae
EP
BRIP 54847
AC
C
D. pascoei
T
TE
D. oxe
T
JX069860
KY435674
KY435654
KY435633
KY435664
Crous et al. (2012b)
T
CBS 132527
D. passifloricola
CBS 141329
KX228292
KX228387
KX228367
-
-
Crous et al. (2016b)
D. penetriteum
LC 3353
KP714505
KP714529
KP714493
KP714517
-
Gao et al. (2016)
T
D. perjuncta
CBS 109745
KC343172
KC344140
KC343656
KC343898
KC343414
Gomes et al. (2013)
D. perniciosa
D. perseae
CBS 124030
CBS 151.73
KC343149
KC343173
KC344117
KC344141
KC343633
KC343657
KC343875
KC343899
KC343391
KC343415
Gomes et al. (2013)
Gomes et al. (2013)
ACCEPTED MANUSCRIPT
MFLUCC 16-0105
D. phaseolorum
CBS 113425
D. phragmitis
T
T
CBS 138897
T
KU557555
KU557579
-
KU557623
KU557603
KC343174
KC344142
KC343658
KC343900
KC343416
Dissanayake et al.
(2017c)
Gomes et al. (2013)
KP004445
KP004507
KP004503
-
-
Crous et al. (2014b)
KX986774
KX999207
KX999246
KX999167
KX999278
Gao et al. (2017)
KC343907
KC343423
Gomes et al. (2013)
KC343910
KC343426
Gomes et al. (2013)
KY964181
KY964138
KF777245
-
Dissanayake et al.
(2017a)
Crous et al. (2013)
-
-
Crous et al. (2013)
CGMCC3.18281
CBS 101339T
KC343181
KC344149
KC343665
D. pseudophoenicicola
CBS 462.69T
KC343184
KC344152
KC343668
D. pseudotsugae
MFLU 15-3228
KY964225
KY964108
-
D. psoraleae
CBS 136412
T
KF777158
KF777251
-
T
KF777159
KF777252
-
M
AN
U
MFLUCC 10-0571
JQ619899
JX275460
-
JX275416
JX197451
Udayanga et al. (2012)
MFLUCC 10-0580a
JQ619887
JX275441
-
JX275403
JX197433
Udayanga et al. (2012)
D. pulla
CBS 338.89
T
KC343152
KC344120
KC343636
KC343878
KC343394
Gomes et al. (2013)
D. pustulata
D. pyracanthae
CBS 109742
T
CBS142384
KC343185
KY435635
KC344153
KY435666
KC343669
KY435645
KC343911
KY435625
KC343427
KY435656
Gomes et al. (2013)
Santos et al. (2017)
D. racemosae
CBS 143770
MG600223
MG600227
MG600221
MG600225
MG600219
Present study
KC343188
KC344156
KC343672
KC343914
KC343430
Gomes et al. (2013)
KU900335
KX432254
-
KX365197
-
KC343189
KC344157
KC343673
KC343915
KC343431
Dissanayake et al.
(2017a)
Gomes et al. (2013)
KP208847
KP208855
KP208851
KP208853
KP208849
Fan et al. (2015)
KC343234
KC344202
KC343718
KC343960
KC343476
KC343190
KC344158
KC343674
KC343916
KC343432
Udayanga et al.
(2014b)
Gomes et al. (2013)
KJ197287
KJ197267
-
KJ197249
-
Thompson et al. (2015)
JX862531
KF170923
-
JX862537
-
Tan et al. (2013)
T
KY852495
KY852511
KY852503
KY852507
KY852499
Yang et al. (2018)
T
KC343191
KC344159
KC343675
KC343917
KC343433
Gomes et al. (2013)
D. raonikayaporum
D. ravennica
T
T
CBS 133182
MFLUCC 15–0479
CBS 146.27
CFCC 50062
D. rudis
CBS 113201
D. saccarata
CBS 116311
D. sackstonii
T
AC
C
D. rhoina
D. rostrata
T
TE
D. pterocarpi
D. pterocarpicola
D
CBS 136413
EP
D. psoraleae-pinnatae
SC
D. podocarpimacrophylli
D. pseudomangiferae
RI
PT
D. pescicola
T
BRIP 54669b
T
T
D. salicicola
BRIP 54825
D. sambucusii
CFCC 51986
D. schini
CBS 133181
ACCEPTED MANUSCRIPT
CFCC 51988T
D. schoeni
MFLU 15-1279
D. sclerotioides
CBS 296.67
T
T
KY852497
KY852513
KY852505
KY852509
KY852501
Yang et al. (2018)
KY964226
KY964109
-
KY964182
KY964139
KC343193
KC344161
KC343677
KC343919
KC343435
Dissanayake et al.
