Genera of Phytopathogenic Fungi GOPHY3

available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 94: 1–124 (2019).
Genera of phytopathogenic fungi: GOPHY 3
Y. Marin-Felix1,2*, M. Hernandez-Restrepo1, I. Iturrieta-Gonzalez2, D. García2, J. Gene2, J.Z. Groenewald1, L. Cai3, Q. Chen3,

W. Quaedvlieg4, R.K. Schumacher5, P.W.J. Taylor6, C. Ambers7, G. Bonthond1,8, J. Edwards9,10, S.A. Krueger-Hadfield11,
J.J. Luangsa-ard12, L. Morton13, A. Moslemi6, M. Sandoval-Denis1,14, Y.P. Tan15,16, R. Thangavel17, N. Vaghefi18,
R. Cheewangkoon19, and P.W. Crous1,20,21*
1
Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; 2Mycology Unit, Medical School and IISPV, Universitat Rovira i Virgili, Sant
Llorenç 21, 43201, Reus, Spain; 3State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing, 100101, China; 4Royal Van Zanten,
P.O. Box 265, 1430 AG, Aalsmeer, The Netherlands; 5Hölderlinstraße 25, 15517, Fürstenwalde/Spree, Germany; 6Faculty of Veterinary and Agricultural Sciences,
University of Melbourne, Melbourne, VIC, 3010, Australia; 7P.O. Box 631, Middleburg, VA, 20118, USA; 8Benthic Ecology, GEOMAR Helmholtz Centre for Ocean
Research Kiel, Hohenbergstraße 2, 24105, Kiel, Germany; 9Agriculture Victoria Research, Department of Jobs, Precincts and Regions, AgriBio Centre, Bundoora,
Victoria, 3083, Australia; 10School of Applied Systems Biology, La Trobe University, Bundoora, Victoria, 3083, Australia; 11Department of Biology, University of
Alabama at Birmingham, 1300 University Blvd, CH464, Birmingham, AL, 35294, USA; 12Plant Microbe Interaction Research Team, Integrative Crop Biotechnology
and Management Research Group, Bioscience and Biotechnology for Agriculture, NSTDA 113, Thailand Science Park Phahonyothin Rd., Khlong Nueng, Khlong
Luang, Pathum Thani, 12120, Thailand; 13P.O. Box 5607, Charlottesville, VA, 22905, USA; 14Faculty of Natural and Agricultural Sciences, Department of Plant
Sciences, University of the Free State, P.O. Box 339, Bloemfontein, 9300, South Africa; 15Department of Agriculture and Fisheries, Biosecurity Queensland,
Ecosciences Precinct, Dutton Park, 4012, QLD, Australia; 16Microbiology, Department of Biology, Utrecht University, Utrecht, Netherlands; 17Plant Health and
Environment Laboratory, Ministry for Primary Industries, P.O. Box 2095, Auckland, 1140, New Zealand; 18Centre for Crop Health, University of Southern
Queensland, Queensland, 4350, Australia; 19Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, 50200,
Thailand; 20Department of Biochemistry, Genetics & Microbiology, Forestry & Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, South Africa;
21
Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB, Wageningen, The Netherlands
*Correspondence: Y. Marin-Felix, y.marin@wi.knaw.nl; P.W. Crous, p.crous@wi.knaw.nl
Abstract: This paper represents the third contribution in the Genera of Phytopathogenic Fungi (GOPHY) series. The series provides morphological descriptions,
information about the pathology, distribution, hosts and disease symptoms for the treated genera, as well as primary and secondary DNA barcodes for the currently
accepted species included in these. This third paper in the GOPHY series treats 21 genera of phytopathogenic fungi and their relatives including: Allophoma, Alternaria,
Brunneosphaerella, Elsinoe, Exserohilum, Neosetophoma, Neostagonospora, Nothophoma, Parastagonospora, Phaeosphaeriopsis, Pleiocarpon, Pyrenophora,
Ramichloridium, Seifertia, Seiridium, Septoriella, Setophoma, Stagonosporopsis, Stemphylium, Tubakia and Zasmidium. This study includes three new genera, 42 new
species, 23 new combinations, four new names, and three typifications of older names.
Key words: DNA barcodes, Fungal systematics, New taxa.

Taxonomic novelties: New genera: Arezzomyces Y. Marín & Crous, Globoramichloridium Y. Marín & Crous, Wingfieldomyces Y. Marín & Crous; New species:
Allophoma pterospermicola Q. Chen & L. Cai, Alternaria aconidiophora Iturrieta-Gonzalez, Dania García & Gene, Alternaria altcampina Iturrieta-Gonzalez, Dania
García & Gene, Alternaria chlamydosporifera Iturrieta-Gonzalez, Dania García & Gene, Alternaria curvata Iturrieta-Gonzalez, Dania García & Gene, Alternaria fimeti
Studies in Mycology
Iturrieta-Gonzalez, Dania García & Gene, Alternaria inflata Iturrieta-Gonzalez, Dania García & Gene, Alternaria lawrencei Iturrieta-Gonzalez, Dania García & Gene,
Alternaria montsantina Iturrieta-Gonzalez, Dania García & Gene, Alternaria pobletensis Iturrieta-Gonzalez, Dania García & Gene, Alternaria pseudoventricosa
Iturrieta-Gonzalez, Dania García & Gene, Brunneosphaerella roupeliae Crous, Elsinoe picconiae Crous, Elsinoe veronicae Crous, Thangavel & Y. Marín,
Neosetophoma aseptata Crous, R.K. Schumach. & Y. Marín, Neosetophoma phragmitis Crous, R.K. Schumach. & Y. Marín, Neosetophoma sambuci Crous, R.K.
Schumach. & Y. Marín, Neostagonospora sorghi Crous & Y. Marín, Parastagonospora novozelandica Crous, Thangavel & Y. Marín, Parastagonospora phragmitis
Crous & Y. Marín, Phaeosphaeriopsis aloes Crous & Y. Marín, Phaeosphaeriopsis aloicola Crous & Y. Marín, Phaeosphaeriopsis grevilleae Crous & Y. Marín,
Phaeosphaeriopsis pseudoagavacearum Crous & Y. Marín, Pleiocarpon livistonae Crous & Quaedvl., Pyrenophora avenicola Y. Marín & Crous, Pyrenophora
cynosuri Y. Marín & Crous, Pyrenophora novozelandica Y. Marín & Crous, Pyrenophora pseudoerythrospila Y. Marín & Crous, Pyrenophora sieglingiae Y. Marín &
Crous, Pyrenophora variabilis Hern.-Restr. & Y. Marín, Septoriella germanica Crous, R.K. Schumach. & Y. Marín, Septoriella hibernica Crous, Quaedvl. & Y. Marín,
Septoriella hollandica Crous, Quaedvl. & Y. Marín, Septoriella pseudophragmitis Crous, Quaedvl. & Y. Marín, Setophoma brachypodii Crous, R.K. Schumach. & Y.
Marín, Setophoma pseudosacchari Crous & Y. Marín, Stemphylium rombundicum Moslemi, Y.P. Tan & P.W.J. Taylor, Stemphylium truncatulae Moslemi, Y.P. Tan &
P.W.J. Taylor, Stemphylium waikerieanum Moslemi, Jacq. Edwards & P.W.J Taylor, Vagicola arundinis Phukhams., Camporesi & K.D. Hyde, Zasmidium thailandicum
Crous; New combinations: Arezzomyces cytisi (Wanas. et al.) Y. Marín & Crous, Globoramichloridium indicum (Subram.) Y. Marín & Crous, Phaeosphaeria
phoenicicola (Crous & Thangavel) Y. Marín & Crous, Pyrenophora poae (Baudys) Y. Marín & Crous, Pyrenophora wirreganensis (Wallwork et al.) Y. Marín & Crous,
Seiridium cupressi (Nattrass et al.) Bonthond, Sandoval-Denis & Crous, Seiridium pezizoides (de Not.) Crous, Septoriella agrostina (Mapook et al.) Y. Marín &
Crous, Septoriella artemisiae (Wanas. et al.) Y. Marín & Crous, Septoriella arundinicola (Wanas. et al.) Y. Marín & Crous, Septoriella arundinis (W.J. Li et al.) Y.
Marín & Crous, Septoriella bromi (Wijayaw. et al.) Y. Marín & Crous, Septoriella dactylidis (Wanas. et al.) Y. Marín & Crous, Septoriella elongata (Wehm.) Y. Marín
& Crous, Septoriella forlicesenica (Thambug. et al.) Y. Marín & Crous, Septoriella garethjonesii (Thambug. et al.) Y. Marín & Crous, Septoriella italica (Thambug.
et al.) Y. Marín & Crous, Septoriella muriformis (Ariyaw. et al.) Y. Marín & Crous, Septoriella rosae (Mapook et al.) Y. Marín & Crous, Septoriella subcylindrospora
(W.J. Li et al.) Y. Marín & Crous, Septoriella vagans (Niessl) Y. Marín & Crous, Wingfieldomyces cyperi (Crous & M.J. Wingf.) Y. Marín & Crous, Zasmidium
ducassei (R.G. Shivas et al.) Y. Marín & Crous; New names: Pyrenophora nisikadoi Y. Marín & Crous, Septoriella dactylidicola Y. Marín & Crous, Septoriella
neoarundinis Y. Marín & Crous, Septoriella neodactylidis Y. Marín & Crous; Typification: epitypification: Ascochyta chrysanthemi F. Stevens, Pestalotia unicornis
Cooke & Ellis, Rhynchosphaeria cupressi Nattrass et al.
Available online 13 June 2019; https://doi.org/10.1016/j.simyco.2019.05.001.
Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.

© 2019 Westerdijk Fungal Biodiversity Institute. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
1
MARIN-FELIX ET AL.
INTRODUCTION (VPRI). For fresh collections, we followed the procedures pre-

viously described in Waksman (1922), Crous et al. (1991) and
Genera of Phytopathogenic Fungi (GOPHY) is a series of papers Calduch et al. (2004). Colonies were transferred to different
with the main focus to provide a stable platform for the taxonomy media, i.e. cornmeal agar (CMA), 2 % malt extract agar (MEA),
of phytopathogenic fungi. All genera included here are associ- potato carrot agar (PCA), 2 % potato dextrose agar (PDA),
ated with plant disease, but note that many species treated are synthetic nutrient-poor agar (SNA), oatmeal agar (OA), water
not well-known plant pathogens, or Koch’s postulates remain to agar (WA) (Crous et al. 2019b), pine needle agar (PNA; Smith
be completed for them. This series links to a larger initiative et al. 1996), and incubated under different conditions to induce
known as the “The Genera of Fungi project” (www. sporulation. Requirements of media and conditions of incubation
GeneraOfFungi.org, Crous et al. 2014a, 2015a, Giraldo et al. are specified for each genus. Reference strains and specimens
2017), which aims to revise the generic names of all currently are maintained at ATCC, BRIP, CBS, CPC, CGMCC, FMR and
accepted fungi (Kirk et al. 2013). Specific aims were detailed by VPRI.
Marin-Felix et al. (2017), when this series was launched. One of Vegetative and reproductive structures were mounted in
the most important aims is to resolve generic and species 100 % lactic acid or Shear’s solution either directly from speci-
concepts of the fungi studied, since many taxa have been shown mens or from colonies sporulating on CMA, MEA, OA, PCA,
to represent species complexes, or to comprise poly- or para- PDA, PNA, SNA or WA. For cultural characterisation, isolates
phyletic genera (Crous et al. 2015b). Other issues to resolve were grown and incubated on different culture media and tem-
include the fact that type material for many genera and species peratures as indicated for each genus. Colour notations were
has not been designated or is missing, and that the vast majority rated according to the colour charts of Kornerup & Wanscher
of these taxa were described before the DNA era (Hibbett et al. (1978) for Alternaria, and Rayner (1970) for all other genera.
2011) and thus lack DNA barcodes (Schoch et al. 2012). Taxonomic novelties were deposited in MycoBank (www.
Therefore, another important aim is to generate DNA barcodes of MycoBank.org; Crous et al. 2004).
type species and type specimens in order to fix the application of
these names. Moreover, in cases where no type material has
DNA isolation, amplification and analyses
been preserved, taxa need to be recollected, epi- or neotypes
designated, and registered in MycoBank to ensure traceability of
Fungal DNA was extracted and purified directly from the colonies
the nomenclatural act (Robert et al. 2013). Finally, it is necessary
or host material as specified for each genus. Primers and pro-
to designate a single scientific name for fungi (Crous et al.
tocols for the amplification and sequencing of gene loci, and
2015b) for which sexual-asexual links have been resolved.
software used for phylogenetic analyses can be found in the
Two issues of GOPHY have already been published, in which
bibliographies provided for each genus. Phylogenetic analyses
41 genera were treated, including a total of two new genera, 46
consisted of Maximum-Likelihood (ML) and Bayesian Inference
new species, 15 new combinations and 10 typifications of older
(BI). ML was inferred as described in Hernandez-Restrepo et al.
names (Marin-Felix et al. 2017, 2019). In this third contribution, a
(2016b), or by using MEGA v. 6.0 (Tamura et al. 2013). BI was
further 21 genera are treated, resulting in the clarification of their
carried out as described by Hernandez-Restrepo et al. (2016b),
taxonomy and classification, and the introduction of three new
or by using MrBayes on XSEDE v. 3.2.6 on the CIPRES portal
genera, 42 new species, 23 new combinations, four new names
(www.phylo.org). Sequence data generated in this study were
and the typification of three older names.
deposited in GenBank and the alignments and trees in Tree-
For submissions to future issues in the GOPHY series, my-
BASE (http://www.treebase.org).
cologists are encouraged to contact Pedro Crous (p.crous@wi.
knaw.nl) to ensure there is no overlap with activities arising
from other research groups. Preference will be given to genera
that include novel species, combinations or typifications. Generic RESULTS
contributions published in each issue will also be placed in the
database displayed on www.plantpathogen.org. Allophoma Q. Chen & L. Cai, Stud. Mycol. 82: 162. 2015. Fig. 1.
Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
sporales, Didymellaceae.
MATERIAL AND METHODS
Type species: Allophoma tropica (R. Schneid. & Boerema) Q.
Chen & L. Cai, basionym: Phoma tropica R. Schneid. & Boer-
Isolates and morphological analysis
ema, Phytopathol. Z. 83: 361. 1975. Isotype and ex-isotype
strain: CBS H-7629, CBS 436.75 = DSM 63365.
Descriptions of the new taxa and typifications are based on
cultures obtained from the collection at the American Type DNA barcodes (genus): LSU, ITS.
Culture Collection, Manassas, Virginia, USA (ATCC), the
DNA barcodes (species): rpb2, tub2. Table 1. Fig. 2.
Queensland Plant Pathology Herbarium, Brisbane, Australia
(BRIP), the Westerdijk Fungal Biodiversity Institute, Utrecht, The Conidiomata pycnidial, globose to flask-shaped, ovoid, superfi-
Netherlands (CBS), the working collection of P.W. Crous (CPC), cial or (semi-)immersed, solitary or confluent, ostiolate, some-
housed at the Westerdijk Fungal Biodiversity Institute (WI), the times with an elongated neck; conidiomatal wall
Chinese General Microbiological Culture Collection Center, pseudoparenchymatous, multi-layered. Conidiogenous cells
Beijing, China (CGMCC), the Facultat de Medicina i Ciencies de phialidic, hyaline, smooth-walled, ampulliform to doliiform,
la Salut, University Rovira i Virgili, Reus, Spain (FMR), and the sometimes flask-shaped or isodiametric. Conidia hyaline,
Victorian Plant Pathogen Herbarium, Bundoora, Australia smooth- and thin-walled, aseptate or 1-septate, variable in shape
2
GENERA OF PHYTOPATHOGENIC FUNGI
Fig. 1. Allophoma spp. A, B. Disease symptoms. A. Symptoms caused by Allophoma hayatii (ex-type CBS 142859) on Lantana camara. B. Symptoms caused by Allophoma
pterospermicola (ex-type CGMCC 3.19245) on Pterospermum xylocarpum. C–O. Asexual morph. C. Conidiomata of Allophoma pterospermicola (LC12181) sporulating on
Maesa montana. D. Conidiomata of Allophoma oligotrophica (ex-type CGMCC 3.18114) sporulating on OA. E. Conidiomata of Allophoma minor (ex-type CBS 325.82). F.
Section of the conidiomatal wall of Allophoma minor (ex-type CBS 325.82). G, H. Conidiogenous cells. G. Allophoma piperis (ex-epitype CBS 268.93). H. Allophoma oli-
gotrophica (ex-type CGMCC 3.18114). I–M. Conidia. I. Allophoma minor (ex-type CBS 325.82). J. Allophoma piperis (ex-epitype CBS 268.93). K. Allophoma oligotrophica (ex-
type CGMCC 3.18114). L. Allophoma cylindrispora (ex-type CBS 142453). M. Allophoma nicaraguensis (ex-type CBS 506.91). N. Swollen cells of Allophoma hayatii (ex-type
CBS 142859). O. Chlamydospores of Allophoma hayatii (ex-type CBS 142859). Scale bars: D, E = 100 μm; O = 50 μm; N = 20 μm; F, I–M = 10 μm; G, H = 5 μm. Pictures A,
N–O taken from Babaahmadi et al. (2018); D, H, K, from Chen et al. (2017); E, F, I from Aveskamp et al. (2010); G, J, M from Chen et al. (2015); L from Valenzuela-Lopez et al.
(2018).
and size, i.e. ovoid, oblong, ellipsoidal to cylindrical, or slightly Culture characteristics: Colonies on OA white when young, grey
allantoid, mostly guttulate. Chlamydospores uni- or multicellular to olivaceous or dull green, brown, floccose to woolly, sometimes
(pseudosclerotioid and dictyosporous), solitary or in chains, with rosy-buff tinges near the colony margins or yellow pigment
intercalary or terminal, smooth-walled, brown, where multicellular in the sterile sectors, margins regular.
variable in shape and size. Swollen cells (pseudo-chlamydo-
Optimal media and cultivation conditions: OA or sterile pine
spores) pale brown, terminal or intercalary, solitary or in clusters,
needles placed on OA under near-ultraviolet light (12 h light, 12 h
variable in size and shape, commonly in aerial mycelia. Sexual
dark) to promote sporulation at 25 °C.
morph unknown (adapted from Chen et al. 2015, Babaahmadi
et al. 2018). Distribution: Worldwide.
www.studiesinmycology.org 3
MARIN-FELIX ET AL.
Table 1. DNA barcodes of accepted Allophoma spp.