(2017a)
Gomes et al. (2013)
D. scobina
CBS 251.38
KC343195
KC344163
KC343679
D. sennae
CFCC 51636
T
KY203724
KY228891
-
CFCC 51634
T
KY203722
KY228889
-
T
KJ197274
KJ197254
-
D. sennicola
RI
PT
D. schisandrae
KC343921
KC343437
Gomes et al. (2013)
KY228885
KY228875
Yang et al. (2017b)
KY228883
KY228873
Yang et al. (2017b)
KJ197236
-
Thompson et al. (2015)
BRIP 55665a
D. siamensis
D. sojae
MFLUCC 10-0573a
CBS 139282T
JQ619879
KJ590719
JX275429
KJ610875
KJ659208
JX275393
KJ590762
KJ612116
Udayanga et al. (2012)
Udayanga et al. (2015)
D. spartinicola
CBS 140003T
KR611879
KR857695
KR857696
-
-
Crous et al. (2015c)
D. sterilis
CBS 136969T
KJ160579
KJ160528
MF418350
KJ160611
KJ160548
Lombard et al. (2014)
D. stewartii
CBS 193.36
FJ889448
-
-
GQ250324
-
Santos et al. (2010)
D. stictica
CBS 370.54
KC343212
KC344180
KC343696
KC343938
KC343454
Gomes et al. (2013)
D. subclavata
ICMP20663
KJ490630
KJ490451
KJ490572
KJ490509
-
Huang et al. (2015)
D. subordinaria
D. taoicola
CBS 101711
MFLUCC 16-0117T
KC343213
KU557567
D
M
AN
U
SC
D. serafiniae
KC344181
KU557591
KC343697
-
KC343939
KU557635
KC343455
-
D. tecomae
CBS 100547
T
KC344183
KC343699
KC343941
KC343457
MFLUCC 12-0777
T
KU712430
KU743977
-
KU749359
KU749345
Doilom et al. (2017)
MFLUCC 13-0471
T
KU712439
KU743986
-
KU749367
KU749354
Doilom et al. (2017)
D. tectonigena
MFLUCC 12-0767
T
KU712429
KU743976
-
KU749371
KU749358
Doilom et al. (2017)
D. terebinthifolii
CBS 133180
KC343216
KC344184
KC343700
KC343942
KC343458
Gomes et al. (2013)
KC153098
-
-
KC153089
-
Gao et al. (2014)
D. ternstroemia
EP
D. tectonendophytica
AC
C
D. tectonae
TE
KC343215
Gomes et al. (2013)
Dissanayake et al.
(2017c)
Gomes et al.(2013)
T
CGMCC 3.15183
T
D. thunbergii
MFLUCC 10-0756a
JQ619893
JX275449
-
JX275409
JX197440
Udayanga et al. (2012)
D. torilicola
MFLUCC 17-1051
T
KY964212
KY964096
-
KY964168
KY964127
D. toxica
CBS 534.93
KC343220
KC344188
KC343704
KC343946
KC343462
Dissanayake et al.
(2017a)
Gomes et al.(2013)
T
FFPRI420987
LC275192
LC275224
LC275216
LC275216
LC275200
Ando et al. (2017)
D. tulliensis
D. ueckerae
BRIP 62248a
FAU 656
KR936130
KJ590726
KR936132
KJ610881
KJ659215
KR936133
KJ590747
KJ612122
Crous et al. (2015e)
Huang et al. (2015)
D. undulata
CGMCC 3.18293
KX986798
KX999230
KX999269
KX999190
-
Gao et al. (2017)
KJ490587
KJ490408
KJ490529
KJ490466
-
Huang et al. (2015)
AF317578
KC344196
KC343712
GQ250326
KC343470
Gomes et al. (2013)
KJ869137
KJ869247
-
-
-
Crous et al. (2014c)
D. vaccinii
CGMCC3.17569
CBS 160.32
T
T
T
D. vangueriae
CBS 137985
D. vawdreyi
BRIP 57887a
D. velutina
CGMCC 3.18286
D. vexans
D. virgiliae
D. woodii
KR936126
KR936128
KX986790
KX999223
CBS 127.14
CBS 138788T
KC343229
KP247573
KC344197
KP247582
CBS 558.93
KC343244
T
KR936129
-
Crous et al. (2015e)
KX999182
-
Gao et al. (2017)
KC343713
-
KC343955
-
KC343471
-
KC344212
KC343728
KC343970
KC343486
Gomes et al.(2013)
Machingambi et al.
(2015)
Gomes et al. (2013)
KX999261
M
AN
U
D. unshiuensis
T
SC
D. toxicodendri
RI
PT
ACCEPTED MANUSCRIPT
CBS 148.27
KC343245
KC344213
KC343729
KC343971
KC343487
Gomes et al. (2013)
CGMCC 3.18282
T
KX986783
KX999216
KX999255
KX999175
-
Gao et al. (2017)
CGMCC 3.18289
T
KX986796
KX999228
KX999267
KX999188
KX999290
Gao et al. (2017)
D. yunnanensis
1
D
D. woolworthii
D. xishuangbanica
AC
C
EP
TE
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands;
CFCC: China Forestry Culture Collection Center, Beijing, China; CGMCC: Chinese General Microbiological Culture Collection Center, Beijing,
China; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute; DAOM: Plant Research Institute, Department of
Agriculture (Mycology), Ottawa, Canada; DAOMC: Canadian Collection of Fungal Cultures, Ottawa, Canada; FAU: Isolates in culture collection of
Systematic Mycology and Microbiology Laboratory; ICMP: International Collection of Micro-organisms from Plants, Landcare Research, Private
Bag 92170, Auckland, New Zealand; IFRDCC: International Fungal Research and Development Culture Collection; MFLU: Mae Fah Luang
University herbarium, Thailand; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand; LC: Working collection of Lei Cai,
T
housed at Institute of Microbiology, Chinese Academy of Sciences, Beijing, China. indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; tub2: partial β-tubulin gene; his3: partial histone H3 gene; tef1: partial elongation
factor 1-alpha gene; cal: partial calmodulin gene.
ACCEPTED MANUSCRIPT
Table 11. DNA barcodes of accepted Microdochium spp.
Isolates
1
GenBank accession numbers
rpb2
tub2
References
CBS 290.79
KP859014
KP859123
KP859077
Hernández-Restrepo et al. (2016a)
Mi. bolleyi
CBS 540.92
KP859010
KP859119
KP859073
Hernández-Restrepo et al. (2016a)
LT990657
LT990643
LT990610
Present study
KP859003
KP859112
KP859066
KP858999
KP859108
KP859062
Hernández-Restrepo et al. (2016a)
KU746690
-
KP859015
CBS 624.94
T
Mi. chrysanthemoides
CGMCC3.17929
Mi. fisheri
CBS 242.91T
T
T
Hernández-Restrepo et al. (2016a)
KU746781
Zhang et al. (2017)
KP859124
KP859078
Hernández-Restrepo et al. (2016a)
KP859016
KP859125
KP859079
Hernández-Restrepo et al. (2016a)
KP859001
KP859110
KP859064
Hernández-Restrepo et al. (2016a)
KP859111
KP859065
Hernández-Restrepo et al. (2016a)
KP859117
KP859071
Hernández-Restrepo et al. (2016a)
Mi. lycopodinum
CBS 122885
Mi. majus
CBS 741.79
Mi. neoqueenslandicum
CBS 108926T
KP859002
T
KP859008
M
AN
U
Mi. colombiense
CBS 109067
D
Mi. citrinidiscum
T
SC
Microdochium albescens
CPC 29378
CBS 116205
Mi. novae-zelandiae
CBS 143847
LT990655
LT990641
LT990608
Present study
CPC 29693
LT990656
LT990642
LT990609
Present study
KP859013
KP859122
KP859076
Hernández-Restrepo et al. (2016a)
KP859038
KP859147
KP859101
Hernández-Restrepo et al. (2016a)
Mi. phragmitis
CBS 285.71
Mi. seminicola
CBS 139951T
Mi. sorghi
CBS 691.96
KP859000
KP859109
KP859063
Hernández-Restrepo et al. (2016a)
CBS 269.76
T
KP859009
KP859118
KP859072
Hernández-Restrepo et al. (2016a)
CBS 623.77
T
KP858998
KP859107
KP859061
Hernández-Restrepo et al. (2016a)
AC
C
Mi. trichocladiopsis
EP
ET
TE
Mi. nivale
Mi. tainanense
1
ITS
2
RI
PT
Species
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Personal collection of Pedro Crous housed at the
Westerdijk Fungal Biodiversity Institute; CGMCC: China General Microbiological Culture Collection Center. T and ET indicate extype and ex-epitype strains.
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 RNA polymerase II second largest subunit gene; tub2:
partial β-tubulin gene.