Species Isolates1 GenBank accession numbers2 References
ITS LSU rpb2 tub2
Allophoma cylindrispora CBS 142453T LT592920 LN907376 LT593058 LT592989 Valenzuela-Lopez et al. (2018)
Al. hayatii CBS 142859T KY684812 KY684814 MF095108 KY684816 Babaahmadi et al. (2018)
CBS 142860 KY684813 KY684815 MF095109 KY684817 Babaahmadi et al. (2018)
Al. labilis CBS 124.93 GU237765 GU238091 KT389552 GU237619 Aveskamp et al. (2010), Chen et al. (2015)
Al. minor CBS 325.82T GU237831 GU238107 KT389553 GU237632 Aveskamp et al. (2010), Chen et al. (2015)
Al. nicaraguensis CBS 506.91T GU237876 GU238058 KT389551 GU237596 Aveskamp et al. (2010), Chen et al. (2015)
T
Al. oligotrophica CGMCC 3.18114 KY742040 KY742194 KY742128 KY742282 Chen et al. (2017)
CGMCC 3.18115 KY742041 KY742195 KY742129 KY742283 Chen et al. (2017)
ET
Al. piperis CBS 268.93 GU237816 GU238129 KT389554 GU237644 Aveskamp et al. (2010), Chen et al. (2015)
CBS 108.93 GU237921 GU238130 KT389555 GU237645 Aveskamp et al. (2010), Chen et al. (2015)
T
Al. pterospermicola CGMCC 3.19245 MK088573 MK088580 MK088587 MK088594 Present study
LC12181 MK088569 MK088576 MK088583 MK088590 Present study
Al. tropica CBS 436.75IsoT GU237864 GU238149 KT389556 GU237663 Aveskamp et al. (2010), Chen et al. (2015)
Al. zantedeschiae CBS 131.93 FJ427084 GU238159 KT389557 FJ427188 Aveskamp et al. (2010), Chen et al. (2015)
CBS 229.32 KT389473 KT389690 KT389558 KT389767 Aveskamp et al. (2010), Chen et al. (2015)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbiological Culture Collection Center, Beijing, China; LC: Dr Lei
Cai's personal culture collection, housed at CAS, China. T, ET and IsoT indicate ex-type,ex-epitype and ex-isotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; rpb2: partial DNA-directed RNA polymerase II second largest
subunit gene; tub2: partial β-tubulin gene.
Hosts: Wide host range, occurring as pathogens or saprobes, on (Boerema et al. 2004, Aveskamp et al. 2010, Chen et al. 2015,
Araceae, Fabaceae, Gesneriaceae, Myrtaceae, Papaveraceae, 2017, Babaahmadi et al. 2018, Valenzuela-Lopez et al. 2018).
Piperaceae, Primulaceae, Rosaceae, Rubiaceae, Solanaceae,
References: Boerema et al. 2004 (morphology and pathoge-
Sterculiaceae, Verbenaceae and other hosts, including humans.
nicity), Aveskamp et al. 2010, Chen et al. 2015, 2017,
Disease symptoms: Dieback, tissue necrosis, leaf spots, stem Babaahmadi et al. 2018, Valenzuela-Lopez et al. 2018
rot, leaf blotch, but also saprobic or isolated from other sub- (morphology, phylogeny and pathogenicity).
strates and environments, e.g. air from karst caves and human
Allophoma pterospermicola Q. Chen & L. Cai, sp. nov.
infections.
MycoBank MB828313. Fig. 3.
Notes: The genus Allophoma was introduced by Chen et al.
(2015) to accommodate five previously described Phoma spe- Etymology: Name reflects Pterospermum, the host genus from
cies, namely Al. labilis (syn. Pho. labilis), Al. minor (syn. Pho. which it was collected.
minor), Al. piperis (syn. Pho. piperis), Al. tropica (syn. Pho.
Conidiomata pycnidial, solitary or aggregated, globose to sub-
tropica) and Al. zantedeschiae (syn. Pho. zantedeschiae), and a
globose, brown, glabrous or with a few hyphal outgrowths, su-
new species Al. nicaraguensis. Another four species have been
perficial, 60–330 × 67–280 μm, with 1–5 ostioles, sometimes
described in the subsequent years, i.e. Al. cylindrispora
elongated as a long neck, up to 150 μm long, papillate; con-
(Valenzuela-Lopez et al. 2018), Al. hayatii (Babaahmadi et al.
idiomatal wall pseudoparenchymatous, 3–5-layered, 12–20 μm
2018), Al. oligotrophica (Chen et al. 2017) and Al. pter-
thick, composed of isodiametric cells. Conidiogenous cells
ospermicola sp. nov. in the present study. Differentiating Allo-
phialidic, hyaline, smooth-walled, ampulliform to doliiform,
phoma from related phoma-like genera based on morphology
6–10 × 3–6 μm. Conidia oval to oblong, occasionally bacilliform,
alone is sometimes complicated. Furthermore, Allophoma spe-
smooth- and thin-walled, hyaline, aseptate, 3–5.5 × 1.5–2 μm,
cies are morphologically similar and hard to differentiate from
with 1–2 minute guttules. Conidial matrix cream.
one another. Therefore, molecular data are essential for accurate
identification of species within this genus, with ITS, LSU, tub2 Culture characteristics: Colonies on OA, 33–40 mm diam after
and rpb2 being the loci selected for this purpose (Chen et al. 1 wk, margins regular, floccose to woolly, white, pale brownish
2015, 2017, Valenzuela-Lopez et al. 2018). No sexual morph grey, with a pale salmon concentric ring, pale salmon near the
of this genus has been observed to date. margins, black pycnidia visible; reverse concolourous. Colonies
These fungi are generally found in soil, air and regarded as on MEA, 20–25 mm diam after 1 wk, margins regular, aerial
saprobes or as the causal organisms of various diseases of mycelium sparse, olivaceous; reverse concolourous. Colonies on
different herbaceous and woody plants, such as some orna- PDA, 20–30 mm diam after 1 wk, margins regular, floccose to
mental plants, coffee, etc., and even human eye lesions woolly, olivaceous, white near the margins; reverse dull green,
4
Fig. 2. Phylogenetic tree generated from a maximum parsimony analysis based on the combined LSU (860 bp), ITS (480 bp), tub2 (333 bp) and rpb2 (803 bp) sequences of all
accepted species of Allophoma. The tree was rooted to Phoma herbarum CBS 615.75. Values above the branches represent parsimony bootstrap support values (> 50 %).
Novel sequences and novel taxon are printed in bold. GenBank accession numbers are indicated in Table 1. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype strains,
respectively. TreeBASE: S23493.
white near the margins. Application of NaOH results in a pale been recorded on Syzygium aromaticum (Myrtaceae)
brownish olivaceous discolouration of the agar. (Aveskamp et al. 2010).
Typus: China, Guangxi, Nonggang National Nature Reserve, on Authors: Q. Chen & L. Cai
diseased leaves of Pterospermum xylocarpum (Sterculiaceae), Alternaria Nees, Das System der Pilze und Schw€amme: 72.
Jun. 2017, Z.Y. Ma (holotype HMAS 247983, culture ex-type 1816 (1816–1817). Fig. 4.
CGMCC 3.19245 = LC 12185). For synonyms see Woudenberg et al. (2013).
Additional materials examined: China, Guangxi, Nonggang National Nature Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
Reserve, on diseased leaves of Pterospermum xylocarpum (Sterculiaceae), Jun. sporales, Pleosporaceae.
2017, Z.Y. Ma, LC 12183; ibid., LC 12184; Guangxi, Jingxi, Gulongshan, on
diseased leaves of Maesa montana (Primulaceae), Jun. 2017, Z.Y. Ma, LC Type species: Alternaria alternata (Fries) Keissler, basionym:
12181; ibid., LC 12182. Torula alternata Fr., Syst. Mycol. (Lundae) 3: 500. 1832 (nom.
sanct.); additional synonyms listed in Woudenberg et al. (2015).
Notes: Allophoma pterospermicola represents the first report of a
Neotype designated by Simmons (1967): E.G.S. 11.050. Ex-
species in the family Didymellaceae on the two host genera
epitype strain designated by de Hoog & Horre (2002): CBS
Pterospermum (Sterculiaceae) and Maesa (Primulaceae). This
916.96 = E.G.S. 34.016.
species is closely related to Al. minor, but differs in producing
longer conidiogenous cells [6–10 × 3–6 μm in Al. pter- DNA barcodes (genus): LSU, ITS.
ospermicola vs. 4–5.5(–6.2) × 3–4.5(–4.7) in Al. minor] and
DNA barcodes (species): ITS, ATPase, gapdh, rpb2, tef1.
slightly narrower conidia [3–5.5 × 1.5–2 μm in Al. pter-
Table 2. Figs 5–7.
ospermicola vs. (3–)3.5–4.5(–5) × 1.8–2.5(–3) μm in Al. mi-
nor]. In addition, Al. pterospermicola grows much slower on OA, Ascomata small, solitary to clustered, erumpent to almost su-
MEA and PDA than Al. minor, and the latter species has only perficial at maturity, dark brown, globose to ovoid, apically
MARIN-FELIX ET AL.
Fig. 3. Allophoma pterospermicola (ex-type CGMCC 3.19245). A, B. Colony on OA (front and reverse). C, D. Colony on MEA (front and reverse). E, F. Colony on PDA (front
and reverse). G. Conidiomata sporulating on OA. H, I. Conidiomata. J. Section of conidioma. K. Section of conidiomatal wall. L, M. Conidiogenous cells. N. Conidia. Scale bars:
H, J = 20 μm; I = 40 μm; K, N = 10 μm; L, M = 5 μm.
papillate, ostiolate, smooth or setose at maturity, with a thin Culture characteristics: Colonies effuse, grey, olivaceous brown,
ascomatal wall; centrum formed by a hamathecium with cellular dark blackish brown or black; mycelium immersed or partly su-
pseudoparaphyses and asci in basal layer. Asci bitunicate, fis- perficial, composed of colourless, olivaceous brown or brown
situnicate, uni- or biseriate, (4–6–)8-spored, cylindrical to hyphae.
cylindro-clavate, straight or somewhat curved, with a short fur-
Optimal media and cultivation conditions: For morphological
cate pedicel. Ascospores ellipsoid to fusoid, muriform, slightly
examinations the use of PCA and V-8 is recommended, incu-
constricted at septa, 3–7-transverse septa, 1–2 series of lon-
bated at moderate temperatures (ca. 22–25 ºC) under near-
gitudinal septa through the two original central segments, end
ultraviolet light (8 h light, 16 h dark), without humidity control,
cells without septa, or with one longitudinal or oblique septum, or
for 5–7 d or more if necessary (Simmons 2007). We also
with a Y-shaped pair of septa, yellow-brown, smooth-walled,
recommend microscopic examination of OA cultures due to the
without guttules. Conidiophores macronematous or semi-
alterations observed on the conidial wall when grown on PCA.
macronematous, mononematous, simple or branched, pale
brown or brown. Conidiogenous cells integrated, terminal Distribution: Worldwide.
becoming intercalary, mono- or polytretic and sympodial,
Hosts: Mainly pathogens of a wide range of plant families, such
cicatrised. Conidia solitary or in simple or branched chains, dry,
as Apiaceae, Asteraceae, Brassicaceae, Cyperaceae, Poaceae,
ovoid, obovoid, cylindrical, narrowly ellipsoid or obclavate,
Rosaceae, Rutaceae, Solanaceae, among others (Thomma
beaked or non-beaked, pale or medium olivaceous brown to
2003, Lawrence et al. 2016). Some are implicated as human
brown, smooth-walled or verrucose, with transverse and with or
pathogens (de Hoog et al. 2011).
without oblique or longitudinal septa; septa can be thick, dark, an
internal cell-like structure can be formed. Species with meriste- Disease symptoms: Most species are foliar pathogens, causing
matic growth are known (adapted from Ellis 1976, Woudenberg necrotic lesions as brown/black spots or “target spot” with the
et al. 2013, 2014, Grum-Grzhimaylo et al. 2016). fungus residing in the central area, but also inducing leaf blight;
6
Fig. 4. Alternaria spp. A–D. Disease symptoms. A. Alternaria dauci on Daucus carota. B. Alternaria linariae on Solanum lycopersicum. C. Alternaria neoipomoeae on
Ipomoeae batatas (Photo A.H. Thompson, ARC, South Africa). D. Alternaria solani on Solanum tuberosum (Photo J.E. van der Waals, University of Pretoria, South Africa). E–V.
Asexual morph. E–O. Conidiophores. E. Alternaria caricis. F. Alternaria chartarum. G. Alternaria cinerariae. H. Alternaria conjuncta. I. Alternaria elegans. J. Alternaria
embellisia. K. Alternaria indefessa. L. Alternaria japonica. M. Alternaria penicillata. N. Alternaria proteae. O. Alternaria tenuissima. P–T. Conidia. P. Alternaria blumeae. Q.
Alternaria calendulae. R. Alternaria perpunctulata. S. Alternaria carotiincultae. T. Alternaria triglochinicola. U, V. Conidia producing secondary conidia. U. Alternaria mimicola. V.
Alternaria molesta. Scale bars: 10 μm. Pictures A–D, P, Q taken from Woudenberg et al. (2014); E–O, R–V from Woudenberg et al. (2013).
8
MARIN-FELIX
Table 2. DNA barcodes of accepted Alternaria spp.
Species Section Isolates1 GenBank accession numbers2 References
ET AL.
ITS gapdh rpb2 tef1 ATPase
Alternaria abundans Chalastospora CBS 534.83T JN383485 KC584154 KC584448 KC584707 JQ671802 Woudenberg et al. (2013), Deng et al. (2018)
A. acalyphicola Porri CBS 541.94T KJ718097 KJ717952 KJ718271 KJ718446 – Woudenberg et al. (2014)
A. aconidiophora Infectoriae FMR 17111T LR133931 LR133965 LR133967 LR133968 LR133969 Present study
A. agerati Porri CBS 117221 KJ718098 KJ717953 KJ718272 KJ718447 – Woudenberg et al. (2014)
A. agripestis Porri CBS 577.94T KJ718099 JQ646356 KJ718273 KJ718448 – Woudenberg et al. (2014)
A. allii Porri CBS 107.28T KJ718100 KJ717954 KJ718274 KJ718449 – Woudenberg et al. (2014)
A. alstroemeriae Alternaria CBS 118809 T
KP124297 KP124154 KP124765 KP125072 – Woudenberg et al. (2015)
A. altcampina Pseudoalternaria FMR 16476T LR133895 LR133900 – – LR133906 Present study
A. alternantherae Althernantherae CBS 124392 KC584179 KC584096 KC584374 KC584633 – Woudenberg et al. (2013)
A. alternariacida Porri CBS 105.51T KJ718105 KJ717959 KJ718279 KJ718454 – Woudenberg et al. (2014)
A. alternariae Ulocladium CBS 126989T AF229485 AY278815 KC584470 KC584730 – Woudenberg et al. (2013)
A. alternarina Infectoriae CBS 119396T JQ693648 JQ646289 JQ905199 LR134367 JQ671817 Poursafar et al. (2018), Geng et al.
(unpubl. data), present study
A. alternata Alternaria CBS 916.96T AF347031 AY278808 KC584375 KC584634 – Woudenberg et al. (2013)
A. anagallidis Porri CBS 117128 KJ718106 JQ646338 KJ718280 EU130544 – Woudenberg et al. (2014)
A. anigozanthi Eureka CBS 121920T KC584180 KC584097 KC584376 KC584635 – Woudenberg et al. (2013)
A. anodae Porri PPRI 12376 KJ718110 KJ717963 KJ718284 KJ718458 – Woudenberg et al. (2014)
A. aragakii Porri CBS 594.93T KJ718111 KJ717964 KJ718285 KJ718459 – Woudenberg et al. (2014)
A. arborescens Alternaria CBS 102605T AF347033 AY278810 KC584377 KC584636 – Woudenberg et al. (2013)
A. arbusti Infectoriae CBS 596.93 T
JQ693644 JQ646365 LR134184 – JQ671940 Poursafar et al. (2018), present study
A. argyranthemi CBS 116530T KC584181 KC584098 KC584378 KC584637 – Woudenberg et al. (2013)
A. argyroxiphii Porri CBS 117222T KJ718112 JQ646350 KJ718286 KJ718460 – Woudenberg et al. (2014)
A. armoraciae Chalastospora CBS 118702T KC584182 KC584099 KC584379 KC584638 LR134098 Woudenberg et al. (2013), present study
A. arrhenatheri Pseudoalternaria LEP 140372 T
JQ693677 JQ693635 – – JQ693603 Poursafar et al. (2018)
A. aspera Pseudoulocladium CBS 115269T KC584242 KC584166 KC584474 KC584734 – Woudenberg et al. (2013)
A. atra Ulocladioides CBS 195.67T AF229486 KC584167 KC584475 KC584735 – Woudenberg et al. (2013)
A. avenicola Panax CBS 121459T KC584183 KC584100 KC584380 KC584639 – Woudenberg et al. (2013)
A. axiaeriisporifera Gypsophilae CBS 118715 T
KC584184 KC584101 KC584381 KC584640 – Woudenberg et al. (2013)
A. azadirachtae Porri CBS 116444T KJ718115 KJ717967 KJ718289 KJ718463 – Woudenberg et al. (2014)
A. bataticola Porri CBS 531.63 T
KJ718117 JQ646349 KJ718291 KJ718465 – Woudenberg et al. (2014)
A. betae-kenyensis Alternaria CBS 118810T KP124419 KP124270 KP124888 KP125197 – Woudenberg et al. (2015)
Table 2. (Continued).

www.studiesinmycology.org
A. blumeae Porri CBS 117364T KJ718126 AY562405 KJ718300 KJ718474 – Woudenberg et al. (2014)
A. bornmuelleri Undifilum DAOM 231361T FJ357317 FJ357305 KC584491 KC584751 – Woudenberg et al. (2013)
A. botryospora Embellisioides CBS 478.90 T
AY278844 AY278831 KC584461 KC584720 – Woudenberg et al. (2013)
A. botrytis Ulocladium CBS 197.67T KC584243 KC584168 KC584476 KC584736 – Woudenberg et al. (2013)
A. brassicae CBS 116528 KC584185 KC584102 KC584382 KC584641 – Woudenberg et al. (2013)
A. brassicaepekinensis Ulocladioides CBS 121493T KC584244 KC584170 KC584478 KC584738 – Woudenberg et al. (2013)
A. brassicicola Brassicicola CBS 118699 JX499031 KC584103 KC584383 KC584642 – Woudenberg et al. (2013)
A. brassicifolii Pseudoalternaria CNU 111118T JQ317188 KM821537 – – KY412558 Deng et al. (2018)
A. breviramosa Chalastospora CBS 121331T FJ839608 KC584148 KC584442 KC584700 LR134099 Woudenberg et al. (2013), present study
T
A. broccoli-italicae Infectoriae CBS 118485 KM821536 KM821538 LR134194 LR134262 KY412557 Deng et al. (2018), present study
A. burnsii Alternaria CBS 107.38T KP124420 JQ646305 KP124889 KP125198 – Woudenberg et al. (2015)
A. caespitosa Infectoriae CBS 177.80T KC584250 KC584178 KC584492 KC584752 LR134114 Woudenberg et al. (2013), present study
A. calendulae Porri CBS 224.76T KJ718127 KJ717977 KJ718301 KJ718475 – Woudenberg et al. (2014)
A. californica Infectoriae CBS 119409T JQ693645 JQ646285 LR134181 LR134245 JQ671813 Poursafar et al. (2018), present study
A. calycipyricola Panax CBS 121545T KC584186 KC584104 KC584384 KC584643 – Woudenberg et al. (2013)
A. cantlous Ulocladioides CBS 123007T KC584245 KC584171 KC584479 KC584739 – Woudenberg et al. (2013)
A. capsici-annui Ulocladium CBS 504.74 KC584187 KC584105 KC584385 KC584644 – Woudenberg et al. (2013)
A. caricis Nimbya CBS 480.90T AY278839 AY278826 KC584467 KC584726 JQ671780 Woudenberg et al. (2013), Deng et al. (2018)
A. carotiincultae Radicina CBS 109381T KC584188 KC584106 KC584386 KC584645 – Woudenberg et al. (2013)
A. carthami Porri CBS 635.80 KJ718131 KJ717981 KJ718305 KJ718479 – Woudenberg et al. (2014)
A. carthamicola Porri CBS 117092T KJ718134 KJ717984 KJ718308 KJ718482 – Woudenberg et al. (2014)
A. cassiae Porri CBS 478.81 KJ718135 KJ717985 KJ718309 KJ718483 – Woudenberg et al. (2014)
A. catananches Porri CBS 137456T KJ718139 KJ717989 KJ718313 KJ718487 – Woudenberg et al. (2014)
GENERA
A. centaureae Porri CBS 116446 T
KJ718140 KJ717990 KJ718314 KJ718488 – Woudenberg et al. (2014)
A. cerasidanica Infectoriae CBS 121923T LR135744 LR135747 LR135746 LR135745 LR135748 Present study
OF
A. cesenica Infectoriae MFLUCC 13-0450 T
KP711383 – – KP711386 – Liu et al. (2015)
PHYTOPATHOGENIC FUNGI
A. cetera Chalastospora CBS 121340T JN383482 AY562398 KC584441 KC584699 LR134101 Woudenberg et al. (2013), Deng et al. (2018)
A. chartarum Pseudoulocladium CBS 200.67 T
AF229488 KC584172 KC584481 KC584741 – Woudenberg et al. (2013)
A. cheiranthi Cheiranthus CBS 109384 AF229457 KC584107 KC584387 KC584646 – Woudenberg et al. (2013)
A. chlamydospora Phragmosporae CBS 491.72T KC584189 KC584108 KC584388 KC584647 – Woudenberg et al. (2013)
(continued on next page)
9
10
MARIN-FELIX
ET AL.
A. chlamydosporigena Embellisia CBS 341.71 KC584231 KC584156 KC584451 KC584710 – Woudenberg et al. (2013)
A. chlamydosporifera Radicina FMR 17360T LR133924 LR133927 LR133926 LR133929 – Present study
A. cichorii Porri CBS 102.33T KJ718141 KJ717991 KJ718315 KJ718489 – Woudenberg et al. (2014)
A. cinerariae Sonchi CBS 116495 KC584190 KC584109 KC584389 KC584648 – Woudenberg et al. (2013)
A. cirsinoxia Porri CBS 113261T KJ718143 KJ717993 KJ718317 KJ718491 – Woudenberg et al. (2014)
A. citrullicola Porri CBS 103.32T KJ718144 KJ717994 KJ718318 KJ718492 – Woudenberg et al. (2014)
A. concatenata Pseudoulocladium CBS 120006 T
KC584246 AY762950 KC584480 KC584740 – Woudenberg et al. (2013)
A. conidiophora Porri CBS 137457T KJ718145 KJ717995 - KJ718493 – Woudenberg et al. (2014)
T
A. conjuncta Infectoriae CBS 196.86 FJ266475 AY562401 KC584390 KC584649 JQ671824 Woudenberg et al. (2013), Poursafar et al. (2018)
A. conoidea Brassicicola CBS 132.89 AF348226 FJ348227 KC584452 KC584711 – Woudenberg et al. (2013)
A. consortialis Ulocladioides CBS 104.31T KC584247 KC584173 KC584482 KC584742 – Woudenberg et al. (2013)
A. crassa Porri CBS 110.38T KJ718147 KJ717997 KJ718320 KJ718495 – Woudenberg et al. (2014)
A. cyamopsidis Porri CBS 364.67 KJ718156 KJ718003 KJ718329 KJ718504 – Woudenberg et al. (2014)
A. cumini Eureka CBS 121329T KC584191 KC584110 KC584391 KC584650 – Woudenberg et al. (2013)
A. cucumerina Porri CBS 116114T KJ718153 KJ718000 KJ718326 KJ718501 – Woudenberg et al. (2014)
A. cucurbitae Ulocladioides CBS 483.81 FJ266483 AY562418 KC584483 KC584743 – Woudenberg et al. (2013)
A. curvata Infectoriae FMR 16901T LR133898 LR133899 LR133901 LR133902 LR133905 Present study
A. dactylidicola Infectoriae MFLUCC 15-0466 T
KY703616 – KY750720 – – Thambugala et al. (2017)
A. dauci Porri CBS 111.38T KJ718158 KJ718005 KJ718331 KJ718506 – Woudenberg et al. (2014)
A. daucicaulis Infectoriae CBS 119398T JQ693653 JQ646294 LR134177 LR134241 JQ671822 Poursafar et al. (2018), present study
A. dennisii CBS 476.90T JN383488 JN383469 KC584454 KC584713 – Woudenberg et al. (2013)
A. deserticola Porri CBS 110799T KJ718249 KJ718077 KJ718424 KJ718595 – Woudenberg et al. (2014)
A. dianthicola Dianthicola CBS 116491 KC584194 KC584113 KC584394 KC584653 – Woudenberg et al. (2013)
A. dichondrae Porri CBS 200.74T KJ718167 KJ718012 KJ718340 KJ718515 – Woudenberg et al. (2014)
A. didymospora Phragmosporae CBS 766.79 FJ357312 FJ357300 KC584455 KC584714 JQ671796 Woudenberg et al. (2013), Deng et al. (2018)
A. doliconidium Alternaria KUMCC 17-0263T MG828864 – – – – Wanasinghe et al. (2018)
A. echinaceae Porri CBS 116117T KJ718170 KJ718015 KJ718343 KJ718518 – Woudenberg et al. (2014)
A. eichhorniae Alternaria CBS 489.92T KC146356 KP124276 KP124895 KP125204 – Woudenberg et al. (2015)
A. elegans Dianthicola CBS 109159T KC584195 KC584114 KC584395 KC584654 – Woudenberg et al. (2013)
A. ellipsoidea Gypsophilae CBS 119674 T
A. embellisia Embellisia CBS 339.71 KC584230 KC584155 KC584449 KC584708 – Woudenberg et al. (2013)