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
x10
Gliocladiopsis sagariensis CBS 199.55
Cylindrocladiella clavata CBS 129563
Cylindrocladiella clavata CBS 129564
Cylindrocladiella humicola CBS 142777
Cylindrocladiella humicola CBS 142779
Cylindrocladiella reginae CBS 142782
ACCEPTED MANUSCRIPT
EP
TE
D
M
AN
U
SC
RI
PT
Cylindrocladiella parva CBS 114524
Cylindrocladiella stellenboschensis CBS 115611
Cylindrocladiella stellenboschensis CBS 110668
Cylindrocladiella pseudoparva CBS 129560
Cylindrocladiella pseudoparva CBS 113624
Cylindrocladiella lanceolata CBS 129565
Cylindrocladiella lanceolata CBS 129566
Cylindrocladiella lageniformis CBS 340.92
Cylindrocladiella lageniformis CBS 111060
Cylindrocladiella hahajimaensis PD684
Cylindrocladiella variabilis CBS 375.93
Cylindrocladiella variabilis CBS 129561
Cylindrocladiella australiensis CBS 129567
Cylindrocladiella australiensis CBS 129568
Cylindrocladiella longistipitata CBS 112953
Cylindrocladiella longistipitata CBS 116075
Cylindrocladiella nauliensis CBS 143791
Cylindrocladiella nauliensis CBS 143792
Cylindrocladiella ellipsoidea CBS 129572
Cylindrocladiella ellipsoidea CBS 129573
Cylindrocladiella thailandica CBS 129570
Cylindrocladiella thailandica CBS 129571
Cylindrocladiella horticola CBS 142785
Cylindrocladiella horticola CBS 142784
Cylindrocladiella queenslandica CBS 129575
Cylindrocladiella queenslandica CBS 129574
Cylindrocladiella pseudoinfestans CBS 114531
Cylindrocladiella pseudoinfestans CBS 114530
Cylindrocladiella kurandica CBS 129576
Cylindrocladiella kurandica CBS 129577
Cylindrocladiella brevistipitata CBS 142786
Cylindrocladiella infestans CBS 191.50
Cylindrocladiella infestans CBS 111795
Cylindrocladiella lateralis CBS 142788
Cylindrocladiella lateralis CBS 142787
Cylindrocladiella hawaiiensis CBS 129569
Cylindrocladiella hawaiiensis CBS 118704
Cylindrocladiella pseudohawaiiensis CBS 210.94
Cylindrocladiella pseudohawaiiensis CBS 115610
Cylindrocladiella viticola CBS 112897
Cylindrocladiella viticola CBS 114682
x4
Cylindrocladiella elegans CBS 338.92
Cylindrocladiella elegans CBS 110801
Cylindrocladiella novaezelandica CBS 486.77
Cylindrocladiella addiensis CBS143794
x4
Cylindrocladiella addiensis CBS143793
Cylindrocladiella addiensis CBS143795
Cylindrocladiella cymbiformis CBS 129553
Cylindrocladiella pseudocamelliae CBS 129556
Cylindrocladiella pseudocamelliae CBS 129555
x4
Cylindrocladiella microcylindrica CBS 111794
Cylindrocladiella natalensis CBS 114944
x4
Cylindrocladiella natalensis CBS 114943
Cylindrocladiella terrestris CBS 142790
Cylindrocladiella terrestris CBS 142789
Cylindrocladiella peruviana IMUR1843
Cylindrocladiella peruviana CBS 114697
Cylindrocladiella longiphialidica CBS 129558
Cylindrocladiella longiphialidica CBS 129557
Cylindrocladiella camelliae CBS 114891
Cylindrocladiella camelliae IMI 346845
Cylindrocladiella nederlandica CBS 152.91
Cylindrocladiella nederlandica CBS 146.94
0.2
AC
C
x10
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
CBS 121124T Corylus sp. China
Diaporthella corylina
Diaporthe eleagni
D. saccarata
D. oncostoma
D. incospicua
D. anacardii
D. cytosporella
D. baccae
D. foeniculina
D. hongkongensis
D. pseudophoenicicola
D. pseudomangifaerae
D. arengae
D. perseae
D. arecae
D. decedens
D. citri
D. citrichinensis
D. heterophyllae sp. nov.
D. neilliae
D. alnea
D. helicis
D. pulla
AC
C
EP
TE
D
M
AN
U
SC
CBS 504.72 Eleangus sp. Netherlands
CBS 116311T Protea repens South Africa
1/100
CBS 589.78 Robinia pseudoacacia
FranceMANUSCRIPT
0.73/100
ACCEPTED
T
CBS 133813 Maytenus ilicifolia Brazil
CBS 720.97T Anacardium occidentale Eastern Africa
0.94/CBS 137020 T Citrus limon Spain
0.89
CBS 136972 T Vaccinium corymbosum Italy
1/100
CBS 187.27 Camellia sinensis Italy
1/100
CBS 111553T Foeniculum vulgare Spain
1/98
CBS 111554 Foeniculum vulgare Portugal
0.99/75 CBS 123208 Foeniculum vulgare Portugal
CBS 123209 Foeniculum vulgare Portugal
CBS 115448 T Dichroa febrifuga China
1/100
CBS 462.69T Phoenix dactylifera Spain
CBS 101339 T Mangifera indica Dominican Republic
1/100
CBS 114979 T Arenga engleri Hong Kong
1/81
1/CBS 151.73 Persea gratissima Netherlands
CBS 161.64T Areca catechu India
1/59
1/- CBS 535.75 Citrus sp. Suriname
CBS 109772 Coryllus avellana Austria
CBS 134237 Citrus reticulata China
1/100
CBS 134239T Citrus sinensis USA
0.76/99
1/99 CBS 135422 Citrus sp. USA
1/100
CBS 134242T Citrus sp. China
CPC 26215 T Acacia heterophylla La Réunion (France)
1/52
CBS 144.27T Spiraea sp. USA
CBS 146.46T Alnus sp. Netherlands
1/CBS 138596T Hedera helix France
1/100
CBS 338.89T Hedera helix Yugoslavia
1/99
0.81/CBS 200.39 Laurus nobilis Germany
1/92
CBS 116953 Pyrus pyrifolia New Zealand
1/91
CBS 113470 Castanea sativa Australia
CBS 587.79 Pinus pentaphylla Japan
0.67/CBS 439.82T Cotoneaster sp. Scotland
CBS 101742 Fraxinus sp. Netherlands
1/100
1/93
CBS 138594 Ulmus laevis Germany
1/81
CBS 139.27T Celastrus sp. USA
CBS 121004T Junglans sp. USA
1/90
CBS 495.72T Betula alleghaniensis Canada
0.81/1/89
CBS 160.32T Vaccinium macrocarpon USA
1/96
CBS 118571 Vaccinium corymbosum USA
1/100
0.05
0.98/59 CBS 122114 Vaccinium corymbosum USA
RI
PT
4X
D. eres
D. celastrina
D. bicincta
D. alleghaniensis
D. vaccinii
D. carpini
D. detrusa
D. impulsa
D. fibrosa
D. ampelina
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
CBS 114437 Carpinus betulus Sweden
CBS 109770 Barberis vulgaris Austria
1/100
ACCEPTED
MANUSCRIPT
CBS
114434 Sorbus aucuparia
Sweden
0.99/CBS 109751 Rhamnus cathartica Austria
1/0.8/CBS 111888 Vitis vinifera USA
0.99/1/100 CBS 114016 T Vitis vinifera France
CBS 109745 T Ulmus glabra Austria
CBS 129521 T Acacia retinodes Australia
CBS 126679T Prunus dulcis Portugal
1/100
1/100
0.89/CBS 136969 T Vaccinium corymbosum Italy
0.98/CBS 534.93 T Lupinus angustifolius Australia
4X
0.83
BRIP 54801 T Nothofagus cunninghamii Australia
1/CBS 111886 T Vitis vinifera Australia
1/100
CBS 113201 Vitis vinifera Portugal
1/100
1/100 CBS 114436 Sambucus cf. racemosa Sweden
CBS 134240 T Citrus unshiu China
CBS 117167 Aspalathus linearis South Africa
1/100 CBS 187.87 Helianthus annuus Italy
1/100
CBS 133812 T Schinus terebinthifolius Brazil
1/99
CBS 127271T Glycine max Croatia
0.99/76
CBS 111592 T Heracleum sphondylium Austria
1/100
0.99/79
CBS 117499 T Aspalathus linearis South Africa
CBS 133180 T Schinus terebinthifolius Brazil
1/100 1/100
CPC 26646T Euclea racemosa South Africa
1/99
CBS 133181 T Schinus terebinthifolius Brazil
CBS 344.94 Helianthus annuus
1/98
1/100 CBS 592.81 T Helianthus annuus Serbia
CBS 507.78T Cucumis melo
0.78/CGMCC3 17569 T Citrus unshiu China
1/95
CBS 199.39 Unknown Italy
0.94/1/100 CBS 230.52 T Citrus sinensis Suriname
1/75
CBS 133811 T Schinus terebinthifolius
1/100
CBS 139282T Glycine max USA
1/100 CBS 116019 Caperonia palustris USA
CBS 113425 Olearia cf. rani New Zealand
0.99/94
0.05
1/85 CBS 127465 Actinidia chinensis New Zealand
D. perjuncta
D. acaciigena
D. amygdali
D. sterilis
D. toxica
D. nothofagi
D. australafricana
D. rudis
D. citriasiana
D. ambigua
D. infecunda
D. novem
D. angelicae
D. cuppatea
D. terebinthifolii
D. racemosae sp. nov.
D. schini
D. helianthi
D. melonis
D. unshiuensis
D. infertilis
D. endophytica
D. sojae
D. phaseolorum
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
ACCEPTED MANUSCRIPT
CBS 584.71
100
CBS 145.57T
Dichotomophthora lutea
CBS 585.71
100
100
CBS 518.78
100
CPC 33016T
100
RI
PT
CBS 132.81
Dichotomophthora basellae sp. nov.