A. eryngii Panax CBS 121339 JQ693661 AY562416 KC584397 KC584656 – Woudenberg et al. (2013)
A. ethzedia Infectoriae CBS 197.86T AY278833 AY278795 KC584398 KC584657 JQ671805 Woudenberg et al. (2013), Poursafar et al. (2018)
A. euphorbiicola Euphorbiicola CBS 119410 KJ718173 KJ718018 KJ718346 KJ718521 – Woudenberg et al. (2014)
A. eureka Eureka CBS 193.86T JN383490 JN383471 KC584456 KC584715 – Woudenberg et al. (2013)
A. fimeti Infectoriae FMR 17110T LR133920 LR133921 LR133923 LR133922 LR133925 Present study
A. forlicesenensis Infectoriae MFLUCC 13-0456 T
KY769657 – – – – Thambugala et al. (2017)
A. frumenti Infectoriae CBS 119401T JQ693654 JQ646295 LR134172 LR134370 JQ671823 Poursafar et al. (2018), present study
A. gaisen Alternaria CBS 632.93 KC584197 KC584116 KC584399 KC584658 – Woudenberg et al. (2013)
A. geniostomatis Eureka CBS 118701T KC584198 KC584117 KC584400 KC584659 – Woudenberg et al. (2013)
A. gossypina Alternaria CBS 104.32 T
KP124430 JQ646312 KP124900 KP125209 – Woudenberg et al. (2015)
A. graminicola Infectoriae CBS 119400T JQ693650 JQ646291 LR134180 LR134249 JQ671819 Poursafar et al. (2018), present study
A. grandis Porri CBS 109158 T
KJ718239 JQ646341 KJ718414 EU130547 – Woudenberg et al. (2014)
A. gypsophilae Gypsophilae CBS 107.41T KC584199 KC584118 KC584401 KC584660 – Woudenberg et al. (2013)
A. hampshirensis Infectoriae MFLUCC 17-0783 T
MG828866 – MG829247 – – Wanasinghe et al. (2018)
A. helianthiinficiens CBS 208.86T JX101649 KC584120 KC584403 EU130548 – Woudenberg et al. (2013)
A. heterospora Ulocladioides CBS 123376T KC584248 KC584176 KC584488 KC584748 – Woudenberg et al. (2013)
A. hordeiaustralica Infectoriae CBS 119402T JQ693641 JQ646283 LR134179 LR134243 JQ671811 Poursafar et al. (2018), present study
T
A. hordeicola Infectoriae CBS 121458 JQ693642 JQ646284 LR134175 LR134371 JQ671812 Poursafar et al. (2018), present study
A. humuli Infectoriae CBS 119404T JQ693652 JQ646293 LR134174 LR134199 JQ671821 Poursafar et al. (2018), present study
A. hyacinthi Embellisioides CBS 416.71 T
A. incomplexa Infectoriae CBS 121330T JQ693658 JQ646287 LR134185 LR134250 JQ671815 Poursafar et al. (2018), present study
A. indefessa Cheiranthus CBS 536.83 T
A. infectoria Infectoriae CBS 210.86T AF347034 AY278793 KC584404 KC584662 JQ671804 Woudenberg et al. (2013), Poursafar et al. (2018)
GENERA
A. inflata Pseudoalternaria FMR 16477 T
LR133930 LR133938 – – LR133966 Present study
A. intercepta Infectoriae CBS 119406T JQ693656 JQ646297 LR134170 – JQ671826 Poursafar et al. (2018), present study
OF
A. ipomoeae Porri CBS 219.79 T
A. iridiaustralis Alternaria CBS 118486T KP124435 KP124284 KP124905 KP125214 – Woudenberg et al. (2015)
A. japonica Japonicae CBS 118390 KC584201 KC584121 KC584405 KC584663 – Woudenberg et al. (2013)
A. jacinthicola Alternaria CBS 133751T KP124438 KP124287 KP124908 KP125217 – Woudenberg et al. (2015)
A. jesenskae Porri CBS 133855 T
11
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MARIN-FELIX
ET AL.
A. juxtiseptata Gypsophilae CBS 119673T KC584202 KC584122 KC584406 KC584664 – Woudenberg et al. (2013)
A. kordkuyana Pseudoalternaria IRAN 16888FT MF033843 MF033826 – – MF033860 Poursafar et al. (2018)
FMR 17061 LR133970 LR133998 – – LR134001 Present study
A. kulundii Soda CBS 137525T KJ443262 KJ649618 KJ443176 KJ443219 – Grum-Grzhimaylo et al. (2016)
A. lawrencei Infectoriae FMR 17004T LR133907 LR133908 LR133911 LR133912 LR133914 Present study
A. leptinellae Eureka CBS 477.90 T
A. leucanthemi Teretispora CBS 421.65T KC584240 KC584164 KC584472 KC584732 – Woudenberg et al. (2013)
T
A. limaciformis Phragmosporae CBS 481.81 KC584203 KC584123 KC584407 KC584665 JQ671798 Woudenberg et al. (2013), Deng et al. (2018)
A. limicola Euphorbiicola CBS 483.90T KJ718178 JQ646329 KJ718351 KJ718526 – Woudenberg et al. (2014)
A. linariae Porri CBS 105.41T KJ718180 KJ718024 KJ718353 KJ718528 – Woudenberg et al. (2014)
A. longipes Alternaria CBS 540.94 AY278835 AY278811 KC584409 KC584667 – Woudenberg et al. (2013)
A. lolii Embellisioides CBS 115266T JN383492 JN383473 KC584460 KC584719 – Woudenberg et al. (2013)
A. macrospora Porri CBS 117228T KC584204 KC584124 KC584410 KC584668 – Woudenberg et al. (2013)
A. malorum Chalastospora CBS 135.31 JQ693638 JQ646278 – – JQ671800 Poursafar et al. (2018)
A. merytae Infectoriae CBS 119403T JQ693651 JQ646292 LR134119 LR134198 JQ671820 Poursafar et al. (2018), present study
T
A. metachromatica Infectoriae CBS 553.94 JQ693660 AY562404 JQ905189 FJ214931 JQ671809 Andersen et al. (2009), Poursafar et al. (2018),
Geng et al. (unpubl. data)
A. mimicula Brassicicola CBS 118696T FJ266477 AY562415 KC584411 KC584669 – Woudenberg et al. (2013)
A. molesta Phragmosporae CBS 548.81T KC584205 KC584125 KC584412 KC584670 – Woudenberg et al. (2013)
A. montanica Porri CBS 121343 T
A. montsantina Infectoriae FMR 17060T LR133913 LR133915 LR133918 LR133919 LR133916 Present study
A. mouchaccae Phragmosporae CBS 119671 T
KC584206 AY562399 KC584413 KC584671 – Woudenberg et al. (2013)
A. multiformis Ulocladioides CBS 102060T FJ266486 KC584174 KC584484 KC584744 – Woudenberg et al. (2013)
A. multirostrata Porri CBS 712.68 T
KJ718195 JQ646362 KJ718368 EU130546 – Woudenberg et al. (2014)
A. murispora Infectoriae MFLU 14-0758T NR_137964 – – – – Ariyawansa and Hyde (unpubl. data)
A. neoipomoeae Porri PPRI 11845 T
A. nepalensis Japonicae CBS 118700T KC584207 KC584126 KC584414 KC584672 – Woudenberg et al. (2013)
A. nitrimali Porri CBS 109163 T
A. nobilis Gypsophilae CBS 116490 KC584208 KC584127 KC584415 KC584673 – Woudenberg et al. (2013)

A. novae-guineensis Porri CBS 116120T KJ718202 KJ718039 KJ718375 KJ718548 – Woudenberg et al. (2014)
A. novae-zelandiae Infectoriae CBS 119405T JQ693655 JQ646296 LR134120 LR134197 JQ671825 Poursafar et al. (2018), present study
T
A. obclavata Chalastospora CBS 124120 KC584225 KC584149 KC584443 KC584701 LR134100 Woudenberg et al. (2013), present study
A. obovoidea Ulocladioides CBS 101229 FJ266487 FJ266498 KC584485 KC584745 – Woudenberg et al. (2013)
A. obtecta Porri CBS 117367 KJ718204 KJ718041 KJ718377 KJ718550 – Woudenberg et al. (2014)
A. oregonensis Infectoriae CBS 542.94T FJ266478 FJ266491 KC584416 KC584674 JQ671827 Woudenberg et al. (2013), Poursafar et al. (2018)
A. oudemansii Ulocladium CBS 114.07 T
FJ266488 KC584175 KC584486 KC584746 – Woudenberg et al. (2013)
A. panax Panax CBS 482.81 KC584209 KC584128 KC584417 KC584675 – Woudenberg et al. (2013)
A. papavericola Crivellia CBS 116606T FJ357310 FJ357298 KC584446 KC584705 – Woudenberg et al. (2013)
A. paralinicola Porri CBS 116652T KJ718206 KJ718043 KJ718379 KJ718552 – Woudenberg et al. (2014)
A. passiflorae Porri CBS 113.38 KJ718207 JQ646353 KJ718380 KJ718553 – Woudenberg et al. (2014)
A. parvicaespitosa Pseudoalternaria LEP 014858T MF033859 MF033842 – – KJ908217 Poursafar et al. (2018)
A. penicillata Crivellia CBS 116608 T
FJ357311 FJ357299 KC584440 KC584698 – Woudenberg et al. (2013)
A. perpunctulata Althernantherae CBS 115267T KC584210 KC584129 KC584418 KC584676 – Woudenberg et al. (2013)
A. petroselini Radicina CBS 112.41T KC584211 KC584130 KC584419 KC584677 – Woudenberg et al. (2013)
A. petuchovskii Soda CBS 137517 T
KJ443254 KJ649616 KJ443170 KJ443211 – Grum-Grzhimaylo et al. (2016)
A. peucedani CNU 111485T KF728231 KF889361 – – – Deng et al. (2014)
T
A. photistica Panax CBS 212.86 KC584212 KC584131 KC584420 KC584678 JQ671807 Woudenberg et al. (2013), Deng et al. (2018)
A. phragmospora Phragmosporae CBS 274.70T JN383493 JN383474 KC584462 KC584721 – Woudenberg et al. (2013)
A. pipionipisi Porri CBS 116115T KJ718214 KJ718049 KJ718387 KJ718560 – Woudenberg et al. (2014)
A. planifunda Embellisioides CBS 537.83T FJ357315 FJ357303 KC584463 KC584722 – Woudenberg et al. (2013)
A. poaceicola Infectoriae MFLUCC 13-0346 T
KY026587 – KY460971 – – Thambugala et al. (2017)
A. pobletensis Chalastospora FMR 16448T LR133896 LR133897 – – LR133903 Present study
GENERA
A. porri Porri CBS 116699T KJ718218 KJ718053 KJ718391 KJ718564 – Woudenberg et al. (2014)
A. proteae Embellisioides CBS 475.90T AY278842 KC584161 KC584464 KC584723 – Woudenberg et al. (2013)
–
OF
A. protenta Porri CBS 116696 KJ718221 JQ646335 KJ718394 KJ718567 Woudenberg et al. (2014)
A. pseudorostrata Porri CBS 119411T JN383483 AY562406 KC584422 KC584680 – Woudenberg et al. (2013)
T
A. pseudoventricosa Infectoriae FMR 16900 LR133928 LR133935 LR133934 LR133936 LR133937 Present study
A. radicina Radicina CBS 245.67T KC584213 KC584133 KC584423 KC584681 – Woudenberg et al. (2013)
A. ranunculi Porri CBS 116330T KJ718225 KJ718058 KJ718398 KJ718571 – Woudenberg et al. (2014)
13
14
MARIN-FELIX
ET AL.
A. ricini Porri CBS 215.31T KJ718226 KJ718059 KJ718399 KJ718572 – Woudenberg et al. (2014)
A. rosae Pseudoalternaria CBS 121341T JQ646279 JQ646279 – – JQ671803 Poursafar et al. (2018)
A. roseogrisea Infectoriae CBS 121921T LR134102 LR134103 LR134192 LR134260 LR134104 Present study
A. rostellata Porri CBS 117366 T
A. saponariae Gypsophilae CBS 116492 KC584215 KC584135 KC584425 KC584683 – Woudenberg et al. (2013)
A. scirpicola Nimbya CBS 481.90 KC584237 KC584163 KC584469 KC584728 JQ671781 Woudenberg et al. (2013), Deng et al. (2018)
A. scirpinfestans Nimbya EGS 49-185T JN383499 JN383480 – JQ672404 JQ671783 Lawrence et al. (2012), Lawrence et al. (unpubl. data)
A. scirpivora Nimbya EGS 50-021 T
JN383500 JN383481 – JQ672405 JQ671782 Lawrence et al. (2012), Lawrence et al. (unpubl. data)
A. scorzonerae Porri CBS 103.46 KJ718190 JQ646363 KJ718363 KJ718537 – Woudenberg et al. (2014)
A. selini Radicina CBS 109382T AF229455 AY278800 KC584426 KC584684 – Woudenberg et al. (2013)
A. sennae Porri CBS 477.81T KJ718230 JQ646344 KJ718403 EU130543 – Woudenberg et al. (2014)
A. septospora Pseudoulocladium CBS 109.38 FJ266489 FJ266500 KC584487 KC584747 – Woudenberg et al. (2013)
A. septorioides Brassicicola CBS 106.41T KC584216 KC584136 KC584427 KC584685 – Woudenberg et al. (2013)
A. sesami Porri CBS 115264 JF780939 KJ718061 KJ718405 KJ718577 – Woudenberg et al. (2014)
A. shukurtuzii Soda CBS 137520T KJ443257 KJ649620 KJ443172 KJ443214 – Grum-Grzhimaylo et al. (2016)
A. sidae Porri CBS 117730T KJ718232 KJ718062 KJ718406 KJ718578 – Woudenberg et al. (2014)
A. simsimi Dianthicola CBS 115265 T
JF780937 KC584137 KC584428 KC584686 – Woudenberg et al. (2013)
A. silybi Porri CBS 134092T KJ718233 KJ718063 KJ718407 KJ718579 – Woudenberg et al. (2014)
T
A. slovaca Infectoriae CBS 567.66 KC584226 KC584150 KC584444 KC584702 LR134368 Woudenberg et al. (2013), present study
A. smyrnii Radicina CBS 109380 AF229456 KC584138 KC584429 KC584687 – Woudenberg et al. (2013)
A. solani Porri CBS 106.21 KJ718236 KJ718066 KJ718410 KJ718582 – Woudenberg et al. (2014)
A. solani-nigri Porri CBS 113403 KJ718243 KJ718071 KJ718418 KJ718589 – Woudenberg et al. (2014)
A. soliaridae CBS 118387T KC584218 KC584140 KC584431 KC584689 – Woudenberg et al. (2013)
A. solidaccana Brassicicola CBS 118698T KC584219 KC584141 KC584432 KC584690 – Woudenberg et al. (2013)
A. sonchi Sonchi CBS 119675 KC584220 KC584142 KC584433 KC584691 – Woudenberg et al. (2013)
A. steviae Porri CBS 117362T KJ718252 KJ718079 KJ718427 KJ718598 – Woudenberg et al. (2014)
A. subcucurbitae Ulocladioides CBS 121491 T
KC584249 EU855803 KC584489 KC584749 – Woudenberg et al. (2013)
A. tagetica Porri CBS 479.81 KC584221 KC584143 KC584434 KC584692 – Woudenberg et al. (2013)

A. tellustris Embellisia CBS 538.83T FJ357316 AY562419 KC584465 KC584724 JQ671794 Woudenberg et al. (2013), Deng et al. (2018)
A. terricola Ulocladioides CBS 202.67T FJ266490 KC584177 KC584490 KC584750 – Woudenberg et al. (2013)
A. tillandsiae Porri CBS 116116T KJ718260 KJ718087 KJ718435 KJ718606 – Woudenberg et al. (2014)
A. thalictrigena CBS 121712T EU040211 KC584144 KC584436 KC584694 – Woudenberg et al. (2013)
A. thunbergiae Porri CBS 116331 T
A. tomato Alternaria CBS 103.30 KP124445 KP124294 KP124915 KP125224 – Woudenberg et al. (2015)
A. triglochinicola Eureka CBS 119676T KC584222 KC584145 KC584437 KC584695 – Woudenberg et al. (2013)
A. triticimaculans Infectoriae CBS 578.94T JQ693657 JQ646280 LR134183 – JQ671806 Poursafar et al. (2018), present study
T
A. triticina Infectoriae CBS 763.84 AY278834 JQ646281 LR134186 FJ214942 JQ671808 Andersen et al. (2009), Poursafar et al. (2018), Present study
A. tropica Porri CBS 631.93T KJ718261 KJ718088 KJ718436 KJ718607 – Woudenberg et al. (2014)
A. tumida Embellisioides CBS 539.83T FJ266481 FJ266493 KC584466 KC584725 – Woudenberg et al. (2013)
A. quercicola Infectoriae CBS 141466T KX228295 KX228362 LR134188 LR134259 LR134115 Crous et al. (2013), present study
A. vaccariae Gypsophilae CBS 116533 KC584223 KC584146 KC584438 KC584696 – Woudenberg et al. (2013)
A. vaccariicola Gypsophilae CBS 118714T KC584224 KC584147 KC584439 KC584697 – Woudenberg et al. (2013)
A. venezuelensis Porri CBS 116121T KJ718263 KJ718090 KJ718438 KJ718609 – Woudenberg et al. (2014)
T
A. ventricosa Infectoriae CBS 121546 JQ693649 JQ646290 LR134134 KY352501 JQ671818 Poursafar et al. (2018), Fotedar et al. (unpubl. data), present study
A. viburni Infectoriae CBS 119407T JQ693647 JQ646288 LR134166 LR134200 JQ671816 Poursafar et al. (2018), present study
A. zinniae Porri CBS 117223 KJ718270 KJ718096 KJ718445 KJ718616 – Woudenberg et al. (2014)
1
CBS: Culture collection of the Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CNU: Culture Collection Center of the Chungnam National University; DAOM: Canadian Collection of Fungal Cultures, Ottawa, Canada; EGS: Personal
collection of Dr. E.G. Simmons; FMR: Facultat de Medicina, Universitat Rovira i Virgili, Reus, Spain; IRAN: Fungal Culture Collections of the Iranian Research Institute of Plant Protection; KUMCC, Culture collection of Kunming Institute of Botany,
Kunming, China; LEP: Mycological Herbarium of All-Russian Institute of Plant Protection, Saint Petersburg, Russia; MFLU and MFLUCC: Herbarium and culture collection of Mae Fah Luang University, Chiang Rai, Thailand, respectively; PPRI: ARC-
Plant Protection Research Institute, Roodeplaat, South Africa. T indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial translation elongation factor 1-
alpha gene; ATPase: partial plasma membrane ATPase gene.
GENERA
OF
15
MARIN-FELIX ET AL.
CBS 119398T Alternaria daucicaulis

CBS 119402T Alternaria hordeiaustralica
CBS 121458T Alternaria hordeicola
CBS 119409T Alternaria californica
CBS 119407T Alternaria viburni
T
79 CBS 196.86 Alternaria conjuncta
88 CBS 121923T Alternaria cerasidanica
CBS 119404T Alternaria humuli
CBS 118485T Alternaria broccoli-italicae
90 CBS 210.86T Alternaria infectoria
CBS 119401T Alternaria frumenti
CBS 119396T Alternaria alternarina section
FMR 17004T Alternaria lawrencei Infectoriae
CBS 119403T Alternaria merytae
CBS 119405T Alternaria novae-zelandiae
119400T Alternaria graminicola
85 CBS 578.94T Alternaria triticimaculans
CBS 197.86T Alternaria ethzedia
CBS 763.84T Alternaria triticina
100 CBS 553.94T Alternaria metachromatica
CBS 119406T Alternaria intercepta
75
CBS 121330T Alternaria incomplexa
80 CBS 542.94T Alternaria oregonensis
77 596.93T Alternaria arbusti
87 CBS 121921T Alternaria roseogrisea
FMR 17110T Alternaria fimeti
FMR 17111T Alternaria aconidiophora
CBS 141466T Alternaria quercicola
CBS 567.66T Alternaria slovaca
CBS 177.80T Alternaria caespitosa
FMR 16901T Alternaria curvata
FMR 17060T Alternaria montsantina
CBS 121546T Alternaria ventricosa
FMR 16900T Alternaria pseudoventricosa
CBS 534.83T Alternaria abundans
CBS 121331T Alternaria breviramosa section Chalastospora
0.01
Fig. 5. Maximum Likelihood (ML) tree constructed with ITS (529 bp), ATPase (1180 bp), gapdh (489 bp), rpb2 (573 bp) and tef1 (239 bp) sequences of ex-type strains of the
species in section Infectoriae. The phylogenetic tree was rooted to Alternaria abundans CBS 534.83 and Alternaria breviramosa CBS 121331 (section Chalastospora). Bootstrap
support values above 70 % are shown at the nodes. GenBank accession numbers are indicated in Table 2. The novel species described in this study are indicated in bold. T
indicates ex-type strain. TreeBASE: S23786.
seed-borne species may attack seedlings, resulting in damping- Notes: Alternaria is characterised mainly by its asexual morph
off, stem lesions or collar rot; sunken and dark lesions are with darkly pigmented multi-celled conidia, which are typically
present in roots, tubers, stems and fruits infections; some rots dictyosporous, some phragmosporous, and arranged single or
and decay are typical symptoms of post-harvest diseases in chains on the conidiophore. Some of these morphological
(Laemmlen 2001, Thomma 2003, Lawrence et al. 2008). Phy- features can also be observed in other closely related genera
totoxins are also produced during the invasion process as viru- such as Paradendryphiella (Woudenberg et al. 2013) or
lence factors which affect a wide spectrum of plant species. Stemphylium (Woudenberg et al. 2017). However, Paraden-
Alternaria toxins diffuse into host tissues resulting in a chlorotic or dryphiella mainly differs by its denticulate conidiogenous cells
yellow halo around lesions, exacerbating the severity of the with prominent conidial scars aggregated at the apex of simple
symptoms (Singh et al. 2015). or branched conidiophores, and Stemphylium by showing
16
IRAN 16888FT Alternaria kordkuyana

97 FMR 17061 Alternaria kordkuyana
99 FMR 17372 Alternaria kordkuyana
86 X1272T Alternaria parvicaespitosa

88 FMR 16476T Alternaria altcampina
100 FMR 16477T Alternaria inflata section Pseudoalternaria
LEP 140372T Alternaria arrhenatheri
CBS 121341T Alternaria rosae
100 100
FMR 17376 Alternaria rosae
97
FMR 15720 Alternaria rosae
83 FMR 17377 Alternaria rosae
CNU 111118T Alternaria brassicifolii
100 CBS 135.31 Alternaria malorum
FMR 17369 Alternaria malorum
99 CBS 121340T Alternaria cetera
94
CBS 124120T Alternaria obclavata
section Chalastospora
76 FMR 16448T Alternaria pobletensis
80 CBS 121331T Alternaria breviramosa

98 CBS 534.83T Alternaria abundans
100 CBS 118702T Alternaria armoraciae
CBS 480.90T Alternaria caricis
100 CBS 481.90 Alternaria scirpicola section Nimbya
0.01
Fig. 6. Maximum Likelihood (ML) tree constructed with ITS (576 bp), ATPase (1198 bp) and gapdh (491 bp) sequences of ex-type strains of species in the sections
Pseudoalternaria and Chalastospora. The phylogenetic tree was rooted to Alternaria caricis CBS 480.90 and A. scirpicola CBS 481.90 (section Nimbya). Bootstrap support
values above 70 % are shown at the nodes. GenBank accession numbers are indicated in Table 2. The novel species described in this study are indicated in bold. T indicates
ex-type strain. TreeBASE: S23787.
T
100 CBS 112.41 Alternaria petroselini
CBS 109382T Alternaria selini
95 FMR 17360T Alternaria chlamydosporifera
section Radicina
100 CBS 109380 Alternaria smyrnii
CBS 109381T Alternaria carotiincultae
100 CBS 245.67T Alternaria radicina
CBS 481.90 Alternaria scirpicola
CBS 480.90T Alternaria caricis

section Nimbya
0.01
Fig. 7. Maximum Likelihood (ML) tree constructed with ITS (523 bp), gapdh (503 bp), rpb2 (860 bp) and tef1 (247 bp) sequences of ex-type strains of species in
section Radicina. The phylogenetic tree was rooted to Alternaria caricis CBS 480.90 and A. scirpicola CBS 481.90 (section Nimbya). Bootstrap support values above 70 % are
shown at the nodes. GenBank accession numbers are indicated in Table 2. The novel species described in this study are indicated in bold. T indicates ex-type strain.
TreeBASE: S23788.
MARIN-FELIX ET AL.
percurrent conidiophores with apically swollen conidiogenous crops, including cereals, ornamentals, oil crops, vegetables such
cells. as broccoli, cauliflower, carrot, onion and potato, and fruits like
Extensive morphological investigations of the genus Alternaria apple, citrus, pear and strawberry, among others. Species in
were carried out by Emory G. Simmons, which culminated with section Alternaria, such as A. alternata, A. arborescens or
his monograph on Alternaria species identification (Simmons A. tenuissima, as well as others from sections Alternantherae,
2007). Based on the sporulation patterns and conidial Brassiccicola, Crivellia, Gypsophilae, Nimbya, Radicina or Son-
morphology, he described several Alternaria species-groups chi, are frequently reported causing such diseases, but the
which were typified by representative species (Simmons largest group of phytopathogens in the genus is concentrated in
1992). In recent years, based on molecular phylogenetic ap- section Porri (Lawrence et al. 2016, Meena et al. 2017). The
proaches using DNA sequence data, it has been shown that the most relevant plant pathogens in this latter section are
main morphological groups identified by Simmons represent A. bataticola, A. porri, A. solani and A. tomatophila (Woudenberg
monophyletic species groups. Lawrence et al. (2013) provided et al. 2014). Alternaria species also produce diverse phytotoxins,
the first strongly supported phylogenetic hypothesis among which affect their host plants at different stages of pathogenesis
Alternaria lineages and elevated several of those monophyletic (Thomma 2003, Lawrence et al. 2008, Meena et al. 2017). Some
species groups to the taxonomic status of sections, each with a of these phytotoxins have been evaluated by the European Food
type species. Successive phylogenetic investigations added Safety Authority as potentially causing risks to human health
additional sections within the genus by synonymising genera (Meena et al. 2017).
such as Allewia, Brachycladium, Chalastospora, Chmelia, Cri- In humans, Alternaria species are commonly associated with
vellia, Embellisia, Nimbya, Pseudoalternaria, Sinomyces, Ter- hypersensitivity pneumonitis, bronchial asthma, allergic sinusitis
etispora, Ulocladium, Undiphilum and Ybotromyces and rhinitis. To a lesser extent, they have been also described as
(Woudenberg et al. 2013, 2014, Lawrence et al. 2016). There- causing paranasal sinusitis, ocular infections, onychomycosis,
fore, the genus Alternaria currently comprises close to 280 cutaneous and subcutaneous infections, granulomatous pulmo-
species, most of them classified in 27 sections. Taxonomic traits nary disease, soft palate perforation and disseminated disease
and species composition of all Alternaria sections are summar- (Pastor & Guarro 2008, de Hoog et al. 2011).
ised in Lawrence et al. (2016). In several surveys of microfungi from Spanish regions with
Considering, however, the overlap of morphological traits different climates and biodiversity, samples of plant litter (leaves,
among Alternaria sections/species and that the culture condi- bark and twigs) and dung of wild and farm herbivore animals
tions can greatly influence the morphology of these fungi, (rabbits, rodents, goats, cattle and horses) were collected. From
molecular identification is practically mandatory for the clas- these samples, we found 16 interesting Alternaria isolates,
sification of Alternaria isolates. Although the ITS barcode is belonging to sections Infectoriae, Pseudoalternaria, Chalasto-
considered a good phylogenetic marker to define sections, it spora and Radicina. The multi-locus phylogenetic analysis based
has limited discriminatory power to distinguish species, making on five above-mentioned gene markers showed that 10 of them
multi-locus sequence analysis with several protein-coding loci were undescribed species for the genus, and the others were
essential for accurate species identification. While identified as A. kourtkuyana, A. rosae and A. malorum (Figs
Woudenberg et al. (2013), in addition to the nrDNA regions, 5–7). Most of these novel species have been isolated from
used the combination of gapdh, rpb2 and tef1 loci for rede- herbivore dung, which appear to represent a reservoir of inter-
fining the genus, the combination of other phylogenetic esting Alternaria species which could represent potential plant
markers has since been analysed to determine relationships pathogens.
and species delineation in studies on a particular section; i.e.
References: Ellis 1976, Simmons 2007 (morphology); Laemmlen
ITS, Alt a-1, endoPG, gapdh, OPA10-2, rpb2 and tef1 for
2001, Thomma 2003, Lawrence et al. 2008, Meena et al. 2017
section Alternaria (Woudenberg et al. 2015); ITS, ATPase, tef1
(plant infections); Pastor & Guarro 2008, de Hoog et al. 2011
and gapdh for sections Infectoriae and Pseudoalternaria
(human infections); Woudenberg et al. 2013, 2014, 2015,
(Andersen et al. 2009, Deng et al. 2018, Poursafar et al.
Grum-Grzhimaylo et al. 2016, Lawrence et al. 2016, Poursafar
2018); ITS, Alt a-1, gapdh, rpb2 and tef1 for section Porri
et al. 2018 (morphology and phylogeny).
(Woudenberg et al. 2014); and ITS, Alt a-1 and gapdh for
section Sonchi (Lawrence et al. 2012, Deng et al. 2014). Alternaria aconidiophora Iturrieta-Gonzalez, Dania García &
Nevertheless, according to Lawrence et al. (2013) the plasma Gene, sp. nov. MycoBank MB829626. Fig. 8.
membrane ATPase, cmdA, and Alt a-1 loci are the most
Etymology: Name refers to the lack of conidiophores from
informative markers for Alternaria species delimitation. How-
vegetative hyphae.
ever, considering that the latter locus unreliably amplifies some
species within sect. Infectoriae, they suggested that the most Asexual morph on PCA: Mycelium superficial and immersed.
suitable genetic markers for molecular identification at the Hyphae 1–4 μm wide, septate, branched, hyaline to greyish,
species level are ATPase and cmdA genes (Lawrence et al. smooth-walled. Conidiophores absent. Conidiogenous loci
2013, 2016). Unfortunately, the latter marker has not been inconspicuous on vegetative hyphae, scarce. Conidia commonly
used for the phylogeny of any of the above-mentioned solitary at centre of the colony, globose, ovoid, near ellipsoid or
sections. obclavate, 12–31 × 7–12 μm, with some darkened middle
Alternaria is a very successful pathogenic genus that causes transverse septa, 1–5 transverse, 0–1(–2) longitudinal or obli-
disease on a great number of economically important plants, que septa per transverse segment, brown, smooth-walled.
causing large economic losses due to the number of plant Secondary conidiophores present, may be formed apically
species affected and worldwide distributions of several Alternaria from the conidial body as a short extension often geniculate, with
species (Meena et al. 2017). They are commonly described one or two, terminal or subterminal conidiogenous loci. Sexual
causing stem canker, leaf blight or leaf spot on a large variety of morph not observed.
18
Fig. 8. Alternaria aconidiophora (ex-type FMR 17111). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–F. Conidia. Scale bars = 10 μm.
Culture characteristics: Colonies on PDA reaching 64 mm diam Asexual morph on PCA: Mycelium superficial and immersed.
after 1 wk at 25 °C, flat, cottony at centre, slightly radially folded Hyphae 2–4 μm wide, branched, pale yellowish brown to brown,
towards the periphery, aerial mycelium abundant, margins reg- septate, smooth-walled to verruculose. Conidiophores macro-
ular; surface white (1A1); reverse yellowish white (4A2). On PCA nematous, arising laterally or terminally from aerial hyphae, erect
attaining 54 mm diam, flat, aerial mycelium scarce, margins to slightly flexuous, unbranched, occasionally branched, up to
regular; surface greyish green to greenish grey (1D3/1B1); 10-septate, 12–88 × 3–4 μm, brown becoming pale towards
reverse greenish grey (1C2/1B1). On OA reaching 61 mm diam, apex, smooth-walled, with 1 terminal and up to 3 subterminal
flat, aerial mycelium scarce, margins regular; surface and conidiogenous loci. Conidia in branched chains, occasionally
reverse colourless. solitary, ovoid, obclaviform, ellipsoidal or somewhat cylindrical,
9–43 × 6–8 μm, with darkened middle transverse septa, (1–)
Cardinal temperature for growth: Optimum 25 °C, maximum 2–3(–6) transverse, 0–1 longitudinal or oblique septa in up to 4
37 °C, minimum 5 °C. of the transverse segments, usually inconspicuous, pale
Typus: Spain, Catalonia, Alta Ribagorça, Vall de Boí, isolated yellowish to yellowish brown, verrucose. Secondary co-
from forest leaf litter, Dec. 2017, J. Gene (holotype CBS H- nidiophores commonly formed apically as a beak from conidial
23891, culture ex-type CBS 145419 = FMR 17111). body, or as a lateral conidiogenous loci from body cells bearing
conidia in short chains. Sexual morph not observed.
Notes: Alternaria aconidiophora together with A. fimeti, both
species introduced here from herbivore dung, are placed in an Culture characteristics: Colonies on PDA reaching
unsupported clade in Alternaria section Infectoriae (Fig. 5). 73 mm diam after 1 wk at 25 °C, flat, densely floccose, aerial
Morphologically, the latter differs from A. aconidiophora in mycelium abundant, margins fimbriate; surface olive brown to
having conspicuous sporulation with well-differentiated co- blond (4D3/4C4), white at the periphery; reverse yellowish
nidiophores and verrucose conidia up to 44 μm long. The brown to orange-grey (5E4/5B2). On PCA attaining
conidia of A. aconidiophora are smooth-walled and 12–31 μm 66 mm diam, flat, granular, aerial mycelium scarce, margins
long. regular; surface dark green (30F8); reverse dull green (30E4).
On OA reaching 70 mm diam, flat, loosely floccose at centre,
Alternaria altcampina Iturrieta-Gonzalez, Dania García & aerial mycelium scarce, margins regular; surface dark green
Gene, sp. nov. MycoBank MB829627. Fig. 9. (28F4); reverse dull green (29E3).
Etymology: Name refers to the region of Alt Camp (Catalonia), Cardinal temperature for growth: Optimum 25 °C, maximum
from where the fungus was collected. 37 °C, minimum 5 °C.
MARIN-FELIX ET AL.
Fig. 9. Alternaria altcampina (ex-type FMR 16476). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–I. Conidiophores and conidia. Scale bars: D–E = 20 μm;
F–I = 10 μm.
Typus: Spain, Catalonia, Alt Camp, isolated from goat dung, Mar. Alternaria parvicaespitosa differs in having smaller conidia
2017, I. Iturrieta-Gonzalez, M. Guevara-Suarez & J. Guarro (10–25 × 7–12 μm) with smooth to slightly punctulate outer walls
(holotype CBS H-23892, culture ex-type CBS 145420 = FMR (Gannibal & Lawrence 2016), and A. kordkuyana by its larger
16476). conidia [30–50(−60) × 7–11 μm] and shorter conidiophores
(10–40 × 3–4 μm) (Poursafar et al. 2018).
Notes: Based on the phylogeny of ITS, ATPase and gapdh,
A. altcampina is classified in Alternaria section Pseudoalternaria
Alternaria chlamydosporifera Iturrieta-Gonzalez, Dania García
(Fig. 6). It is closely related to the recently described species
& Gene, sp. nov. MycoBank MB829628. Fig. 10.
A. parvicaespitosa, which was isolated from harvested blueberry
fruit (California, USA), and A. kordkuyana, isolated from symp- Etymology: Name refers to the production of abundant chla-
tomatic wheat heads of Triticum aestivum (Kordkuy, Iran). mydospores in culture.
20
Fig. 10. Alternaria chlamydosporifera (ex-type FMR 17360). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–H. Conidiophores and conidia. I. Chla-
mydospores. Scale bars: D = 50 μm; E, I = 20 μm; F–H = 10 μm.
Asexual morph on PCA: Mycelium superficial and immersed. Secondary conidiophores can be formed apically from conidial
Hyphae 3–6 μm wide, septate, branched, pale brown to brown, body as a beak, geniculate, with 1–3 terminal or lateral con-
smooth-walled. Conidiophores macronematous, arising directly idiogenous loci, bearing solitary or short chains of conidia.
from aerial hyphae, erect to slightly flexuous, occasionally Chlamydospores abundant, immersed, intercalary, irregular
geniculate at apex, 1–4-septate, unbranched or scarcely shape, rarely broadly ellipsoidal or clavate, muriform, sometimes
branched, 14–140 × 3–5 μm, dark brown, verruculose, with 1–2 showing central constriction, 60–91 × 32–57 μm, dark brown to
conidiogenous loci. Conidia mostly solitary, occasionally in short black. Sexual morph not observed.
chains with up to two conidia, ellipsoidal or ovoid, occasionally
subglobose, 12–41 × 7–20 μm, with darkened middle transverse Culture characteristics: Colonies on PDA reaching 82 mm diam
septa, 1–3(–4) transverse, and 0–1(–2) longitudinal septa per after 1 wk at 25 °C, flat, densely floccose, aerial mycelium
transverse segments, brown to dark brown, verruculose. abundant, margins regular; surface greyish brown (5E3); reverse
MARIN-FELIX ET AL.
black to greyish brown (5E3). On PCA attaining 68 mm diam, flat, Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park,
with granular appearance by the presence of abundant chla- isolated from goat dung, Oct. 2017, G. Siso & C. Gonzalez-
mydospores, aerial mycelium scarce, margins regular; surface García (holotype CBS H-23894, culture ex-type CBS
dark green (29F5); reverse dark green (30F8). On OA reaching 145422 = FMR 16901).
71 mm diam, flat, loosely floccose at centre, slightly granular
Notes: Alternaria curvata was included in the section
towards the periphery, aerial mycelium scarce, margins slightly
Infectoriae, forming an unsupported basal clade together
lobate; surface dark green (29F4); reverse dark green (29F4).
with A. montsantina and A. pseudoventricosa, both intro-
Cardinal temperature for growth: Optimum 25 °C, maximum duced here, and A. ventricosa (Fig. 5). Morphologically, the
37 °C, minimum 5 °C. former two species differ from A. curvata in lacking curved
conidia. Alternaria ventricosa has asymmetrical conidia, due
Typus: Spain, Huesca, Baells, isolated from rabbit dung, Apr.
to the hyperplasia and hypertrophy of cells, especially on
2018, G. Siso & D. García (holotype CBS H-23893, culture ex-
one side of the conidia (Roberts 2007). Other morphologi-
type CBS 145421 = FMR 17360).
cally similar species are A. fimeti and A. triticina, which also
Notes: Alternaria chlamydosporifera belongs to Alternaria have curved conidia. However, A. triticina is phylogenetically
section Radicina (Fig. 7). It is included in a well-supported more distant and its conidia are strongly inequilateral and
clade (95 % BS) with A. petroselini, A. selini and A. smyrnii, wider (up to 22 μm) (Simmons 2007) than those of
which are pathogens of Apiaceae (Lawrence et al. 2016). A. curvata (4 – 16 μm wide). Alternaria fimeti can be differ-
Alternaria petroselini and A. selini can be easily differentiated entiated from A. curvata by the production of longer co-
by the lack of chlamydospores in culture and their larger nidiophores (22– 182 μm in A. fimeti vs. 23– 80 μm in
(50–66 μm in A. petroselini vs. 48–65 μm in A. selini) and A. curvata) and the absence or scarce development of
usually ellipsoidal conidia (Simmons 1995). Although secondary conidiophores.
A. smyrnii, the closest relative to A. chlamydosporifera, has
Alternaria fimeti Iturrieta-Gonzalez, Dania García & Gene, sp.
been described as producing sclerotial knots in culture that
nov. MycoBank MB829630. Fig. 12.
are able to form fertile conidiophores, its conidia are consid-
erably longer (67–96 μm) (Simmons 1995) than those of Etymology: Name refers to the substrate where the species was
A. chlamydosporifera (12–41 μm long). In addition, we have isolated, herbivore dung.
never observed conidiophores associated with the chlamydo-
Asexual morph on PCA: Mycelium superficial and immersed.
spores of the latter species.
Hyphae 2–5 μm wide, septate, branched, hyaline to subhya-
Alternaria curvata Iturrieta-Gonzalez, Dania García & Gene, line to pale yellowish, verruculose. Conidiophores semi- to
sp. nov. MycoBank MB829628. Fig. 11. macronematous, arising laterally or terminally from aerial hy-
phae, erect to slightly flexuous, unbranched (can be slightly
Etymology: Name refers to the presence of curved conidia.
branched on OA), up to 9-septate, 22–182 × 1–5 μm, pale
Asexual morph on PCA: Mycelium superficial and immersed. brown, smooth-walled, with 1 terminal conidiogenous locus.
Hyphae 2–6 μm wide, septate, branched, hyaline to yellowish Conidia solitary or in short chains of up to six conidia, ovoid,
brown to brown, smooth-walled to verruculose. Conidiophores obpyriform or obclavate, some slightly curved,
macronematous, arising laterally or terminally from aerial hy- 9–44 × 5–14(–23) μm, with darkened middle transverse
phae, erect to slightly flexuous, usually unbranched, up to 14- septa, 0–5 transverse (up to 7 in OA), and 0–1(–2) longi-
septate, 23–80 × 3–5 μm, brown to dark brown, smooth- tudinal or oblique septa per segment, brown, verrucose.
walled, with a terminal or occasionally a sub-terminal con- Secondary conidiophores only scarcely produced on OA as
idiogenous loci. Conidia forming branched chains, with up to 5 apical or lateral extension from conidial body, up to 25 μm
conidia in unbranched part, ovoid or nearly ellipsoidal, often long. Sexual morph not observed.
slightly curved, 13–47(–70) × 4–16 μm, with darkened middle
Culture characteristics: Colonies on PDA reaching 66 mm diam
transverse septa, (0–)1–5(–7) transverse, and 0–2(–3) longi-
after 1 wk at 25 °C, flat, densely floccose, aerial mycelium
tudinal or oblique septa per transverse segment, brown to dark
abundant, margins fimbriate; surface yellowish grey to yellowish
brown, verrucose to tuberculate. Secondary conidiophores can
white (3C2/3A2); reverse yellowish brown to light yellow (5E8/
be formed apically or laterally from the conidial body as a short
4A5). On PCA attaining 65 mm diam, flat, slightly floccose at
extension bearing conidia in short chains. Sexual morph not
centre, aerial mycelium scarce, margins regular; surface olive-
observed.
brown (4F5); reverse olive-brown (4F8/4E4). On OA reaching
Culture characteristics: Colonies on PDA reaching 63 mm diam 64 mm diam, flat, slightly floccose, scarce aerial mycelium,
after 1 wk at 25 °C, flat, densely floccose, aerial mycelium margins regular; surface dull green (30E5) with grey floccose
abundant, margins regular; surface white to dull green (1A1/ area; reverse dull green (30E4).
30D4); reverse dark green to olive yellow (30F8/2D6), white at
Cardinal temperature for growth: Optimum 25 °C, maximum
the periphery. On PCA attaining 62 mm diam, flat, loosely floc-
37 °C, minimum 5 °C.
cose, aerial mycelium scarce, margins regular; surface olive
(3F4); reverse dark green to grey (29F4/29B1). On OA reaching Typus: Spain, Catalonia, Priorat, Montsant Natural Park, Arbolí,
61 mm diam, scarce aerial mycelium towards the periphery, isolated from small rodent dung, Feb. 2018, I. Iturrieta-Gonzalez,
margins regular; surface greyish green (30E5), with greyish E. Carvalho & J. Gene (holotype CBS H-23895, culture ex-type
mycelium tufts at centre; reverse dull green (29E4). CBS 145423 = FMR 17110).
Cardinal temperature for growth: Optimum 25 °C, maximum Note: Alternaria fimeti is placed in a clade of section Infectoriae
37 °C, minimum 5 °C. together with A. aconidiophora (see notes of this latter species).
22
Fig. 11. Alternaria curvata (ex-type FMR 16901). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–F. Conidiophore and conidia. Scale bars = 10 μm.
Alternaria inflata Iturrieta-Gonzalez, Dania García & Gene, sp. scarce aerial mycelium, margins regular; surface dull green to
nov. MycoBank MB829631. Fig. 13. grey (30E4/30B1); reverse dark green to grey (30E4/30B1). On
OA reaching 61 mm diam, flat, loosely floccose, margins regular;
Etymology: Name refers to the presence of swollen cells in the
surface pale grey (1B1); reverse pale grey (1B1).
conidial body.
Asexual morph on PCA: Mycelium superficial and immersed.
Hyphae of 1–3 μm wide, septate, branched, hyaline to pale
brown, smooth-walled to verruculose. Conidiophores arising
Typus: Spain, Catalonia, Conca de Barbera, Poblet, isolated
laterally or terminally from aerial hyphae, erect to slightly flex-
from rabbit dung, Mar. 2017, J. Guarro & I. Iturrieta-Gonzalez
uous, semi- to macronematous, up to 10-septate, commonly
(holotype CBS H-23896, culture ex-type CBS 145424 = FMR
unbranched, 9–73(–105) × 2–5 μm, pale brown to brown, 16477).
smooth-walled, with one terminal conidiogenous loci or up to
three subterminal conidiogenous loci. Conidia solitary or in short
Notes: Our phylogeny shows that A. inflata belongs to section
chains with up to four conidia, broadly ellipsoidal or ovoid,
Pseudoalternaria (Fig. 6). It clustered in a well-supported clade
13–41 × 5–14 μm, often with some swollen cells protruding the
(86 % BS) with A. altcampina, A. kordkuyana and
conidium outline, some with darkened middle transverse septa,
A. parvicaespitosa, but formed a single basal lineage repre-
(1–)2–3(–5) transverse septa, and 0–2 longitudinal or oblique
sentative of a distinct species. Alternaria inflata can be differ-
septa per transverse segment, brown, verruculose. Secondary
entiated from all the species in the section by the production of
conidiophores scarcely produced, as an apical extension up to
chlamydospores and by the formation of broadly ellipsoidal
15 μm long, bearing conidia in short chains. Chlamydospores
conidia, usually with swollen cells protruding from the conidial
present, consisting of intercalary, thick-walled, brown swollen
body. In addition, A. altcampina also differs in the production of
cells, up to 8 × 6 μm, arranged in chains or in clusters. Sexual
secondary conidiophores, A. parvicaespitosa in its shorter co-
morph not observed.
nidiophores (up to 70 μm) and conidia (10–25 μm) (Gannibal &
Culture characteristics: Colonies on PDA reaching 62 mm diam Lawrence 2016), and A. kordkuyana in the production of longer
after 1 wk at 25 °C, flat, aerial mycelium abundant, floccose, conidial chains [up 5–8(–10) conidia] and conidia measuring
margins fimbriate; surface white (1A1); reverse greyish yellow to 30–50(–60) × 7–11 μm with up to seven transverse septa
yellowish white (4C6/4A2). On PCA attaining 67 mm diam, flat, (Poursafar et al. 2018).
MARIN-FELIX ET AL.
Fig. 12. Alternaria fimeti (ex-type FMR 17110). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–G. Conidiophores and conidia. Scale bars: D = 20 μm;
E–G = 10 μm.
With the additions of A. altcampina and A. inflata section longitudinal or oblique septa, pale brown to brown, verrucose to
Pseudoalternaria now comprises seven species. It is of note, tuberculate. Secondary conidiophores commonly formed apically
however, that most of these taxa are only known from a single on conidia as a geniculate extension with several conidiogenous
collection. In our survey of asexual fungi from herbivore dung, we loci, or as lateral extensions from cells of conidial body, up to
identified several Spanish isolates belonging to other species in 35 μm long, producing conidia solitary or in short chains. Sexual
the section, namely A. kordkuyana and A. rosae (Fig. 6). morph not observed.
Considering that most of the species in section Pseudoalternaria
are mainly associated with herbaceous plants of the families
after 1 wk at 25 °C, low convex, cottony, aerial mycelium
Brassicaceae, Ericaceae, Poacea or Rosaceae, it is not sur-
abundant, margins regular; surface white (1A1); reverse
prising to find these fungi in faeces of herbivorous animals.
yellowish brown to greyish yellow (5E7/4B6). On PCA attaining
Alternaria lawrencei Iturrieta-Gonzalez, Dania García & Gene, 69 mm diam, low convex, slightly floccose, aerial mycelium
sp. nov. MycoBank MB829632. Fig. 14. relatively abundant at centre, margins regular; surface yellowish
grey to olive (2D2/1E3); reverse dark green to olive (30F8/1E3).
Etymology: Name in honour of Daniel P. Lawrence for his
On OA reaching 63 mm diam, loosely floccose at centre, flat and
contribution to the taxonomy of the genus Alternaria.
scarce aerial mycelium towards the periphery, margins regular
Asexual morph on PCA: Mycelium superficial and immersed. and diffuse; surface olive (2F4) to olive-grey (2B1); reverse olive
Hyphae 1–3 μm wide, septate, branched, pale brown, smooth- to yellowish grey (2F8/2D2).
walled. Conidiophores macronematous, solitary, arising directly
from aerial hyphae, erect to slightly flexuous, occasionally Cardinal temperature for growth: Optimum 25 °C, maximum
geniculate at apex, usually unbranched, up to 10-septate, 37 °C, minimum 5 °C.
9–125 × 3–4(–5) μm, brown, smooth-walled, with 1–2 lateral or Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park,
terminal conidiogenous loci; micronematous conidiophores also isolated from goat dung, Oct. 2017, G. Siso & C. Gonzalez-
present, reduced to intercalary conidiogenous cells with a single García (holotype CBS H-23897, culture ex-type CBS
conidiogenous locus on hyphae. Conidia solitary or in short 145425 = FMR 17004).
chains, up to six conidia in the unbranched part, ovoid, obpyri-
form or obclavate, 6–71 × 7–15 μm, with darkened middle Notes: Although A. lawrencei is clearly placed in section Infec-
transverse septa, (1–)2–7(–9) transverse, and 0–2(–3) toriae, the multi-locus analysis did not reveal any phylogenetic
24
Fig. 13. Alternaria inflata (ex-type FMR 16477). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D, F–G. Conidiophores and conidia. E. Chlamydospores. Scale
bars = 10 μm.
relationship with any species included in the analysis (Fig. 4). It is brown, smooth-walled to verruculose. Conidia solitary or in
of note, however, that eight species of the section (i.e. A. cerasi, short chains with up to five conidia, subglobose, ovoid or
A. cesenica, A. dactylidicola, A. forlicesenensis, obpyriform, 8 – 65 × 6 –12 μm, with 1– 3( – 11) transverse
A. hampshirensis, A. litorea, A. murispora and A. poaceicola) septa, and 0 – 2 longitudinal or oblique septa, brown,
could not be included in our analysis due to their limited mo- verrucose to tuberculate. Secondary conidiophores
lecular data. Nevertheless, A. cesenica, A. dactylidicola, commonly produced apically as a long, often geniculate
A. forlicesenensis, A. hampshirensis, A. murispora and extension, up to 105 μm long and 10-septate, bearing
A. poaceicola can be distinguished from A. lawrencei by the terminal conidial chains. Sexual morph not observed.
production of a sexual morph (Ariyawansa et al. 2015b, Liu et al.
Culture characteristics: Colonies on PDA reaching
2015, Thambugala et al. 2017, Wanasinghe et al. 2018),
76 mm diam after 1 wk at 25 °C, flat, densely floccose,
A. cerasi by its inequilateral conidia (Potebnia 1907, Simmons
aerial mycelium abundant, margins regular; surface pastel
2007), and A. litorea by the production of shorter primary co-
grey to greyish yellow (1C1/2C4) and with a white edge;
nidiophores (40–50 μm long) and smooth-walled conidia that are
reverse blond to white (4C4/1A1). On PCA attaining
22–32 μm long (Simmons 2007).
70 mm diam, flat, loosely floccose at centre, aerial myce-
Alternaria montsantina Iturrieta-Gonzalez, Dania García & lium moderate, margins regular; surface olive brown to
Gene, sp. nov. MycoBank MB829633. Fig. 15. white (4D4/1A1); reverse olive to white (3D4/1A1). On OA
reaching 75 mm diam, flat, cottony, margins regular; sur-
Etymology: Name refers to the place, Montsant Natural Park
face yellowish grey to olive (3D2/2F4) and white edge;
(Catalonia), where the fungus was collected.
reverse olive to white (2F4/1A1).
Asexual morph on PCA: Mycelium superficial and
immersed. Hyphae 1 – 7 μm wide, septate, branched, usu-
ally forming hyphal coils, subhyaline to pale brown, smooth-
walled to verruculose. Conidiophores macronematous, Typus: Spain, Catalonia, Priorat, Montsant Natural Park,
arising laterally or terminally from aerial hyphae, erect to Swamp of Siurana, isolated from an unidentified twig, Feb.
slightly flexuous, unbranched, up to 15-septate, 2018, I. Iturrieta-Gonzalez, E. Carvalho & J. Gene (holo-
12 –137 × 3 – 6 μm, often with geniculate apical portion type CBS H-23898, culture ex-type CBS 145426 = FMR
containing intercalary and terminal conidiogenous loci, 17060).
MARIN-FELIX ET AL.
Fig. 14. Alternaria lawrencei (ex-type FMR 17004). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–K. Conidiophores and conidia. Scale bars: D = 20 μm;
E–K = 10 μm.
Notes: Alternaria montsantina is placed in a weakly supported Alternaria pobletensis Iturrieta-Gonzalez, Dania García &
basal clade of the section Infectoriae, together with A. curvata, Gene, sp. nov. MycoBank MB829634. Fig. 16.
A. pseudoventricosa and A. ventricosa (Fig. 5). Morphologically,
Etymology: Name refers to the place, Poblet (Catalonia), from
this new species can be distinguished from A. curvata and
where the species was collected.
A. ventricosa by the absence of curved or inequilateral inflated
conidia. Alternaria montsantina differs from A. pseudoventricosa Asexual morph on PCA: Mycelium superficial and immersed.
in the production of longer (12–137 μm) and often geniculate Hyphae 2–5 μm wide, branched, pale brown, septate, smooth-
primary and secondary conidiophores, bearing solitary conidia or walled. Conidiophores semi- to macronematous, solitary,
arranged in short chains (up to five conidia). Conidiophores in arising directly from aerial hyphae, erect to slightly flexuous,
A. pseudoventricosa are 30–44 μm long, and the conidial chains occasionally slightly geniculate at apex, unbranched or
include up to 19 conidia. branched, up to 8-septate, 14–82 × 4–5(–6) μm, brown,
26
Fig. 15. Alternaria montsantina (ex-type FMR 17060). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–H. Conidiophores and conidia. Scale bars: D = 50 μm;
E–H = 10 μm.
smooth-walled, with 1–2 lateral or terminal conidiogenous loci. Culture characteristics: Colonies on PDA reaching 46 mm diam
Conidia commonly in short, scarcely branched chains, with up to after 1 wk at 25 °C, flat, floccose at the centre, velvety towards
seven conidia, obpyriform or obclavate, some ellipsoidal or the periphery, aerial mycelium moderate, margins regular; sur-
subcylindrical, 8–50 × 5–20 μm, (1–)3–7(–9) transverse septa, face olive (3F8), whitish at the periphery; reverse black to
often middle septa darker, and 0–1(–2) longitudinal or oblique yellowish brown (5D5). On PCA attaining 58 mm diam, flat,
septa per transverse segment, pale brown to brown, smooth- velvety, margins regular; surface dark green to dull green (30F8/
walled or verruculose. Secondary conidiophores commonly 28D3); reverse dark green to dull green (30F8/28D3). On OA
produced apically as a short beak up to 11 μm long, or laterally reaching 55 mm diam, flat, loosely floccose, margins regular;
from cells of conidial body, bearing conidia in short chains. surface greyish green to dull green (29C3/29E4); reverse dark
Sexual morph not observed. green to dull green (30F8/30E3).
MARIN-FELIX ET AL.
Fig. 16. Alternaria pobletensis (ex-type FMR 16448). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–I. Conidiophores and conidia. Scale bars: D = 20 μm;
E–I = 10 μm.
Cardinal temperature for growth: Optimum 25 °C, maximum septa (Woudenberg et al. 2013); however, the conidia in
35 °C, minimum 5 °C. A. pobletensis usually have two or more longitudinal or oblique
septa. Its closest relative is A. breviramosa. This was originally
Typus: Spain, Catalonia, Conca de Barbera, Poblet, isolated
described as Chalastospora ellipsoidea, found on Triticum
from unidentified herbivore dung, Mar. 2017, J. Guarro & I.
(Poaceae) in Australia (Crous et al. 2009a), but later its name
Iturrieta-Gonzalez (holotype CBS H-23899, culture ex-type CBS
was changed to avoid confusion with Alternaria ellipsoidea, an
145427 = FMR 16448).
already described species from section Gypsophilae
Notes: Alternaria pobletensis clustered in section Chalastospora (Woudenberg et al. 2013). Section Gypsophilae contains all
in a single branch clearly separated from the other six species Alternaria species that occur on Caryophyllaceae (Lawrence
that currently comprise the section (Fig. 7). Other species of et al. 2016). Alternaria breviramosa differs from A. pobletensis
section Chalastospora rarely produce conidia with longitudinal by having shorter conidiophores (up to 25 μm), often reduced to
28
conidiogenous cells, and ellipsoidal, subcylindrical to fusoid floccose, margins regular; surface dull green (29E4); reverse dull
conidia with only transverse septa (Crous et al. 2009a). green (29E4).
Alternaria pseudoventricosa Iturrieta-Gonzalez, Dania García Cardinal temperature for growth: Optimum 25 °C, maximum
& Gene, sp. nov. MycoBank MB829635. Fig. 17. 37 °C, minimum 5 °C.
Etymology: Name refers to the apparent phylogenetic relation- Typus: Spain, Catalonia, Tarragona, Els Ports Natural Park,
ship to A. ventricosa. isolated from horse dung, Oct. 2017, G. Siso & C. Gonzalez-
García (holotype CBS H-23900, culture ex-type CBS
Asexual morph on PCA: Mycelium superficial and immersed. 145428 = FMR 16900).
Hyphae 1–7 μm wide, septate, branched, hyaline to pale brown,
Notes: Alternaria pseudoventricosa and A. ventricosa clustered
smooth-walled. Conidiophores macronematous, arising laterally
in an unsupported monophyletic basal clade in section Infec-
from aerial hyphae, erect to slightly flexuous, up to 4-septate,
toriae (Fig. 5). They can be differentiated by their conidial
unbranched, 30–45 × 4–6 μm, brown, smooth-walled, with one
morphology. Conidia in A. ventricosa are usually asymmetric,
terminal conidiogenous locus. Conidia commonly in unbranched
laterally swollen, and pale cinnamon brown (Roberts 2007). In
chains, with up to 19 conidia, obpyriform or obclavate,
contrast, those of A. pseudoventricosa are obpyriform or
10–48(–66) × 5–14 μm, with darkened middle transverse septa,
obclavate and brown to dark brown.
1–7 transverse, 0–1 longitudinal or oblique septa, brown to dark
brown, verrucose to tuberculate. Secondary conidiophores Authors: I. Iturrieta-Gonzalez, D. García, M. Hernandez-
scarce, as a beak arising from the conidial body. Sexual morph Restrepo & J. Gene
not observed.
Brunneosphaerella Crous, Stud. Mycol. 64: 31. 2009. Fig. 18.
Classification: Dothideomycetes, Dothideomycetidae, Capno-
after 1 wk at 25 °C, flat, cottony at the centre, floccose towards
diales, Mycosphaerellaceae.
the periphery, margins regular; surface white (1A1); reverse
yellowish white (4A2). On PCA attaining 62 mm diam, flat to- Type species: Brunneosphaerella protearum (Syd. & P. Syd.)
wards the periphery, margins regular; surface dark green (29F4), Crous, basionym: Leptosphaeria protearum Syd. & P. Syd.
with tuft of white aerial mycelium at centre; reverse dark green to Epitype and ex-epitype strain designated by Crous et al. (2011):
grey (29F8/29B1). On OA reaching 67 mm diam, flat, loosely CBS H-20335, CBS 130597 = CPC 16338.
Fig. 17. Alternaria pseudoventricosa (ex-type FMR 16900). A. Colonies on PDA. B. Colonies on PCA. C. Colonies on OA. D–F. Conidiophores and conidia. Scale bars:
D = 20 μm; E, F = 10 μm.
MARIN-FELIX ET AL.
Fig. 18. Brunneosphaerella spp. A–E. Disease symptoms. A, B. Brunneosphaerella protearum (epitype CBS H-20335). C. Brunneosphaerella nitidae (holotype CBS H-20334).
D, E. Ascomata visible on lesion surface. D. Brunneosphaerella protearum (CBS H-20335). E. Brunneosphaerella nitidae (holotype CBS H-20334). F–H. Vertical sections through
ascomata showing wall structure. F. Brunneosphaerella jonkershoekensis (holotype PREM 59447). G, H. Brunneosphaerella protearum (CBS H-20335). I–M. Asci. I. Brun-
neosphaerella jonkershoekensis (holotype PREM 59447). J, K. Brunneosphaerella nitidae (holotype CBS H-20334). L, M. Brunneosphaerella protearum (CBS H-20335). N–Q.
Ascospores. N, O. Brunneosphaerella jonkershoekensis (holotype PREM 59447). P. Brunneosphaerella nitidae (holotype CBS H-20334). Q. Brunneosphaerella protearum (CBS
H-20335). R, S. Germinating ascospores. R. Brunneosphaerella nitidae (holotype CBS H-20334). S. Brunneosphaerella protearum (CBS H-20335). Scale bars: G = 75 μm;
F, I = 50 μm; H, J–M, P–S = 10 μm; N, O = 5 μm. Pictures A, B, D, F–I, L–O, Q, S taken from Crous et al. (2009b); C, E, J, K, P, R from Crous et al. (2011).
30
Table 3. DNA barcodes of accepted Brunneosphaerella spp.