CBS 174.35PT
Dichotomophthora portulacae
CBS 149.94T
Dichotomophthora brunnea sp. nov.
M
AN
U
BRIP 14541
SC
100
100
CBS 239.48
AC
C
EP
TE
D
0.004
Curvularia portulacae
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
CPC 26020
Gaeumannomyces graminis
ACCEPTED MANUSCRIPT
100
CBS 141390T
Gaeumannomyces oryzicola
97
CBS 141387T
Gaeumannomyces australiensis
CBS 141377T
Gaeumannomyces californicus
100
CBS 141379T
Gaeumannomyces fusiformis
CBS 352.93T
Gaeumannomyces graminicola
CBS 141378T
Gaeumannomyces floridanus
90
RI
PT
100
99
CBS 235.32T
Gaeumannomyces oryzinus
CBS 296.53T
SC
Gaeumannomyces radicicola
98
CBS 350.77T
Gaeumannomyces hyphopodioides
M
AN
U
100
BRIP 60376A
Gaeumannomyces wongoonoo
79
CBS 141394T
Gaeumannomyces setariicola
100
CBS 903.73T
94
CBS 141400T
CBS 905.73
CPC 26258ET
TE
89 94
D
92
CPC 26057T
98
EP
100
CBS 109354T
AC
C
0.03
Gaeumannomyces tritici
Gaeumannomyces avenae
Gaeumannomyces glycinicola
Gaeumannomyces ellisiorum
CM12M9T
CBS 125863T
Gaeumannomyces walkeri
Gaeumannomyces amomi
CBS 387.81T
100
Gaeumannomyces arxii
Pseudophialophora eragrostis
Falciphora oryzae
AC
C
EP
TE
D
M
AN
U
SC
RI
PT
ACCEPTED MANUSCRIPT
3x
Cryphonectria
parasitica
ACCEPTED
MANUSCRIPT
85/0.99
AC
C
EP
TE
D
89/0.95
0.05
Harknessia pilularis sp. nov.
Harknessia corymbiae sp. nov.
Harknessia syzygii
Harknessia eucalyptorum
Harknessia renispora
Harknessia pellitae
Harknessia banksiae-repens
Harknessia banksiae
Harknessia banksiigena
Harknessia communis
RI
PT
90/0.99
Harknessia rhabdosphaera
Harknessia fusiformis
SC
9x
M
AN
U
81/-
CBS 122373
100/1 CPC 33356
CPC 33218T
CPC 33289T
CBS 111124T
100/1 CBS 113620
CBS 111115T
CBS 153.71IsoT
CBS 142543T
CBS 142541T
CBS 142539T
96/0.99
73/CBS 142540T
CBS 142538T
CPC 11124
100/1
90/1
CBS 110785T
CBS 120030T
CBS 115647T
98/1
CBS 132127ET
99/1
CBS 132128ET
CBS 113074
CBS 110729
CBS
110729
99/87/1 CPC 16277T
CPC 13596
100/1
CBS 132119T
CBS 143913T
95/0.96
CBS 142544T
96/1
100/1 CPC 17209
CBS 132125T
T
100/1 CBS 132121
CPC 17113
CBS 142542T
92/1
CBS 115648
T
96/1 CBS 132124
CPC 13001
95/1
CBS 114811
100/1 CBS 111122
CPC 14924
99/1
CBS 111578T
93/1 CPC 13643
CBS 342.97
97/1
CBS 114877T
CBS 136426T
CBS 137228ET
86/CBS 120033T
86/0.95
CPC 17642
CBS 111829T
CBS 115061
99/1
CBS 112618T
72/CBS 111830
96/1
CBS 112620
100/1 CBS 775.97T
CBS 143914T
100/1 CPC 30174
Harknessia ipereniae
Harknessia viterboensis
Harknessia spermatoidea
Harknessia weresubiae
Harknessia uromycoides
Harknessia kleinzeeina
Harknessia australiensis
Harknessia bourbonica sp. nov.
Harknessia malayensis
Harknessia ravenstreetina
Harknessia ellipsoidea
Harknessia platyphyllae
Harknessia karwarrae
Harknessia pseudohawaiiensis
Harknessia hawaiiensis
Harknessia globispora
Harknessia eucalypti
Harknessia molokaiensis
Harknessia proteae
Harknessia arctostaphyli
Harknessia gibbosa
Harknessia capensis
Harknessia protearum
Harknessia leucospermi
Harknessia cupressi sp. nov.
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CMW 9986
ACCEPTED
MANUSCRIPT
100
Huntiella moniliformopsis
CMW 13013
Huntiella tribiliformis
100
CMW 10134
Huntiella moniliformis
100
Huntiella sublaevis
CMW 22449
CMW 28932
Huntiella bhutanensis
RI
PT
CMW 8217
CMW 17300
Huntiella savannae
Huntiella oblonga
CMW 23803
100
SC
CMW 30855
CMW 25911
M
AN
U
86
CMW 15245
CMW 24658
CMW 21109
79
TE
D
CMW 21117
AC
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0.04
Huntiella tyalla
CMW 21092
CMW 21106
Huntiella decipiens
Huntiella salinaria
Huntiella ceramica
Huntiella chinaeucensis
Huntiella sumatrana
Huntiella microbasis
Huntiella abstrusa sp. nov.