ITS chs rpb2 tef1
Brunneosphaerella jonkershoekensis CPC 13902ET JN712439 JN712609 MF951441 JN712571 Crous et al. (2011), Videira et al. (2017)
B. nitidae CBS 130595T GU214625 JN712619 MF951442 JN712581 Crous et al. (2009b, 2011), Videira et al. (2017)
ET
B. protearum CBS 130597 GU214626 JN712620 MF951443 JN712582 Crous et al. (2009b, 2011), Videira et al. (2017)
B. roupeliae CBS 144602T MK539950 – MK540080 – Present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at the Westerdijk Fungal Biodiversity Institute. T
and ET indicate ex-type and ex-epitype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; chs: partial chitin synthase-1 gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit
gene; tef1: partial translation elongation factor 1-alpha gene.
DNA barcodes (genus): LSU, ITS. Hosts: Protea spp. (Proteaceae).

DNA barcodes (species): chs, rpb2, tef1. Table 3. Disease symptoms: Leaf spots and Brunneosphaerella leaf
blight.
Ascomata amphigenous, immersed to semi-immersed, black,
single, gregarious, substomatal, pyriform or globose with a Notes: Brunneosphaerella was introduced by Crous et al.
papillate, periphysate ostiole; ascomatal wall consisting of three (2009b) to accommodate Leptosphaeria protearum, which is
strata of slightly compressed textura angularis, an outer stratum of a major leaf spot and blight pathogen of Protea spp. causing
dark brown, thick-walled cells, becoming paler in the central severe losses in plantations of South African Protea spp.
stratum, and hyaline, thin-walled in the inner stratum. Pseudo- wherever they are cultivated. Morphologically, Brunneos-
paraphyses absent. Asci clavate to cylindro-clavate, often curved, phaerella is distinct from Leptosphaeria in that ascospores are
tapering to a pedicel, narrowing slightly to a rounded apex with an always brown at maturity, and asexual morphs have brown,
indistinct ocular chamber, 8-spored, bitunicate with fissitunicate percurrently proliferating conidiogenous cells. A new species
dehiscence. Ascospores biseriate, fusiform, broader at the apical isolated from leaves of Protea repens in South Africa,
end, initially hyaline and 1-septate, becoming yellow-brown and 3- B. jonkershoekensis, was included in the genus when it was
septate at maturity, slightly constricted at median to supra-median introduced (Crous et al. 2009b). This species appears to be a
septum (adapted from Crous et al. 2009b). serious pathogen of Pr. repens in the Western Cape Province
of South Africa. Subsequently, Crous et al. (2011) described
Culture characteristics: Colonies on OA spreading, flat, with
the third species known from the genus, B. nitidae. This was
sparse to moderate aerial mycelium, lobate and smooth, rarely
isolated from the same area as B. jonkershoekensis, but
feathery, margins. On OA surface pale luteous, smoke grey with
B. nitidae was isolated from leaves of Pr. nitida, causing leaf
submerged iron-grey margins, or olivaceous grey with iron-grey,
spots on this host. Thus, the genus comprises four species, all
pale olivaceous grey and smoke grey patches. On PDA surface
of which were isolated from species of Protea in South Africa.
olivaceous grey, sometimes with pale olivaceous grey to smoke
The ITS sequences of the four species are highly similar.
grey patches, or smoke grey with iron-grey margins; reverse
However, these can be easily delimited based on the chs,
iron-grey. On MEA surface pale olivaceous grey, smoke grey,
rpb2 and tef1 sequences.
dirty white with patches of smoke grey, or smoke grey with dirty
white and olivaceous grey patches and submerged iron-grey References: Crous et al. 2009b, 2011, Videira et al. 2017
margins; reverse iron-grey or olivaceous grey. (morphology and phylogeny).
Optimal media and cultivation conditions: MEA, OA, PDA and Brunneosphaerella roupeliae Crous, sp. nov. MycoBank
SNA at 25 °C under near-ultraviolet light to promote sporulation. MB829609. Fig. 19.
Distribution: Africa, mainly reported from South Africa. Also re- Etymology: Name refers to Protea roupeliae, the host species
ported from Pacific Islands (Hawaii) and Europe (Portugal and from which it was collected.
Spain).
Fig. 19. Brunneosphaerella roupeliae (ex-type CBS 144602). A. Close-up of leaf spot with ascomata. B, C. Asci with ascospores. D. Germinating ascospores. Scale
bars = 10 μm.
MARIN-FELIX ET AL.
Fig. 20. Disease symptoms of Elsinoe spp. A. Elsinoe ampelina on Vitis vinifera. B. Elsinoe asclepiadea on Asclepias mellodora. C. Elsinoe bidentis on Bidens segetum. D.
Elsinoe erythrinae on Erythrina sp. E. Elsinoe eucalypticola on Eucalyptus sp. F. Elsinoe fawcettii on Citrus sp. G. Elsinoe freyliniae on Freylinia lanceolata. H. Elsinoe perseae
on Persea americana. I. Elsinoe othonnae on Othonna quinquedentata. J. Elsinoe poinsettiae on Euphorbia sp. K. Elsinoe punicae on Punica granatum. L. Elsinoe terminaliae
on Terminalia catappa. Pictures taken from Fan et al. (2017).
32
Fig. 21. Elsinoe spp. A–J. Sexual morph. A. Subcutilar ascoma of Elsinoe fecunda. B–D. Asci. B. Elsinoe eucalypticola (ex-type CBS 124765). C, D. Elsinoe fecunda
(holotype, PREM 56503). E–J. Ascospores. E–H. Elsinoe eucalypticola (ex-type CBS 124765). I, J. Elsinoe fecunda. K–P. Asexual morph. K, L. Conidiophores. K. Elsinoe
asclepiadea (ex-type CPC 18544). L. Elsinoe othonnae (ex-type CBS 139910). M–P. Conidia. M. Elsinoe asclepiadea (ex-type CPC 18544). N. Elsinoe erythrinae (ex-epitype
CPC 18542). O. Elsinoe tectificae (ex-type CBS 124777). P. Elsinoe othonnae (ex-type CBS 139910). Scale bars: A = 100 μm; B–D, K–U = 10 μm; E–J = 5 μm; C applies to C
and B; J applies I and J. Pictures taken from Fan et al. (2017).
Leaf spots amphigenous, sub-circular, 5–20 mm diam, me- 10 mm diam after 2 wk at 25 °C. On MEA surface pale oliva-
dium brown, with raised, dark brown border. Ascomata ceous grey, reverse olivaceous grey; on PDA surface and
pseudothecial, amphigenous, black, immersed to erumpent, reverse olivaceous grey, and on OA surface pale luteous.
globose, to 250 μm diam, with apical ostiole; ascomatal wall
Typus: South Africa, KwaZulu-Natal Province, Drakensberg,
of 2–3 layers of brown cells of textura angularis. Asci
Monks Cowl, on leaves of Protea roupeliae (Proteaceae), 18
aparaphysate, fasciculate, bitunicate, subsessile, ellipsoid-
Jan. 2010, A. Wood, HPC 1522 (holotype CBS H-23847, culture
fusoid, straight to slightly curved, 8-spored,
ex-type CPC 32914 = CBS 144602).
65–110 × 11–15 μm. Ascospores bi- to triseriate, over-
lapping, guttulate, thick-walled, straight to slightly curved, Notes: Brunneosphaerella roupeliae was isolated from the same
obovoid with obtuse ends, widest in middle of apical cell, 3- host genus in South Africa as the other three species of the
septate, constricted at median septum, tapering towards both genus, Protea. The ITS and LSU sequences located this species
ends, but more prominently towards lower end, (19–) in the genus Brunneosphaerella since both sequences showed
22–23( –25) × 5( –6) μm. Ascospores germinating from both more than 99 % of nucleotide similarity with the ex-type strains of
ends, becoming brown and verruculose, constricted at pri- the other three species. The rpb2 sequence showed a nucleotide
mary septum, with germ tubes parallel to the long axis, similarity of 95.86 % with the ex-epitype strain of B. protearum,
ascospore becoming 7 –9 μm diam. 95.45 % with the ex-epitype strain of B. jonkershoekensis, and
94.5 % with the ex-type strain of B. nitidae. The tef1 sequence
Culture characteristics: Colonies erumpent, spreading, with showed a nucleotide similarity of 95.66 % with the ex-epitype
moderate aerial mycelium, and even, lobate margins, reaching strain of B. protearum and the ex-type strain of B. nitidae, and
MARIN-FELIX ET AL.
Table 4. DNA barcodes of accepted Elsinoe spp.