Huntiella inquinans
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Pyricularia grisea CBS 128304
96/1
Pyricularia penniseticola ML0031T
ACCEPTED MANUSCRIPT
83/0.99
-/1
100/1
96/0.97
Pyricularia oryzae CBS 255.38
Pyricularia graminis-tritici URM7380T
Pyricularia angulata NBRC 9625
Pyricularia
Pyricularia urashimae CBS 142117T
Pyricularia zingibericola RN0001
85/1
Pyricularia ctenantheicola GR0002T
100/1
Pyriculariomyces asari CPC 27442
Pyriculariomyces gen. nov.
Pyriculariomyces asari CPC 27444T
Neopyricularia commelinicola CBS 128308T
78/0.99
Neopyricularia
Proxypiricularia
Proxipyricularia zingiberis CBS 132195
Bambusicularia
SC
Bambusicularia brunnea CBS 133599T
91/0.99
RI
PT
Pyricularia pennisetigena ML0036T
Utrechtiana roumeguerei CBS 128780
100/1
T
Utrechtiana
Utrechtiana arundinacea CPC 33994ET
M
AN
U
Pseudopyricularia cyperi CBS 133595T
71/0.97
100/1
Pseudopyricularia higginsii CBS 121934
100/1
79/0.98
Pseudopyricularia hagahagae CPC 25635T
Pseudopyricularia
Pseudopyricularia bothriochloae CBS 136427T
Pseudopyricularia kyllingae CBS 133597T
80/-
Macgarvieomyces juncicola CBS 610.82
Macgarvieomyces luzulae CPC 32458ET
D
99/1
100/1
Macgarvieomyces luzulae CPC 31571
Macgarvieomyces
Macgarvieomyces luzulae CPC 31555
TE
100/1
Macgarvieomyces borealis CBS 461.65T
AC
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Xenopyricularia zizaniicola CBS 133593NT
0.07
Barretomyces
Barretomyces calatheae CBS 129274
Xenopyricularia
Bussabanomyces longisporus CBS 125232T
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3x
Cladosporium tenuissimum CBS 125995ET
ACCEPTED MANUSCRIPT
CPC 33937
-/1
CBS 161.74ET
Metulocladosporiella musae
CBS 113863
100/1
M. malaysiana sp. nov.
CBS 143919T
M. samutensis sp. nov.
CBS 143921T
M. musigena sp. nov.
CBS 143920T
90/1
RI
PT
3x
M. chiangmaiensis sp. nov.
CBS 143918T
CPC 32970
100/1
CBS 113862
100/1
85/0.98
CBS 113860
M
AN
U
73/0.99
SC
CBS 113864
76/0.99
CBS 113865
CPC 32807
94/0.98
CPC 32849
D
CPC 18124
AC
C
0.05
EP
TE
71/1
-/1
CBS 113873
CBS 113861
CBS 110960T
CBS 110962
99/1
CBS 110964
M. musicola
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ACCEPTED MANUSCRIPT
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ACCEPTED MANUSCRIPT
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ACCEPTED MANUSCRIPT
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ACCEPTED MANUSCRIPT
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CBS 741.79
ACCEPTED MANUSCRIPT
Microdochium majus
CBS 116205T
Microdochium nivale
100
Microdochium colombiense
CBS 624.94T
82
CPC 29378
100
Microdochium bolleyi
98
100
RI
PT
CBS 540.92
CPC 29376T
Microdochium novae-zelandiae sp. nov.
CPC 29693
CBS 139951T
Microdochium seminicola
SC
100
99
100 100
CBS 691.96
100
CBS 269.76T
CBS 285.71ET
CBS 122885T
EP
100
AC
C
CBS 242.91T
100
0.06
D
TE
100
Microdochium citrinidiscum
Microdochium sorghi
Microdochium taiwanense
Microdochium trichocladiopsis
CBS 623.77T
CBS 108926T
M
AN
U
CBS 109067T
100
Microdochium albescens
CBS 290.79
100
Microdochium neoqueenslandicum
Microdochium phragmitis
Microdochium lycopodinum
Microdochium fisheri
Thamnomyces dendroidea CBS 123578
Xylaria polymorpha MUCL 49884
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CBS 496.80T
Oculimacula anguioides
ACCEPTED MANUSCRIPT
CBS 495.80T
2x
-/1
Oculimacula acuformis comb. nov.