ITS LSU rpb2 tef1
Elsinoe abutilonis CBS 510.50T KX887185 KX886949 KX887068 KX886831 Fan et al. (2017)
E. ampelina CBS 208.25 KX887186 KX886950 KX887069 KX886832 Fan et al. (2017)
T
E. anacardii CBS 470.62 KX887189 KX886953 KX887072 KX886835 Fan et al. (2017)
E. annonae CBS 228.64 KX887190 KX886954 KX887073 KX886836 Fan et al. (2017)
T
E. arachidis CBS 511.50 KX887191 KX886955 KX887074 KX886837 Fan et al. (2017)
E. arrudai CBS 220.50T KX887194 KX886958 KX887077 KX886840 Fan et al. (2017)
T
E. asclepiadea CBS 141937 KX887195 KX886959 KX887078 KX886841 Fan et al. (2017)
E. australis CBS 314.32ET KX887198 KX886962 KX887081 KX886844 Fan et al. (2017)
E. banksiae STE-U 2678 T
AF227197 – – – Swart et al. (2001)
E. banksiicola CBS 113734T KX887199 KX886963 KX887082 KX886845 Fan et al. (2017)
T
E. barleriicola CBS 471.62 KX887200 KX886964 KX887083 KX886846 Fan et al. (2017)
E. bidentis CBS 512.50T KX887201 KX886965 KX887084 KX886847 Fan et al. (2017)
E. brasiliensis CPC 18528 ET
KX887204 – KX887087 KX886850 Fan et al. (2017)
E. caleae CBS 221.50T KX887205 KX886968 KX887088 KX886851 Fan et al. (2017)
T
E. centrolobii CBS 222.50 KX887206 KX886969 KX887089 KX886852 Fan et al. (2017)
E. citricola CPC 18535T KX887207 KX886970 KX887090 KX886853 Fan et al. (2017)
T
E. coryli CBS 275.76 KX887209 KX886972 KX887092 KX886855 Fan et al. (2017)
E. diospyri CBS 223.50T KX887210 KX886973 KX887093 KX886856 Fan et al. (2017)
E. eelemani DAR 83016T KX372292 – KX398204 KX398203 Crous et al. (2016a)
E. embeliae CBS 472.62 T
KX887211 KX886974 – KX886857 Fan et al. (2017)
E. erythrinae CPC 18542ET KX887214 KX886977 KX887096 KX886860 Fan et al. (2017)
T
E. eucalypticola CBS 124765 KX887215 KX886978 KX887097 KX886861 Fan et al. (2017)
E. eucalyptorum CBS 120084T KX887216 KX886979 KX887098 KX886862 Fan et al. (2017)
T
E. euphorbiae CBS 401.63 KX887217 KX886980 KX887099 KX886863 Fan et al. (2017)
E. fagarae CBS 514.50T KX887218 KX886981 KX887100 KX886864 Fan et al. (2017)
T
E. fawcettii CBS 139.25 KX887219 KX886982 KX887101 KX886865 Fan et al. (2017)
E. fici CBS 515.50 KX887223 KX886986 KX887105 KX886869 Fan et al. (2017)
E. fici-caricae CBS 473.62 T
KX887224 KX886987 KX887106 KX886870 Fan et al. (2017)
E. flacourtiae CBS 474.62T KX887225 KX886988 KX887107 KX886871 Fan et al. (2017)
T
E. freyliniae CBS 128204 KX887226 KX886989 KX887108 KX886872 Fan et al. (2017)
E. genipae CBS 342.39T KX887227 KX886990 KX887109 KX886873 Fan et al. (2017)
T
E. genipae-americanae CBS 516.50 KX887228 KX886991 KX887110 KX886874 Fan et al. (2017)
E. glycines CBS 389.64ET KX887229 KX886992 KX887111 KX886875 Fan et al. (2017)
T
E. hederae CBS 517.50 KX887231 KX886994 KX887113 KX886877 Fan et al. (2017)
E. ichnocarpi CBS 475.62T KX887232 KX886995 KX887114 KX886878 Fan et al. (2017)
T
E. jasminae CBS 224.50 KX887233 KX886996 KX887115 KX886879 Fan et al. (2017)
E. jasminicola CBS 212.63T KX887234 KX886997 – KX886880 Fan et al. (2017)
ET
E. krugii CPC 18531 KX887235 KX886998 KX887116 KX886881 Fan et al. (2017)
E. lagoa-santensis CBS 518.50T KX887239 KX887002 KX887120 KX886885 Fan et al. (2017)
ET
E. ledi CBS 167.33 KX887240 KX887003 KX887121 KX886886 Fan et al. (2017)
E. lepagei CBS 225.50T KX887241 KX887004 KX887122 – Fan et al. (2017)
T
E. leucopogonis CBS 144439 MH327822 MH327858 MH327885 MH327897 Crous et al. (2018)
E. leucospermi CBS 111207T KX887242 KX887005 KX887123 KX886887 Fan et al. (2017)
T
E. lippiae CBS 166.40 KX887248 KX887011 KX887129 KX886893 Fan et al. (2017)
E. mangiferae CBS 226.50T KX887249 KX887012 KX887130 KX886894 Fan et al. (2017)
E. mattiroloanum CBS 287.64 KX887250 KX887013 KX887131 KX886895 Fan et al. (2017)
E. menthae CBS 322.37ET KX887253 KX887016 KX887134 KX886898 Fan et al. (2017)
E. mimosae CPC 19478ET KX887255 KX887018 KX887136 KX886900 Fan et al. (2017)
E. oleae CBS 227.59T KX887256 KX887019 KX887137 KX886901 Fan et al. (2017)
34
ITS LSU rpb2 tef1
E. othonnae CBS 139910T KR476726 – MK540083 – Crous et al. (2015c), present study
E. perseae CBS 406.34T KX887258 KX887021 KX887139 KX886903 Fan et al. (2017)
T
E. phaseoli CBS 165.31 KX887263 KX887026 KX887144 KX886908 Fan et al. (2017)
E. picconiae CBS 145026T MK539951 MK540022 MK540081 MK540164 Present study
E. piri CBS 163.29 KX887267 KX887030 KX887148 KX886912 Fan et al. (2017)
E. pitangae CBS 227.50T KX887269 KX887032 KX887150 KX886914 Fan et al. (2017)
E. poinsettiae CBS 109333 KX887270 KX887033 KX887151 KX886915 Fan et al. (2017)
E. pongamiae CBS 402.63ET KX887272 KX887035 KX887153 KX886917 Fan et al. (2017)
E. populi CBS 289.64 KX887273 KX887036 KX887154 KX886918 Fan et al. (2017)
E. preissianae CBS 142129T KY173406 KY173498 – – Crous et al. (2016a)
E. proteae STE-U 1349 T
AF097578 – – – Swart et al. (2001)
E. protearum CBS 113618T KX887275 KX887038 KX887156 KX886920 Fan et al. (2017)
E. punicae CPC 19968 KX887276 KX887039 KX887157 KX886921 Fan et al. (2017)
E. quercus-ilicis CBS 232.61T KX887277 KX887040 – KX886922 Fan et al. (2017)
IsoT
E. randii CBS 170.38 KX887278 KX887041 KX887158 KX886923 Fan et al. (2017)
E. rhois CBS 519.50T KX887280 KX887043 KX887160 KX886925 Fan et al. (2017)
E. ricini CBS 403.63 KX887281 KX887044 KX887161 KX886926 Fan et al. (2017)
E. rosarum CBS 212.33ET KX887283 KX887046 KX887163 KX886928 Fan et al. (2017)
T
E. salicina CPC 17824 KX887286 KX887049 KX887166 KX886931 Fan et al. (2017)
E. semecarpi CBS 477.62T KX887287 KX887050 KX887167 KX886932 Fan et al. (2017)
E. sesseae CPC 18549 KX887288 KX887051 KX887168 KX886933 Fan et al. (2017)
E. sicula CBS 398.59T KX887289 KX887052 KX887169 KX886934 Fan et al. (2017)
E. solidaginis CBS 191.37ET KX887290 KX887053 KX887170 KX886935 Fan et al. (2017)
T
E. tectificae CBS 124777 KX887292 KX887055 KX887172 KX886937 Fan et al. (2017)
E. terminaliae CBS 343.39ET KX887293 KX887056 KX887173 – Fan et al. (2017)
T
E. theae CBS 228.50 KX887295 KX887058 KX887175 KX886939 Fan et al. (2017)
E. tiliae CBS 350.73 KX887296 KX887059 KX887176 KX886940 Fan et al. (2017)
E. veneta CBS 164.29ET KX887297 KX887060 KX887177 KX886941 Fan et al. (2017)
E. verbenae CPC 18561ET KX887298 KX887061 KX887178 KX886942 Fan et al. (2017)
E. veronicae CBS 145362 T
MK539952 MK540023 MK540082 – Present study
E. violae CBS 336.35T KX887302 KX887065 KX887182 KX886946 Fan et al. (2017)
E. zizyphi CBS 378.62T KX887303 KX887066 KX887183 KX886947 Fan et al. (2017)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute; DAR:
Plant Pathology Herbarium, New South Wales, Australia; STE-U: Department of Plant Pathology, Stellenbosch University, South Africa. T, ET and IsoT indicate ex-type, ex-
epitype and ex-isotype strains, respectively.
2
subunit gene; tef1: partial translation elongation factor 1-alpha gene.
94.44 % with the epitype strain of B. jonkershoekensis. Brun- Plectodiscella Woron., Mykol. Zentbl. 4: 232. 1914.
neosphaerella roupeliae produces the shortest ascospores of the Isotexis Syd., in Sydow & Petrak, Annls mycol. 29: 261. 1931.
genus [(19–)22–23(–25) μm in B. roupeliae vs. (25–) Melanobasis Clem. & Shear, Gen. fung., Edn 2 (Minneapolis):
27–34(–37) μm in B. jonkershoekensis vs. (20–) 224. 1931.
24–28(–30) μm in B. nitidae vs. (20–)23–26(–30) μm in Melanodochium Syd., Annls mycol. 36: 310. 1938.
B. protearum]. Bitancourtia Thirum. & Jenkins, Mycologia 45: 781. 1953.
Kurosawaia Hara, List of Japanese Fungi, 4th Edn: 172.
Authors: P.W. Crous, J.Z. Groenewald & Y. Marin-Felix 1954.
Elsinoe Racib., Parasit. Alg. Pilze Java's (Jakarta) 1: 14. 1900. Uleomycina Petr., Sydowia 8: 74. 1954.
Figs 20, 21. Melanophora Arx, Verh. K. ned. Akad. Wet., tweede sect. 51: 43.
Synonyms: Sphaceloma de Bary, Ann. Oenol. 4: 165. 1874. 1957.
Manginia Viala & Pacottet, C. r. hebd. Seanc. Acad. Sci., Paris
139: 88. 1904. Classification: Dothideomycetes, Dothideomycetidae, Myr-
Melanobasidium Maubl., Bull. Soc. mycol. Fr. 22: 69. 1906. iangiales, Elsinoaceae.
MARIN-FELIX ET AL.
Fig. 22. RAxML phylogram obtained from the combined ITS (609 bp), LSU (741 bp), rpb2 (747 bp) and tef1 (422 bp) sequence alignment of all accepted species of Elsinoe.
The tree was rooted to Myriangium hispanicum CBS 247.33. The novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS) values above 70 % are
shown at the nodes. GenBank accession numbers are listed in Table 4. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype strains, respectively. TreeBASE: S23834.
36
Type species: Elsinoe canavaliae Racib. Type or reference brown, soft, multi-loculate, locules scattered in upper part of
material not available. ascostromata; ascostromatal wall composed of pseudoparen-
DNA barcode (genus): LSU. chymatous cells of textura globulosa to textura angularis; locules
with few to numerous asci inside each locule, ostiolate; ostiole
DNA barcodes (species): ITS, rpb2, tef1. Table 4. Fig. 22. minute. Periphyses absent. Asci 8-spored, bitunicate, fissituni-
Ascostromata solitary, aggregated, or gregarious, wart-like, or as cate, saccate to globose, with a minute pedicel, and ocular
small distinctively coloured elevations, or pulvinate, immersed to chamber. Ascospores irregularly arranged, oblong or fusiform
semi-immersed, globose to subglobose, white, pale yellow or with slightly acutely rounded ends, with 2–3 transverse septa,
Fig. 22. (Continued).
MARIN-FELIX ET AL.
Fig. 23. Elsinoe picconiae (ex-type CBS 145026). A. Colony on OA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.
hyaline, smooth-walled, lacking a sheath. Acervuli or spor- cork-like appearance of older infected tissues with scab-like
odochia subepidermal, pseudoparenchymatous. Conidiophores appearance. Also, these can produce other disease symptoms
hyaline to pale brown, polyphialidic. Conidiogenous cells formed often called anthracnose, as in the case of infected grapevines
directly from the upper cells of the pseudoparenchyma, mono- to (Barrus & Horsfall 1928, Jenkins 1947, Farr et al. 1989, Pan
polyphialidic, integrated or discrete, determinate, hyaline to pale 1994, Phillips 1994, Gottwald 1995). However, the use of this
brown, without visible periclinal thickening. Conidia hyaline, name is confusing since it is used much broader to include
smooth-walled, aseptate, ellipsoidal, guttulate (adapted from Fan diseases caused by Colletotrichum. Although many species of
et al. 2017). Elsinoe causing scab disease have been described, only few of
them cause important diseases (Holliday 1980), having the main
Culture characteristics: Colonies on MEA, slow growing, raised,
impact on the appearance of the harvested product and its
irregular, erumpent, folded or cerebriform, smooth and irregular
market acceptability rather than on crop productivity (Swart et al.
margins, with sparse to moderate white to grey aerial mycelium.
2001). Species of Elsinoe seem to be host-specific since 77 of
On MEA, surface white to pale luteous, cinnamon, sepia, apricot,
the 81 species accepted in the present study occur on only one
saffron with or without purplish grey in centre, brown with apricot
host species or genus.
margins, rosy buff in centre with cinnamon margins, livid red,
Elsinoe and its asexual morph, Sphaceloma, were recently
scarlet red with diffuse red pigment in agar, or iron-grey; reverse
reviewed by Fan et al. (2017). In that study, 26 new combinations
umber, ochreous, iron-grey, dark vinaceous, or centre scarlet
were proposed for the species originally placed in Sphaceloma.
and orange with cinnamon margins.
Moreover, eight new species were introduced and 13 epitypes
Optimal media and cultivation conditions: MEA, OA, PDA, SNA were designated. Based on phylogenetic data, Fan et al. (2017)
and WA at 22 °C under near-ultraviolet light (12 h light, 12 h accepted 75 species in the genus. However, E. banksiae, which
dark). was described by Swart et al. (2001), and three species
described by Crous et al. (2016a), i.e. E. eelemani,
Distribution: Worldwide.
E. eucalyptigena and E. preissianae, were not included in that
Hosts: Wide range of hosts, including some economically study. Our phylogenetic analysis corroborated the placement of
important crops such as avocado, cassava, citrus, grapevines, these species in the main well-supported clade representing the
ornamentals such as poinsettias, field crops and woody hosts. genus Elsinoe, except for E. eucalyptigena, whose placement
remains unknown and, therefore it is not considered an accepted
Disease symptoms: Scab, leaf and fruit spot and anthracnose
species of Elsinoe in the present study. Moreover, another three
disease.
new species have been subsequently described by Crous et al.
Notes: Elsinoe comprises plant pathogenic species that cause (2018), i.e. E. banksiigena, E. elaeocarpi and E. leucopogonis.
scab and spot anthracnose on a wide range of hosts, including However, in our phylogenetic analysis, the two first species were
some economically important crops and ornamentals. The dis- not located in the Elsinoe s. str. clade, and are thus excluded
ease symptoms that these species produce are easily recog- from the genus at present. Therefore, hitherto a total of 79
nisable, being known as “signature-bearing diseases”, for the species are accepted, plus the new species described in the
Fig. 24. Elsinoe veronicae (ex-type CBS 145362). A. Colony on OA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.
38
present study. Unfortunately, there are no cultures and molecular Notes: Elsinoe veronicae is closely related to E. othonnae.
data of the type species of the genus, E. canavaliae, which Although the ITS sequences of both species showed more than
needs to be epitypified in order to clarify its phylogenetic position. 99 % nucleotide similarity, the rpb2 sequences showed only
96.95 % similarity. Morphologically, these are also similar
References: Fan et al. 2017 (morphology and phylogeny). differing mainly in their conidial size [(4–)
Elsinoe picconiae Crous, sp. nov. MycoBank MB829611. 5–5.5(–6.5) × 2.5(–3) μm in E. veronicae vs. (5–)6–7 × (2.5–)
Fig. 23. 3(–4) μm in E. othonnae]. Moreover, E. veronicae was found on
Veronica (Scrophulariaceae) in New Zealand, while E. othonnae
Etymology: Name refers to Picconia, the host from which this has only been reported on Othonna (Asteraceae) in South Africa
fungus was collected. (Crous et al. 2015c).
Conidiomata sporodochial or acervular on leaves, medium Authors: Y. Marin-Felix, R. Thangavel & P.W. Crous
brown, up to 250 μm diam; conidiomatal wall composed of tex-
Exserohilum K.J. Leonard & Suggs, Mycologia 66: 290. 1974.
tura angularis. Conidiophores subcylindrical to doliiform, hyaline
Fig. 25.
to pale brown, smooth-walled, 0–1-septate, unbranched,
Synonyms: Setosphaeria K.J. Leonard & Suggs, Mycologia 66:
10–15 × 3–4 μm. Conidiogenous cells polyphialidic, hyaline,
294. 1974.
smooth-walled, subcylindrical to doliiform, 5–8 × 3–4 μm.
Luttrellia Khokhr. & Gornostaĭ (as ‘Lutrellia’; non Luttrellia
Conidia hyaline, aseptate, ellipsoid, apex obtuse, base truncate,
Shearer), Vodorosli, Griby i Mkhi Dal’nego Vostoka [Algae, Fungi
(4–)5–6(–7) × (2–)2.5 μm.
and Mosses of the Soviet Far-East] (Vladivostok): 80. 1978.
Culture characteristics: Colonies erumpent, spreading, with
sparse aerial mycelium, folded surface, and smooth, lobate
sporales, Pleosporaceae.
margins, reaching 7 mm diam after 2 wk at 25 °C. On MEA
surface rust, reverse sienna; on PDA surface coral, reverse bay; Type species: Exserohilum turcicum (Pass.) K.J. Leonard &
on OA surface scarlet with diffuse scarlet pigment. Suggs, basionym: Helminthosporium turcicum Pass. Ex-epitype
and ex-epitype strain designated by Hernandez-Restrepo et al.
Typus: Spain, Tenerife, Los Silos, leaf of Picconia excelsa
(2018): CBS H-23323, CBS 690.71.
(Oleaceae), 12 Mar. 2017, A. van Iperen, HPC 2063 (holo-
type CBS H-23848, culture ex-type CBS 145026 = CPC DNA barcodes (genus): LSU, ITS.
33648).
DNA barcodes (species): ITS, gapdh, rpb2. Table 5. Fig. 26.
Notes: Elsinoe picconiae is related to E. freyliniae, E. oleae and
Ascomata superficial, immersed or erumpent, globose to ellip-
E. salicina. However, E. picconiae can be easily distinguished by
soid, unilocular, dark brown to black, with or without a neck,
its narrower conidia [(2–)2.5 μm vs. (2.5–)3–4 μm in
ostiolate, with simple rigid setae around the ostiolar apex and on
E. freyliniae, 3–6 μm in E. oleae, and (2.5–)3–4.5(–5) μm in
the upper half of the ascoma where they are often mixed with
E. salicina]. Moreover, E. picconiae is the first species of the
hyaline, filiform, septate hyphae; ascomatal wall composed of
genus isolated from Picconia excelsa.
pseudoparenchymatous cells, dark brown and thick-walled on
Elsinoe veronicae Crous, Thangavel & Y. Marín, sp. nov. the outside, but with more or less hyaline cells towards the in-
MycoBank MB829610. Fig. 24. side, cells of textura angularis. Pseudoparaphyses filiform, hy-
aline, septate, branched, anastomosing. Asci arising from a
Etymology: Name refers to Veronica, the host from which this basal cushion of thin-walled pseudoparenchymatous cells, bitu-
fungus was collected. nicate, 1–8-spored, cylindrical to cylindrical-clavate, short or
Conidiomata sporodochial, erumpent, 80–200 μm diam, based moderately long-stalked, thick-walled, with an apical nasse and
on a pale brown stroma, giving rise to densely aggregated co- fissitunicate dehiscence. Ascospores fusoid, hyaline to pale
nidiophores. Conidiophores hyaline to pale brown, smooth- brown, smooth-walled, 2–6 or rarely more transversely septate,
walled, subcylindrical, 1–2-septate, 15–30 × 2.5–3.5 μm, un- constricted at the septa, surrounded by a hyaline mucilaginous
branched or branched above. Conidiogenous cells integrated, sheath which often extends some distance beyond the ends of
subcylindrical, hyaline, smooth-walled, 7–10 × 2.5–3.5 μm, the spore. Conidiophores macronematous, mononematous,
polyphialidic. Conidia solitary, aggregating in mucoid mass, septate, cylindrical, olivaceous brown to brown, smooth-walled to
aseptate, hyaline, smooth-walled, guttulate, ellipsoid, apex sub- verruculose, often geniculate above. Conidiogenous cells inte-
obtuse, tapering at base to truncate hilum, (4–) grated, terminal and intercalary, sympodial, mono- or polytretic,
5–5.5(–6.5) × 2.5(–3) μm. cicatrised; conidiogenous nodes smooth to rough. Conidia fusi-
form, cylindrical or obclavate, straight to curved, multi-
Culture characteristics: Colonies erumpent, spreading, surface distoseptate, with a protruding hilum (adapted from
folded with moderate aerial mycelium, and even, lobate margins, Hernandez-Restrepo et al. 2018).
reaching 25 mm diam after 2 wk. On MEA surface brick, reverse
cinnamon; on PDA surface brick to scarlet, reverse brick in Culture characteristics: Colonies on PDA brown or grey oliva-
centre, scarlet in outer region; on OA surface scarlet with diffuse ceous to olivaceous black, sometimes white, pale grey, hairy,
scarlet pigment. cottony to powdery, margins fimbriate.
Optimal media and cultivation conditions: Sterilised Zea mays
Typus: New Zealand, Auckland, St. John, Morrin Road, on
leaves placed on 1.5 % WA or PDA at 25 °C under near-
Veronica sp. (Scrophulariaceae), 2013, R. Thangavel (holotype
ultraviolet light (12 h light, 12 h dark) to induce sporulation.
CBS H-23865, culture ex-type CBS 145362 = CPC
34137 = T17_00408D). Distribution: Worldwide.
MARIN-FELIX ET AL.
Fig. 25. Exserohilum spp. A–E. Sexual morph. A. Ascomata of Exserohilum minor (ex-isotype IMI 294530). B, C. Asci of Exserohilum minor (ex-isotype IMI 294530). D, E.
Ascospores. D. Exserohilum minor (ex-isotype IMI 294530). E. Exserohilum khartoumensis (ex-isotype CBS 132708). F–AA. Asexual morph. F–N. Conidiophores and conidia.
F, L. Exserohilum oryzicola (ex-isotype CBS 502.90). G. Exserohilum turcicum (ex-epitype CBS 690.71). H, N. Exserohilum holmii (ex-isotype CBS 413.65 and BRIP 12679).
40
Table 5. DNA barcodes of accepted Exserohilum spp.

ITS LSU gapdh rpb2
Exserohilum corniculatum BRIP 11426T LT837453 LT883391 LT883533 LT852480 Hernandez-Restrepo et al. (2018)
Ex. holmii CBS 413.65IsoT LT837459 LT715621 LT715890 LT852489 Hernandez-Restrepo et al. (2018)
Ex. khartoumensis IMI 249194 IsoT
LT837461 LT715619 LT715888 LT852490 Hernandez-Restrepo et al. (2018)
Ex. minor BRIP 14616T LT837470 LT883403 LT883545 LT852502 Hernandez-Restrepo et al. (2018)
Ex. monoceras BRIP 12271 A
Ex. neoregeliae CBS 132832 T
Ex. oryzicola CBS 502.90 IsoT
HF934949 HF934886 LT715878 HF934851 Amaradasa et al. (2014), Hernandez-Restrepo
et al. (2018)
Ex. pedicellatum CBS 322.64ET KT265258 HF934889 LT715902 HF934854 Amaradasa et al. (2014), Chowdhary et al.
(2015), Hernandez-Restrepo et al. (2018)
Ex. protrudens BRIP 14814T LT631308 LT715611 LT715880 LT715741 Hernandez-Restrepo et al. (2018)
Ex. rostratum CBS 325.87 KT265237 LT715629 LT715898 LT852492 Chowdhary et al. (2015), Hernandez-Restrepo
et al. (2018)
Ex. turcicum CBS 690.71ET LT837487 LT883415 LT882581 – Hernandez-Restrepo et al. (2018)
1
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; IMI: International Mycological
Institute, CABI-Bioscience, Egham, Bakeham Lane, UK. T, ET, IsoT and A indicate ex-type, ex-epitype, ex-isotype and authentic strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase
gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene.
Host: Mainly pathogens of grasses, but some also on non-grass germination failure in cereals (Drechsler 1923, Leonard 1976,
hosts. Other substrates where they can be found include river Sivanesan 1987, Lin et al. 2011).
sediments, soil, grains, plant debris, and humans. Exserohilum Previously, three different species, Ex. longirostratum, Ex.
rostratum has been reported as a human pathogen. macginnisii and Ex. rostratum, were recognised as human
pathogens (McGinnis et al. 1986, Padhye et al. 1986, de Hoog
Disease symptoms: In plants: leaf blight, leaf spots, melting out,
et al. 2000, da Cunha et al. 2012). However, a multi-locus
root rot, and foot rot, among others. In humans: skin infections,
phylogenetic analysis (Hernandez-Restrepo et al. 2018)
keratitis, non-invasive allergies, invasive sinusitis, and dissemi-
demonstrated that they are actually the same phylogenetic
nated infections.
species. Exserohilum rostratum has been reported as an agent
Notes: Exserohilum is differentiated from the closely related of phaeohyphomycosis and sometimes causing life-threatening
Bipolaris, Curvularia and Pyrenophora by producing conidia with infections in humans (McGinnis et al. 1986, Padhye et al.
a protruding hilum (Leonard & Suggs 1974). Recently the tax- 1986, Aquino et al. 1995, Adler et al. 2006). This species was
onomy and phylogeny of Exserohilum has been revisited by recently implicated in an outbreak of fungal meningitis associated
Hernandez-Restrepo et al. (2018). Based on morphological and with contaminated methylprednisolone in the USA (Kainer et al.
molecular data 11 phylogenetic species are accepted (Fig. 26, 2012).
Table 5). Three species were excluded from the genus, namely
Ex. novae-zelandiae relocated to Sporidesmiella, and Ex. paspali References: Drechsler 1923, 1934, Luttrell 1963, Leonard &
and Ex. sorghicola to Curvularia, while another 15 species were Suggs 1974, Sivanesan 1987 (taxonomy, morphology and
retained in Exserohilum, although some were doubtful. Species pathogenicity), Leonard 1976 (sexual/asexual connection), de
in Exserohilum are morphologically very variable and a molec- Hoog et al. 2000, da Cunha et al. 2012 (human pathogens),
ular analysis is required for a correct species identification. Hernandez-Restrepo et al. 2018 (morphology, phylogeny,
The type species of the genus, Ex. turcicum (= Helmintho- review).
sporium turcicum), was described from Italy causing northern
Author: M. Hernandez-Restrepo
leaf blight of corn (Passerini 1876, Saccardo 1886). Other spe-
cies attacking economically significant crops include Ex. ped- Neosetophoma Gruyter et al., Mycologia 102: 1075. 2010.
icellatum, causing root rot on maize and brown lesions on wheat Fig. 27.
roots (Henry 1924, Sivanesan 1987), and Ex. rostratum, pro-
ducing leaf spot on banana, maize and wheat, foot rot in wheat, Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
damping off of sugarcane seedlings, blackening and seed sporales, Phaeosphaeriaceae.
I. Exserohilum pedicellatum (CBS 375.76). J, M. Exserohilum rostratum (CBS 120380, CBS 196.29). K. Exserohilum monoceras (CBS 198.29). O. Detail of the conidial hilum of
Exserohilum oryzicola (ex-isotype CBS 502.90). P–AA. Conidia. P, Q, Z, AA. Exserohilum rostratum (CBS 128054, CBS 120380, BRIP 11422). R. Exserohilum holmii (BRIP
12679). S. Exserohilum pedicellatum (BRIP 12040). T, U. Exserohilum turcicum (BPI 431157 holotype). V. Exserohilum oryzicola (BRIP 16229). W. Exserohilum protrudens
(BRIP 14816). X. Exserohilum corniculatum (ex-type BRIP 11426). Y. Exserohilum neoregelia (CBS 132833). Scale bars A = 50 μm; others = 10 μm; C applies to B and C; E
applies to D and E. Pictures taken from Hernandez-Restrepo et al. (2018).
MARIN-FELIX ET AL.
Fig. 26. RAxML phylogram obtained from the combined ITS (793 bp), gapdh (578 bp) and rpb2 (860 bp) sequence alignment of all the accepted species of Exserohilum. The
tree was rooted to Curvularia and Bipolaris. RAxML bootstrap support (BS) values above 70 % are shown in the nodes. GenBank accession numbers are indicated in Table 5. T,
ET, IsoT
and A indicate ex-type, ex-epitype, ex-isotype and authentic strains. TreeBASE: S23834.
Type species: Neosetophoma samarorum (Desm.) Gruyter et al., DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 6. Fig. 29.
basionym: Phoma samarorum Desm. Epitype and ex-epitype
Ascomata ostiolate, globose to subglobose, solitary to gregar-
strain designated by Gruyter et al. (2010): CBS H-20319, CBS
ious, dark brown to black, immersed to slightly erumpent or
138.96.
superficial, smooth; ascomatal wall composed of 2–4 layers of
DNA barcode (genus): LSU. Fig. 28. brown to reddish-brown or dark brown to black cells of textura
Fig. 27. Neosetophoma lunariae (ex-type CBS 141409). A. Conidiomata on OA. B. Conidiomata showing ostiolar region. C, D. Conidiogenous cells. E. Conidia. Scale
bars = 10 μm; C applies to C and D. Pictures taken from Hernandez-Restrepo et al. (2016a).
42
Fig. 28. RAxML phylogram obtained from the combined ITS (626 bp) and LSU (852 bp) sequence alignment of members of the family Phaeosphaeriaceae. The tree was
rooted to Coniothyrium glycines CBS 124141 and Coniothyrium sidae CBS 135108. The novelties proposed in this study are indicated in bold. RAxML bootstrap support (BS)
values above 70 %and Bayesian posterior probability scores above 0.95 are shown at the nodes. Numbers between parentheses correspond to GenBank accession numbers
for ITS and LSU sequences, respectively. T, ET, HT, IsoT, LT and NT indicate ex-type, ex-epitype, holotype, ex-isotype, ex-lectotype and ex-neotype strains, respectively. TreeBASE:
S23834.
MARIN-FELIX ET AL.
44
Table 6. DNA barcodes of accepted Neosetophoma spp.

ITS rpb2 tef1 tub2
Neosetophoma aseptata CBS 145363T MK539953 MK540084 – – Present study
Nph. clematidis MFLUCC 13-0734T KP744450 – – – Liu et al. (2015)
Nph. garethjonesii MFLUCC 14-0528T KY496758 – KY514402a – Tibpromma et al. (2017)
Nph. guiyangensis GZCC 18-0111 T
MH018134 – – – Hyde et al. (2018)
Nph. iranianum IBRC-M 30176 T
MF684861 – – – Karunarathna et al. (2017)
Nph. italica MFLUCC 13-0388T KP711356 – – – Liu et al. (2015)
Nph. lunariae CBS 141409 T
KX306763 – – – Hernandez-Restrepo et al. (2016a)
Nph. phragmitis CBS 145364 T
MK539954 MK540085 MK540148 b
– Present study
Nph. poaceicola MFLUCC 16-0886 T
KY568986 – – – Thambugala et al. (2017)
Nph. rosae MFLUCC 15-1073T MG828925 – MG829218a – Wanasinghe et al. (2018)
Nph. rosarum MFLUCC 17-0308T MG828927 – – – Wanasinghe et al. (2018)
Nph. rosigena MFLUCC 17-0768T MG828928 – – – Wanasinghe et al. (2018)
ET b
Nph. samarorum CBS 138.96 KF251160 KF252168 KF253119 KF252655 Quaedvlieg et al. (2013)
Nph. sambuci CBS 145365T MK539955 MK540086 MK540149b – Present study
Nph. shoemakeri MFLUCC 17-0780 MG844346 – MG844352a – Hyde et al. (2018)
Nph. xingrensis GZCC 18-0110 T
MH018135 – – – Hyde et al. (2018)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; GZCC: Guizhou Academy of Agricultural Sciences Culture Collection, Guiyang, China; IBRC:
Herbarium of the Plant bank, Iranian Biological Resource Center, Karaj, Iran; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand. T and ET
indicate ex-type and ex-epitype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; tef1: partial translation
elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used for sequencing: a: EF1-983F / EF1-2218R; b: EF1-728F / EF-2.
angularis to textura prismatica. Hamathecium comprising grey olivaceous, or dark brown. On MEA pale grey to almost
numerous, septate, cellular or filamentous pseudoparaphyses, white, buff, or brown to dark brown; reverse dark brown, or buff
embedded in a hyaline gelatinous matrix. Asci 8-spored, bitu- with patches of grey olivaceous.
nicate, fissitunicate, cylindrical to clavate, short pedicellate, with
Optimal media and cultivation conditions: CMA, PDA, PNA and
a furcate pedicel, apically rounded, with a minute or indistinct
OA at 25 °C under continuous near-ultraviolet light to promote
ocular chamber. Ascospores overlapping, 1–3-seriate, hyaline
sporulation.
or subhyaline when young, becoming pale yellow, pale brown or
yellowish brown to brown at maturity, 1–5-septate, straight to Distribution: Asia, Europe and North America.
slightly curved, fusoid or narrowly fusoid, with rounded or acute
Hosts: Pathogens or saprophytes found on a wide range of
ends, constricted or not at the septum, enlarged at the second
host including Cirsium arvense (Asteraceae), Clematis vitalba
cell below apex, guttulate, smooth-walled, without any muci-
(Ranunculaceae), Iris germanica (Iridaceae), Lunaria annuna
laginous sheath and appendages. Conidiomata pycnidial, soli-
(Brassicaceae), Malva sp. (Malvaceae), Phlox paniculata
tary to confluent, immersed or superficial, globose to
(Polemoniaceae), Phragmites australis (Poaceae), Rosa can-
subglobose or irregular, with mycelial outgrowths, or confluent,
ina (Rosaceae), Sambucus spp., Viburnum opulus (Caprifo-
unilocular, occasionally multi-locular, with papillate ostioles,
liaceae), and Urtica dioica (Urticaceae). Also isolated from
sometimes developing long necks, honey, olivaceous, oliva-
soil.
ceous black, pale brown, brown, dark brown, or black, with up to
10 layers of pseudoparenchymatal cells of textura angularis. Disease symptoms: Leaf spots.
Conidiophores reduced to conidiogenous cells. Conidiogenous
Notes: Neosetophoma was introduced by de Gruyter et al.
cells phialidic, extending percurrently at apex, hyaline, doliiform
(2010) to accommodate Phoma samarorum, which is a path-
to ampulliform, determinate, hyaline, smooth-walled. Conidia
ogen causing leaf spots of grasses. Subsequently, 12 new
hyaline, slightly yellowish or pale brown, 0–3(–4)-septate,
species have been added to this genus, all of which appear to be
ellipsoidal, cylindrical, subcylindrical, fusiform, or tear-drop
saprobes, except for Nph. iranianum, which was isolated from
shape, straight to curved, usually attenuate at one end, or
soil (Karunarathna et al. 2017), and Nph. lunariae, which is
apex and base obtuse, or sometimes with bluntly rounded to
endophytic (Hernandez-Restrepo et al. 2016a).
truncate base, continuous or constricted at the septa, smooth-
This genus is characterised by globose to irregular con-
walled, often guttulate (asexual morph description adapted
idiomata with papillate ostioles, and yellowish to brownish
from Gruyter et al. 2010).
conidia usually attenuated at one end, less frequent with
Culture characteristics: Colonies flat, with a moderate amount of apex and base obtuse or with a bluntly rounded to truncate
aerial mycelium. On PDA surface fluffy, circular or irregular, base. The sexual morph was observed for the first time by
margins entire or filiform, white, pale grey, grey, greenish grey, or Tibpromma et al. (2017), when Nph. garethjonesii was
mouse grey; reverse yellowish, yellowish grey, greyish white, introduced. Subsequently, four new species producing a
MARIN-FELIX ET AL.
sexual morph have been described, of which only one also Neosetophoma aseptata Crous, R.K. Schumach. & Y. Marín,
produces the asexual morph, namely Nph. shoemakeri (Hyde sp. nov. MycoBank MB829639. Fig. 30.
et al. 2018). Etymology: Name refers to its aseptate conidia, which have
References: de Gruyter et al. 2010, Quaedvlieg et al. 2013, never been observed in the other species of the genus.
Tibpromma et al. 2017, Hyde et al. 2018, Wanasinghe et al. Conidiomata solitary, brown, erumpent, globose, 250–350 μm
2018 (morphology and phylogeny). diam with 1–3 ostioles; conidiomatal wall of 3–4 layers of brown
Fig. 29. RAxML phylogram obtained from the combined ITS (585 bp), LSU (848 bp) and rpb2 (744 bp) sequence alignment of all accepted species of Neosetophoma. The tree
was rooted to Phaeosphaeriopsis glaucopunctata MFLUCC 13-0265 and Phaeosphaeriopsis agavacearum CPC 29122. The novelties proposed in this study are indicated in
bold. RAxML bootstrap support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession numbers of LSU
are listed in Fig. 28, and of the other loci in Tables 6 and 10. T and ET indicate ex-type and ex-epitype strains, respectively. TreeBASE: S23834.
46
Fig. 30. Neosetophoma aseptata (ex-type CBS 145363). A. Conidiomata sporulating on OA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 300 μm; others = 10 μm.
cells of textura angularis. Conidiophores reduced to con- clematidis, Nph. iranianum, Nph. lunariae, Nph. rosae and Nph.
idiogenous cells lining inner cavity, hyaline, smooth-walled, shoemakeri. Neosetophoma aseptata can be easily distin-
subcylindrical to ellipsoid, phialidic with minute collarette, guished by its aseptate conidia, being 1-septate in Nph. shoe-
4–8 × 4–5 μm. Conidia solitary, aseptate, hyaline, smooth- makeri, 1–3-septate in Nph. iranianum and Nph. rosae, 3-
walled, subcylindrical to ellipsoid, apex obtuse, base truncate, septate in Nph. clematidis, and (1–)3(–4)-septate in Nph.
(3.5–)4–5 × (1.5–)2 μm. lunariae. Moreover, Nph. aseptata produces the smallest con-
Culture characteristics: Colonies flat, spreading, surface folded, idia in the complex [(3.5–)4–5 × (1.5–)2 μm in Nph. aseptata
with moderate aerial mycelium and even, lobate margins, vs. 4–6 × 2–4 μm in Nph. iranianum vs. 7.5–10.5 × 2.5–3 μm
reaching 50 mm diam after 2 wk. On MEA surface pale oliva- in Nph. shoemakeri vs. 8–14 × 1.5–3 μm in Nph. rosae vs.
ceous grey, reverse umber with diffuse umber pigment; on PDA 11–15 × 2–4 μm in Nph. clematidis vs. (10–)
surface olivaceous grey, reverse umber with diffuse umber 14–17(–22) × (2.5–)3 μm in Nph. lunariae]. Neosetophoma
pigment; on OA surface saffron with patches of grey olivaceous. clematidis produces the largest conidiomata in this complex,
being up to 475 μm diam (up to 300 μm in Nph. aseptata and
Typus: Germany, near Berlin, moist meadow, on Viburnum Nph. lunaria, up to 180 μm in Nph. shoemakeri, up to 130 μm in
opulus (Caprifoliaceae), 7 Jun. 2017, R.K. Schumacher, HPC Nph. rosae, up to 120 μm in Nph. iranianum).
2131, RKS 123 (holotype CBS H-23866, culture ex-type CBS
Neosetophoma aseptata is the first species isolated from
145363 = CPC 33919).
Viburnum. Neosetophoma samarorum and Nph. sambuci are
Notes: Neosetophoma aseptata was located in a large, well- reported in Sambucus spp., which is a member of the same
supported clade (96 % BS / 1 PP) comprising Nph. family, Caprifoliaceae.
Fig. 31. Neosetophoma phragmitis (ex-type CBS 145364). A. Conidiomata sporulating on SNA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 200 μm; others = 10 μm.
Fig. 32. Neosetophoma sambuci (ex-type CBS 145365). A. Conidiomata sporulating on PNA. B. Conidioma on SNA showing papillate neck. C. Conidiogenous cells. D.
Conidia. Scale bars: A, B = 200 μm; others = 10 μm.
MARIN-FELIX ET AL.
Fig. 33. Neostagonospora spp. A, B. Conidioma forming in culture. A. Neostagonospora caricis (ex-type CBS 135092). B. Neostagonospora elegiae (ex-type CBS 135101).
C–G. Conidiogenous cells. C, D. Neostagonospora caricis (ex-type CBS 135092). E–G. Neostagonospora elegiae (ex-type CBS 135101). H, I. Conidia. H. Neostagonospora
caricis (ex-type CBS 135092). I. Neostagonospora elegiae (ex-type CBS 135101). Scale bars: B = 150 μm; all others = 10 μm; C applies to C and D. Pictures taken from
Quaedvlieg et al. (2013).
Neosetophoma phragmitis Crous, R.K. Schumach. & Y. Marín, layers of brown cells of textura angularis. Conidiophores reduced
sp. nov. MycoBank MB829640. Fig. 31. to conidiogenous cells lining inner cavity, hyaline, smooth-walled,
ampulliform, 5–8 × 2.5–3 μm, proliferating percurrently at apex.
Etymology: Name refers to Phragmites, the host from which this Conidia solitary, pale brown, smooth-walled, aseptate, straight,
fungus was collected. apex obtuse, base truncate, (3–)4–5(–6) × 2 μm.
Conidiomata solitary, pycnidial, brown, globose, 180–200 μm Culture characteristics: Colonies flat, spreading, with moderate
diam, neck papillate with central ostiole; conidiomatal wall of 3–4 aerial mycelium and feathery margins, covering dish in 2 wk. On
Table 7. DNA barcodes of accepted Neostagonospora spp.