CBS 114730
Oculimacula aestiva
95/-
CBS 128.31
Oculimacula yallundae
CBS 110655NT
RI
PT
75/0.98
CBS 494.80
2x
SC
Cadophora melinii AF0832052
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0.02
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ACCEPTED MANUSCRIPT BRIP 65178
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BRIP 65179
BRIP 65170
BRIP 65177
BRIP 57989
Pa. chlamydocopiosa
BRIP 65176
BRIP 65174
BRIP 57988
BRIP 65168
BRIP 65173
BRIP 63682
BRIP 63685
Pa. vinacea
BRIP 63684
BRIP 63683
BRIP 65169
Pa. pye
BRIP 65171
Pa. rhaphiolepidis
CBS 142524
CBS 522.66
Pa. chrysanthemicola
CBS 172.70
CBS 111.79
Pa. radicina
CBS 102875
CPC 11361
Pa. dioscoreae
CPC 11355
CBS 135100
CBS 379.67
CBS 550.70
CBS 368.91
Pa. fimeti
CBS 164.31
CBS 119754
CBS 258.68
CBS 170.70
Neosetophoma samarorum CBS 138.96
AC
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0.009
AC
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CBS 211.97
CBS 110212 Phaeoacremonium fraxinopennsylvanicum
0.98/89
CBS 101585T
0.85/65
ICMP 17037T
Phaeoacremonium occidentale
1/99
T
CBS 123037
Phaeoacremonium croatiense
0.99/98
T
CBS 123036
Phaeoacremonium hungaricum
1/100 PARC392
Phaeoacremonium canadense
0.99/92
DAOM 242366T
ACCEPTED MANUSCRIPT
CBS
114512
1/100
CBS 110157
Phaeoacremonium leptorrhynchum
CBS 110156
MFLUCC 14-1130
1/100
MFLUCC 14-1125
Phaeoacremonium tectonae
T
MFLUCC 13-0707
1/89
CBS 777.83T
Phaeoacremonium argentinense
1/100
T
CBS 142704
1/100
Phaeoacremonium oleae
CBS 142701
0.99/86
ICMP 17038
1/100
ICMP 16988T
Phaeoacremonium globosum
ICMP 16987
1/100
ICMP 17421T
Phaeoacremonium armeniacum
0.98/95
1/100 STE-U 6364
Phaeoacremonium africanum
CBS 120863T
0.99/83
1/100 STE-U 5968
1/97
Phaeoacremonium prunicolum
CBS 120858T
CBS 120857T
Phaeoacremonium griseo-olivaceum
1/100
CBS 142715
Phaeoacremonium spadicum
CBS 142711T
0.75/59
Pir-1
1/100
Phaeoacremonium iranianum
CBS 117114
T
CBS 101357
1/100
CBS 246.91T
0.89/69
1/100
Phaeoacremonium minimum
CBS 100397
1/100
CBS 110703
CBS 123033T
Phaeoacremonium tuscanicum
0.72/100 PARC273
Phaeoacremonium roseum
1/97 DAOM 242365T
CBS 114992T
1/100
Phaeoacremonium angustius
1/100
CBS 114991
0.87/62
CBS 113065
1/78
1/81
CBS 101738T
Phaeoacremonium viticola
CBS 101737
CBS 142700
1/100
Phaeoacremonium longicollarum
CBS 142699T
CBS 114994
1/100
CBS 114993
Phaeoacremonium austroafricanum
T
1/100
CBS 112949
1/95
1/100
Phaeoacremonium pseudopanacis
CBS 142101T
0.99/100
CBS 142717
Phaeoacremonium geminum
1/98
CBS 142713T
Phaeoacremonium gamsii
CBS 142712T
T
Phaeoacremonium theobromatis
CBS 111586
T
CBS 120862
Phaeoacremonium pallidum
1/100 CBS 142689
1/97
Phaeoacremonium album
1/98
CBS 142688T
CBS 142694T
Phaeoacremonium bibendum
1/100
T
CBS 142708
Phaeoacremonium rosicola
Psp-2
1/100
Phaeoacremonium amygdalinum
Psp-1
T
CBS 128570
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0.88/94
1/100
CBS 137498T
Phaeoacremonium santali
A4
A37
0.93/57
CBS 566.97
Phaeoacremonium griseorubrum
0.79/65 CBS 111657T
1/99
CBS 113597T
1/99
Phaeoacremonium scolyti
CBS 113593
CBS 112585
1/99
Phaeoacremonium amstelodamense
CBS 110627T
CBS 142687
1/100
ACCEPTED MANUSCRIPT
Phaeoacremonium pravum sp. nov.
CBS 142686T
0.95/91 1/100 CBS 142707T
Phaeoacremonium proliferatum
CBS 142706
0.88/48
CBS 113592
1/100
Phaeoacremonium australiense
1/100
CBS 113589T
1/100
CBS 113587
0.99/85
Phaeoacremonium subulatum
CBS 113584T
1/96
CBS 142698
Phaeoacremonium junior
1/97 CBS 142697T
1/98 CMM 4334
1/100
Phaeoacremonium nordesticola
CMM 4312T
1/100
A34
Phaeoacremonium luteum
1/86
CBS 137497T
1/100
CBS 110573T
Phaeoacremonium tardicrescens
1/100 CBS 137764
1/96 CBS 137763T
Phaeoacremonium italicum
0.99/80
CBS 113590
CBS 408.78
0.98/100
Phaeoacremonium alvesii
T
1/100 CBS 110034
CBS 498.94T
0.98/83
Phaeoacremonium rubrigenum
0.97/97 CBS 112046
Phaeoacremonium paululum
CBS 142705T
0.62/84 CBS 860.73T
1/100
CBS 514.82
Phaeoacremonium parasiticum
0.8/48
CBS 113585
1/100 CBS 142691T
Phaeoacremonium aureum
1/100
CBS 142690
1/100 STE-U 6366
Phaeoacremonium fuscum
1/100
CBS 120856T
CBS 113595
0.81/57
CBS 110119
Phaeoacremonium venezuelense
1/100 CBS 651.85T
Pm4
1/100
0.88/45
Phaeoacremonium cinereum
Pm2
1/100
CBS 123909T
CBS 123910T
Phaeoacremonium hispanicum
1/100
1/100 CBS 117115T
Phaeoacremonium vibratile
1/97 CBS 391.71T
1/100
Phaeoacremonium inflatipes
CBS 113273
CBS 166.75
0.7/99 CBS 110118
1/100
Phaeoacremonium krajdenii
CBS 109479T
0.89/40
CBS 110368
1/100
CBS 142710T
Phaeoacremonium meliae
CBS 142709
1/100
CBS 694.88
Phaeoacremonium sphinctrophorum
CBS 337.90T
1/100
CBS 123035
Phaeoacremonium sicilianum
CBS 123034T
0.95/79
Jattaea algeriensis CBS 120871
Calosphaeria africana CBS 120870
Pleurostoma richardsiae CBS 270.33T
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1/99
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1/100
0.2
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ACCEPTED MANUSCRIPT
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Phyllosticta podocarpi
Phyllosticta carissicola
Phyllosticta pseudotsugae
Phyllosticta owaniana
Phyllosticta vaccinii
Phyllosticta vacciniicola
Phyllosticta philoprina
Phyllosticta ilicis-aquifolii
Phyllosticta mangifera-indica
Phyllosticta mimusopisicola
Phyllosticta brazilianiae
Phyllosticta beaumarisii
Phyllosticta partricuspidatae
Phyllosticta parthenocissi
Phyllosticta ampelicida
Phyllosticta vitis-rotundifoliae
Phyllosticta musarum
Phyllosticta maculata
Phyllosticta cavendishii
Phyllosticta aristolochiicola
Phyllosticta cordylinophila
Phyllosticta paracapitalensis
Phyllosticta capitalensis
Phyllosticta alliacea
Phyllosticta fallopiae
Phyllosticta schimae
Phyllosticta schimicola
Phyllosticta carochlae
Phyllosticta ardisiicola
Phyllosticta styracicola
Phyllosticta eugeniae
Phyllosticta aloeicola
Phyllosticta podocarpicola
Phyllosticta paxistimae
Phyllosticta pachysandricola
Phyllosticta leucothoicola
Phyllosticta cornicola
Phyllosticta gaultheriae
Phyllosticta neopyrolae
Phyllosticta rubella
Phyllosticta yuccae
Phyllosticta hamamelidis
Phyllosticta sphaeropsoidea
Phyllosticta minima
Phyllosticta ligustricola
Phyllosticta abieticola
Phyllosticta telopeae
Phyllosticta cruenta
Phyllosticta hubeiensis
Phyllosticta persooniae sp. nov.
Phyllosticta foliorum
Phyllosticta hymenocallidicola
Phyllosticta citriasiana
Phyllosticta citrimaxima
Phyllosticta paracitricarpa
Phyllosticta citricarpa
Phyllosticta bifrenariae
Phyllosticta catimbauensis
Phyllosticta rhaphiolepidis
Phyllosticta ericarum
Phyllosticta citribraziliensis
Phyllosticta concentrica
Phyllosticta kerriae
Phyllosticta spinarum
Phyllosticta citrichinaensis
Phyllosticta elongata
Phyllosticta cussonia
Phyllosticta iridigena sp. nov.
Phyllosticta hypoglossi
Phyllosticta aspidistricola
Phyllosticta hostae
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CBS 111646
CPC 25665T
CBS 111649
70/CBS 776.97ETACCEPTED MANUSCRIPT
97/1
ATCC 46255ET
CBS 136062T
99/1
CBS 587.69
CGMCC 3.14358T
CBS 136061T
CBS 138899T
CBS 129060T
CBS 535.87T
79/0.99
NBRC 9466T
CBS 111645
72/0.98
ATCC 200578NT
CGMCC 3.17322T
IsoET
93/0.97
100/1 BRIP 55434
CBS 132581ET
88/0.98
BRIP 55420IsoT
T
BRIP 53316a
CBS 136244NT
CPC 26517T
CBS
128856ET
99/1
T
90/0.95 MUCC 0014
T
MUCC 0113
100/1
CGMCC 3.14354T
CGMCC 3.17319T
CGMCC 3.17317T
MUCC 0031T
CGMCC 3.14985T
CBS 445.82
CBS 136058T
CBS 728.79T
CBS 112527T
MUCC 124T
CBS
136073T
73/CBS 111639
CBS 447.70T
CBS 134750T
99/1 CBS 111635T
CBS 117136
MUCC 149
CBS 756.70
90/CBS 585.84NT
MUCC 0024T
CBS 112067T
CBS 777.97T
CBS 858.71
CGMCC 3.14986T
CBS 143409T
CBS 447.68NT
CBS 131309T
75/0.95
99/1 CBS 120486T
CBS 136059T
CBS 141357T
100/1
100/1 CBS 127454ET
CBS 128855T
URM 7672T
-/0.96
MUCC 0432T
98/1
T
100/1 CBS 132534
75/0.98
CBS 100098T
CBS 937.70ET
-/0.95
MUCC 0017T
CBS 292.90
98/1
CBS 130529T
CBS 126.22T
76/CBS 136060ET
CBS 143410T
80/CBS 434.92NT
76/- 95/1
MUCC 0010T
CGMCC 3.14355T
Peyronellaea obtusa CMW8232
99/1
0.05
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ACCEPTED MANUSCRIPT
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ACCEPTED MANUSCRIPT
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ACCEPTED MANUSCRIPT
AC
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ACCEPTED MANUSCRIPT
AC
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ACCEPTED MANUSCRIPT
AC
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ACCEPTED MANUSCRIPT
AC
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ACCEPTED MANUSCRIPT
65
MFLUCC 13-0735T
ACCEPTED MANUSCRIPT
CPC 27468
76
70
Wojnowiciella dactylidis
CPC 30353
CPC 33929
CPC 32741
CBS 141297T
RI
PT
78
Wojnowiciella cissampeli
MFLUCC 13-0737T
Wojnowiciella lonicerae
MFLUCC 13-0402T
Wojnowiciella spartii
Wojnowiciella eucalypti
MFLUCC 12-0733T
M
AN
U
CBS 115684T
SC
CBS 139904T
100
100
AC
C
EP
TE
D
0.02
Wojnowiciella viburni
Wojnowiciella leptocarpi
CBS 120249 Phaeosphaeria caricis
CBS 536.77T
Septoriella hirta