ITS rpb2 tef1 tub2
Neostagonospora arrhenatheri MFLUCC 15-0464T KX926417 – MG520901a – Phookamsak et al. (2017), Thambugala et al. (2017)
Nst. caricis CBS 135092 T
KF251163 KF252171 – KF252658 Quaedvlieg et al. (2013)
T b
Nst. elegiae CBS 135101 KF251164 KF252172 KF253122 KF252659 Quaedvlieg et al. (2013)
Nst. phragmitis MFLUCC 16-0493T KX926416 – MG520902a – Phookamsak et al. (2017), Thambugala et al. (2017)
Nst. sorghi CBS 145366T MK539956 MK540087 MK540150b MK540168 Present study
Nst. spinificis BCRC FU30120 KP676045 LC055104 – – Yang et al. (2016)
1
BCRC: Bioresource Collection and Research Centre, Taiwan; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLUCC: Mae Fah Luang
University Culture Collection, Chiang Ria, Thailand. T indicates ex-type strains.
2
elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used for sequencing: a: EF1-983F / EF1-2218R, b: EF1-728F / EF-2.
48
MEA surface and reverse ochreous; on PDA surface and reverse Culture characteristics: Colonies flat, spreading, covering dish in
hazel; on OA surface hazel. 2 wk, with sparse to moderate aerial mycelium. On MEA surface
ochreous, reverse umber; on PDA surface and reverse oliva-
Typus: Germany, near Berlin, on leaf sheath of Phragmites
ceous grey; on OA surface ochreous.
australis (Poaceae), 16 Apr. 2016, R.K. Schumacher, HPC 1178
(holotype CBS H-23867, culture ex-type CBS 145364 = CPC Typus: Germany, near Berlin, on twig of Sambucus nigra
30680). (Caprifoliaceae), 11 Mar. 2016, R.K. Schumacher, HPC 1072
(holotype CBS H-23868, culture ex-type CBS 145365 = CPC
Notes: In the phylogenetic analysis based on ITS, LSU and
30357).
rpb2 sequences, Nph. phragmitis was located in an indepen-
dent branch removed from the other species of the genus. Notes: In the phylogenetic analysis based on ITS, LSU and rpb2
This is the first species isolated from Phragmitis australis sequences, Nph. sambuci was located in a well-supported clade
(Poaceae). Neosetophoma poaceicola is the only species that (97 % BS / 0.98 PP) together with Nph. garethjonesii, Nph.
was reported before on a member of the Poaceae, being samarorum and Nph. rosigena. Neosetophoma garethjonesii can
isolated from a grass host. However, both species are not be easily distinguished from the other species by only producing a
related, and Nph. phragmitis only produces the asexual sexual morph. The other three species produce an asexual
morph, while only the sexual morph was observed in Nph. morph, and can be differentiated by the size of their conidia
poaceicola. [4–16 × 1.5–3 μm in Nph. samarorum vs. (5–)7–8(–10) × (2–)
2.5(–3) μm in Nph. sambuci vs. 4–6 × 1.5–2.5 μm in Nph.
Neosetophoma sambuci Crous, R.K. Schumach. & Y. Marín,
rosigena]. Moreover, conidia in Nph. sambuci are pale brown
sp. nov. MycoBank MB829641. Fig. 32.
while in Nph. samarorum they are slightly yellowish and in Nph.
Etymology: Name refers to the genus Sambucus, the host from rosigena they are olivaceous brown. Neosetophoma sambuci was
which this fungus was collected. isolated from twigs of Sambucus nigra (Caprifoliaceae). The only
species previously reported on this host was Nph. samarorum.
Conidiomata solitary, erumpent, brown, pycnidial, globose, Authors: Y. Marin-Felix & P.W. Crous
150– 200 μm diam, with central ostiole; conidiomatal wall of
3– 4 layers of brown textura angularis. Conidiophores Neostagonospora Quaedvl. et al., Stud. Mycol. 75: 364. 2013.
reduced to conidiogenous cells lining the inner cavity, hyaline, Fig. 33.
smooth-walled, ampulliform, phialidic, 4 –6 × 3–4 μm. Conidia Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
solitary, pale brown, smooth-walled, guttulate, subcylindrical,
sporales, Phaeosphaeriaceae.
apex obtuse, base truncate, aseptate, becoming 1-septate
and swollen (ellipsoid) with age, (5–)7– 8(– 10) × (2 –) Type species: Neostagonospora caricis Quaedvlieg et al. Holo-
2.5( –3) μm. type and ex-type strain: CBS H-21306, CBS 135092.
Fig. 34. RAxML phylogram obtained from the combined ITS (571 bp), LSU (847 bp), rpb2 (337 bp) and tub2 (304 bp) sequence alignment of all accepted species of
Neostagonospora. The tree was rooted to Parastagonospora avenae CBS 289.69 and Parastagonospora nodorum CBS 110109. The novelty proposed in this study is indicated
in bold. RAxML bootstrap support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession numbers of LSU
are listed in Fig. 28, and of the other loci in Tables 7 and 9. T indicates ex-type strains. TreeBASE: S23834.
MARIN-FELIX ET AL.
DNA barcode (genus): LSU. Fig. 28. Conidiomata immersed, pycnidial, globose, exuding a pale
luteous to creamy conidial mass; conidiomatal wall composed of
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 7. 2–3 layers of pale brown cells of textura angularis. Co-
Fig. 34. nidiophores reduced to conidiogenous cells. Conidiogenous cells
Fig. 35. Neostagonospora sorghi (ex-type CBS 145366). A. Conidiomata sporulating on MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 200 μm; others = 10 μm.
Fig. 36. Nothophoma spp. A, B. Disease symptoms. A. Nothophoma quercina (CGMCC 3.19246) on Osmanthus fragrans. B. Nothophoma quercina (LC12187) on Jasminum
mesnyi. C–I. Asexual morph. C, D. Conidiomata of Nothophoma anigozanthi (ex-epitype CBS 381.91) sporulating on OA. E–H. Conidia. E. Nothophoma infossa (ex-neotype
CBS 123395). F. Nothophoma macrospora (ex-type CBS 140674). G. Nothophoma quercina (CGMCC 3.19246). H. Nothophoma variabilis (ex-type CBS 142457). I. Con-
idiogenous cells of Nothophoma macrospora (ex-type CBS 140674). Scale bars: C = 200 μm; D = 20 μm; E–I = 10 μm. Pictures C, D taken from Chen et al. (2015); E from
Aveskamp et al. (2009); F, I from Crous et al. (2016b); H from Valenzuela-Lopez et al. (2018).
50
Table 8. DNA barcodes of accepted Nothophoma spp.

LSU ITS rpb2 tub2
Nothophoma anigozanthi CBS 381.91ET GU238039 GU237852 KT389655 GU237580 Aveskamp et al. (2010), Chen et al. (2015)
Not. arachidis-hypogaeae CBS 125.93 GU238043 GU237771 KT389656 GU237583 Aveskamp et al. (2010), Chen et al. (2015)
Not. gossypiicola CBS 377.67 GU238079 GU237845 KT389658 GU237611 Aveskamp et al. (2010), Chen et al. (2015)
Not. infossa CBS 123395NT GU238089 FJ427025 KT389659 FJ427135 Aveskamp et al. (2009, 2010), Chen et al. (2015)
Not. macrospora CBS 140674T LN880537 LN880536 LT593073 LN880539 Crous et al. (2016b)
Not. quercina CBS 633.92 EU754127 GU237900 KT389657 GU237609 Aveskamp et al. (2010), Chen et al. (2015)
CGMCC 3.19246 MK088581 MK088574 MK088588 MK088595 Present study
Not. raii MCC 1082 T
– MF664467 – MF664468 Crous et al. (2017b)
Not. variabilis CBS 142457T LN907428 LT592939 LT593078 LT593008 Valenzuela-Lopez et al. (2018)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbiological Culture Collection Center, Beijing, China; LC: Dr Lei
Cai's personal culture collection, housed at CAS, China; MCC: National Centre for Microbial Resources (formerly Microbial Culture Collection), Pune, India. T, ET and NT
indicate ex-type, ex-epitype and ex-neotype strains, respectively.
2
LSU: partial large subunit (28S) nrRNA gene; ITS: internal transcribed spacers and intervening 5.8S nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest
subunit gene; tub2: partial β-tubulin gene.
phialidic, hyaline, smooth-walled, aggregated, lining the inner ellipsoidal to subcylindrical conidia, but Neostagonospora is
cavity, ampulliform to doliiform, tapering at apex with prominent distinguished by having conidiogenous cells that are phialidic,
periclinal thickening. Conidia hyaline, smooth-walled, granular, with prominent periclinal thickening (Quaedvlieg et al. 2013).
thin-walled, narrowly fusoid-ellipsoidal to subcylindrical, apex In our phylogenetic analysis based on ITS and LSU se-
subobtusely rounded, base truncate, widest in middle, aseptate quences of members of the family Phaeosphaeriaceae (Fig. 28),
or transversely euseptate, becoming constricted with age Nst. artemisiae, which was the most recently described species
(adapted from Quaedvlieg et al. 2013). (Wanasinghe et al. 2018), is not included in the clade that rep-
resents the genus Neostagonospora, being located in the Sep-
Culture characteristics: Colonies flat, spreading, erumpent, cir- toriella clade. Therefore, this species is excluded from
cular or undulate, smooth to velvety, even margins, with sparse Neostagonospora and transferred to Septoriella (see Septoriella
to moderate aerial mycelium. On PDA, surface dirty white, below).
greyish sepia to isabelline; reverse pale white to pale pink, pale
pink with white edge, luteous or olivaceous grey to pale oliva- References: Quaedvlieg et al. 2013, Yang et al. 2016,
ceous grey. Thambugala et al. 2017 (morphology and phylogeny).
Optimal media and cultivation conditions: PDA and sterilised Neostagonospora sorghi Crous & Y. Marín, sp. nov. Myco-
Carex leaves or Anthriscus stem placed on 1.5 % water agar at Bank MB829612. Fig. 35.
25 °C under continuous near-ultraviolet light to promote Etymology: Name refers to the genus Sorghum, the host from
sporulation. which this fungus was collected.
Distribution: Africa (South Africa), Asia (Russia and Taiwan), Conidiomata solitary, erumpent, pycnidial, brown, globose,
Australia and Europe (Italy and the Netherlands). 180–200 μm diam, with central ostiole; conidiomatal wall of 2–3
Hosts: Arrhenatherum elatius, Phragmites australis, Sorghum layers of brown textura angularis. Conidiophores reduced to
halepense and Spinifex littoreus (Poaceae), Carex acutiformis conidiogenous cells lining the inner cavity. Conidiogenous cells
(Cyperaceae) and Elegia cuspidata (Restionaceae). hyaline to pale brown, smooth-walled, ampulliform, phialidic with
visible periclinal thickening and collarette, 5–8 × 4–5 μm.
Disease symptoms: Leaf spots. Conidia solitary, aseptate, ellipsoid with obtuse ends, straight,
guttulate, (4–)5–6 × (2–)2.5(–3) μm.
Notes: Neostagonospora was introduced by Quaedvlieg et al.
(2013) to accommodate two taxa associated with leaf spots on Culture characteristics: Colonies flat, spreading with moderate
Carex acutiformis (Nst. caricis) and Elegia cuspidata (Nst. ele- aerial mycelium and even, lobate margins, covering dish in 2 wk.
giae). However, its pathogenicity remains unclear since Koch’s On MEA surface hazel to isabelline, reverse isabelline; on PDA
postulates have not been completed. Subsequently, another surface isabelline with scarlet outer margins, reverse brown
foliicolous fungus was included in this genus, Nst. spinificis (Yang vinaceous; on OA surface isabelline with patches of scarlet.
et al. 2016), associated with green tissues and leaf spots of
Typus: Australia, Western Australia, Denmark, Mount Lindesay
Spinifex littoreus. The most recently introduced species are both
Walk trail, on Sorghum halepense (Poaceae), 19 Sep. 2015,
saprobes on members of Poaceae: Nst. arrhenatheri and Nst.
P.W. Crous, HPC 697A (holotype CBS H-23869, culture ex-type
phragmitis (Thambugala et al. 2017). Further studies are needed
CBS 145366 = CPC 29239).
to prove the pathogenicity of members included in this genus.
Neostagonospora is similar to Stagonospora since both pro- Notes: Neostagonospora sorghi is the first species of the genus
duce pycnidial conidiomata with euseptate, hyaline, fusoid- reported from Sorghum, and occurring in Australia. It is closely
MARIN-FELIX ET AL.
CBS 381.91ET Nothophoma anigozanthi

66
CBS 142457T Nothophoma variabilis
CBS 125.93 Nothophoma arachidis-hypogaeae

100
CBS 123395NT Nothophoma infossa

78
CGMCC 3.19246
84
100 LC12187 Nothophoma quercina
100
CBS 633.92
4x
CBS 377.67 Nothophoma gossypiicola

94
97 MCC 1082T Nothophoma raii
CBS 140674T Nothophoma macrospora
Phoma herbarum CBS 615.75

20.0
Fig. 37. Phylogenetic tree generated from a maximum parsimony analysis based on the combined LSU (868 bp), ITS (490 bp), tub2 (336 bp) and rpb2 (845 bp) sequences of
all accepted species of Nothophoma. The tree was rooted to Phoma herbarum CBS 615.75. Values above the branches represent parsimony bootstrap support values (> 50 %).
GenBank accession numbers are indicated in Table 8. T, ET and NT indicate ex-type, ex-epitype and ex-neotype strains, respectively. TreeBASE: S23494.
related to Nst. caricis, which is associated with leaf spots on Conidiogenous cells phialidic, hyaline, smooth-walled, ampulli-
Carex acutiformis. However, Nst. sorghi can be distinguished form to doliiform, sometimes flask-shaped. Conidia hyaline but
from this species and from other species of the genus by its incidentally brown, smooth- and thin-walled, aseptate, ovoid or
aseptate conidia. Moreover, Nst. sorghi produces smaller conidia ellipsoidal, eguttulate or guttulate. Chlamydospores elongated
than Nst. caricis [(4–)5–6 × (2–)2.5(–3) μm in Nst. sorghi vs. barrel-shaped, olivaceous brown, in chains. Sexual morph un-
(10–)13–16(–19) × (3–)3.5(–4) μm in Nst. caricis]. known (adapted from Chen et al. 2015, Crous et al. 2017b).
Authors: Y. Marin-Felix & P.W. Crous Culture characteristics: Colonies on OA yellow/green to oliva-
ceous grey/brown, dull green, or translucent, aerial mycelium
Nothophoma Q. Chen & L. Cai, Stud. Mycol. 82: 212. 2015.
tenuous, sometimes margins irregular and whitish, flattened or
Fig. 36.
effused, compact, floccose.
Optimal media and cultivation conditions: OA or sterile pine
sporales, Didymellaceae.
needles placed on OA under near-ultraviolet light (12 h light, 12 h
Type species: Nothophoma infossa (Ellis & Everh.) Q. Chen & L. dark) to promote sporulation at 25 °C.
Cai, basionym: Phoma infossa Ellis & Everh., J. Mycol. 4: 102.
1888. Neotype and ex-neotype strain designated by Aveskamp
et al. (2009): CBS H-20145, CBS 123395. Hosts: Wide host range, mainly occurring as pathogens, and also
endophytes or saprobes, on Amaryllidaceae, Anacardiaceae,
DNA barcodes (genus): LSU, ITS.
Fabeceae, Fagaceae, Haemodoraceae, Malvaceae, Oleaceae,
DNA barcodes (species): rpb2, tub2. Table 8. Fig. 37. Rosaceae, Rhamnaceae and Rutaceae. Also isolated from other
substrates and environments, such as soil, fungi and human
Conidiomata pycnidial, globose to elongated, or irregular, su-
infections.
perficial or immersed into the agar, solitary or confluent, ostiolate,
sometimes with an elongated neck; conidiomatal wall pseudo- Disease symptoms: Leaf spots, stem cankers, brown spot of
parenchymatous, multi-layered, outer wall pigmented. fruits, shoot canker.
52
Fig. 38. Parastagonospora spp. A–D. Sexual morph of Parastagonospora nodorum (CBS H-13909). A, B. Ascomata. C, D. Asci and ascospores. E–M. Asexual morph. E, F.
Conidiomata. E. Parastagonospora poagena (ex-type CBS 136776). F. Parastagonospora poae (CBS 135091). G–I. Conidiogenous cells. G. Parastagonospora caricis (ex-type
CBS H-21304). H. Parastagonospora poae (CBS 135091). I. Parastagonospora poagena (ex-type CBS 136776). J–M. Conidia. J. Parastagonospora caricis (ex-type CBS H-
21304). K. Parastagonospora nodorum (CBS H-13909). L. Parastagonospora poae (CBS 135091). M. Parastagonospora poagena (ex-type CBS 136776). Scale bars: C, D,
J–M = 10 μm; G–I = 5 μm. Pictures A–D, F–H, J–L taken from Quaedvlieg et al. (2013); E, I, M from Crous et al. (2014b).
Table 9. DNA barcodes of accepted Parastagonospora spp.

ITS rpb2 tef1 tub2
Parastagonospora allouniseptata MFLUCC 13-0386T KU058711 – MG520914a – Li et al. (2015), Phookamsak et al. (2017)
P. avenae CBS 289.69 KF251174 KF252182 KF253132b KF252669 Quaedvlieg et al. (2013)
MFLUCC 13-0557T of P. forlicesenica
KY769660 – – – Thambugala et al. (2017)
P. caricis CBS 135671T KF251176 KF252184 KF253134b KF252671 Quaedvlieg et al. (2013)
P. dactylidis MFLUCC 13-0375T KU058712 – – – Li et al. (2015)
MFLUCC 13-0376T of P. minima
KU058713 – MG520916a – Li et al. (2015), Phookamsak et al. (2017)
MFLUCC 13-0573 T of P. cumpignensis
KU842388 – – – Li et al. (2016a)
P. fusiformis MFLUCC 13-0215T KX926418 KX863711 – – Thambugala et al. (2017)
P. italica MFLUCC 13-0377 T
KU058714 – MG520915 a
– Li et al. (2015), Phookamsak et al. (2017)
P. nodorum CBS 110109 KF251177 KF252185 KF253135b KF252672 Quaedvlieg et al. (2013)
P. novozelandica CBS 145416T MK539957 MK540088 MK540151 MK540169 Present study
P. phragmitis CBS 143446T MK539958 MK540089 MK540152 – Present study
P. poaceicola MFLUCC 15-0471 T
KX926419 KX880499 – – Thambugala et al. (2017)
P. poae CBS 135089T KF251178 KF252186 KF253136b KF252673 Quaedvlieg et al. (2013)
P. poagena CBS 136776T KJ869116 – – – Crous et al. (2014b)
P. uniseptata MFLUCC 13-0387T KU058715 – MG520917a – Li et al. (2015), Phookamsak et al. (2017)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand. T indicates ex-type
strains.
2
elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used in sequencing: a: EF1-983F / EF1-2218R, b: EF1-728F / EF-2.
MARIN-FELIX ET AL.
Fig. 39. RAxML phylogram obtained from the combined ITS (575 bp), LSU (848 bp), rpb2 (337 bp) and tef1 (866 bp) sequence alignment of all accepted species of Par-
astagonospora. The tree was rooted to Neostagonospora carici CBS 135092 and Neostagonospora elegiae CBS 135101. The novelties proposed in this study are indicated in
are listed in Fig. 28, and of the others loci are indicated in Tables 7 and 9. T indicates ex-type strains. TreeBASE: S23834.
Notes: Nothophoma was one of the genera established recently between species are insignificant, and phylogenies based on
in order to delineate a more natural classification for the Asco- multi-locus sequence data are primarily used to distinguish
chyta-Didymella-Phoma complex (Chen et al. 2015). Currently species.
this genus comprises nine species, including five Phoma species Species in this genus are seed- and soil-borne endophytes or
previously classified in Phoma, and four species that were pathogens mainly causing leaf spots and stem canker of culti-
recently proposed (Crous et al. 2016b, 2017b, Valenzuela-Lopez vated crops and plants, such as groundnut and cotton. Some
et al. 2018). Within Nothophoma morphological differences species are mycophylic on other fungi or occur in soil, as well as
54
Fig. 40. Parastagonospora novozelandica (ex-type CPC 29613). A–C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.
in the respiratory secretion of a patient with pneumonia or in a erumpent or immersed to semi-immersed, subepidermal,
human bronchial wash sample (Boerema et al. 2004, Aveskamp globose to subglobose, ampulliform, or obpyriform, with
et al. 2009, 2010, Chen et al. 2015, Crous et al. 2016b, 2017b, central papillate ostiole, exuding creamy or pinkish conidial
Valenzuela-Lopez et al. 2018). mass; conidiomatal wall composed of 2–4 layers of brown
cells of textura angularis, or composed of 1–5 outer layers of
References: Boerema et al. 2004 (morphology and pathoge-
dark brown cells and 1–3 inner layers of hyaline cells of
nicity), Aveskamp et al. 2010, Chen et al. 2015, Crous et al.
textura angularis, or composed of an outer layer of brown to
2016b, 2017b, Valenzuela-Lopez et al. 2018 (morphology and
dark brown cells of textura globosa and an inner layer of
phylogeny).
pale brown to hyaline cells of textura angularis. Co-
Authors: Q. Chen & L. Cai nidiophores reduced to conidiogenous cells. Conidiogenous
cells phialidic, hyaline, smooth-walled, aggregated, lining the
Parastagonospora Quaedvl. et al., Stud. Mycol. 75: 362. 2013.
inner cavity, ampulliform to subcylindrical, broadly cylindrical
Fig. 38.
or broadly conical, with percurrent proliferation near apex.
Classification: Dothideomycetes, Pleosporomycetidae, Pleo- Conidia hyaline or subhyaline, smooth-walled, thin- or thick-
sporales, Phaeosphaeriaceae. walled, cylindrical, subcylindrical or fusiform, granular to
multi-guttulate, with obtuse or subobtuse apex and truncate
Type species: Parastagonospora nodorum (Berk.) Quaedvlieg
base, rarely rounded at both ends, straight to gently curved,
et al., basionym: Depazea nodorum Berk. Reference strain: CBS
sigmoid, transversely 1–9-euseptate, sometimes constricted
110109.
at the septa (adapted from Quaedvlieg et al. 2013).
DNA barcode (genus): LSU. Fig. 28.
Culture characteristics: Colonies flat, with aerial mycelium, white
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 9. Fig. 39. to pink, olivaceous, grey or vinaceous buff.
Ascomata perithecial, immersed, globose, becoming Optimal media and cultivation conditions: Sterilised Carex leaves
depressed, dark brown to black, with central ostiole with placed on 1.5 % WA at 25 °C under continuous near-ultraviolet
upper region slightly papillate; ascomatal wall thin- or thick- light to promote sporulation.
walled, composed of 2–6 layers of brown cells of textura
angularis. Pseudoparaphyses filiform, hyaline, septate. Asci
bitunicate, clavate, cylindrical, narrowly fusoid or curved, Hosts: Pathogens or saprophytes of grass (Poaceae). Species of
shortly stipitate, thick-walled, 8-spored. Ascospores fusoid or Parastagonospora are directly or indirectly responsible for sig-
ellipsoidal, hyaline or subhyaline to pale brown, smooth- nificant annual crop losses worldwide on wheat, barley and rye.
walled, transversely 3-euseptate, cells above central septum
Disease symptoms: Leaf, glume and node spots.
often broader than the lower ones, with acute rounded ends,
constricted or not at each septum, sometimes with distinct oil Notes: Parastagonospora was recently introduced in order to
droplets in each cell. Conidiomata pycnidial, brown to black, accommodate a clade of several common and serious cereal
Fig. 41. Parastagonospora phragmitis (ex-type CPC 32075). A, B. Conidiogenous cells. C, D. Conidia. Scale bars = 10 μm.
MARIN-FELIX ET AL.
Fig. 42. Phaeosphaeriopsis spp. A. Symptomatic leaves of Agapanthus precox caused by Phaeosphaeriopsis agapanthi. B. Asci and ascospores of Phaeosphaeriopsis
agavacearum (ex-type CBS 142110). C–P. Asexual morph. C, D. Conidiomata sporulating on PNA and OA, respectively, of Phaeosphaeriopsis agapanthi (ex-type CBS
141287). E. Conidiomata sporulating on OA of Phaeosphaeriopsis agavacearum (ex-type CBS 142110). F. Conidioma of Phaeosphaeriopsis agavacearum (ex-type CBS
142110). G–L. Conidiogenous cells giving rise to conidia. G, H. Phaeosphaeriopsis agapanthi (ex-type CBS 141287). I–L. Phaeosphaeriopsis glaucopunctata (CBS 653.86).
M–O. Conidia. M. Phaeosphaeriopsis agapanthi (ex-type CBS 141287). N. Phaeosphaeriopsis agavacearum (ex-type CBS 142110). O. Phaeosphaeriopsis glaucopunctata
(CBS 653.86). P. Chlamydospores of Phaeosphaeriopsis agavacearum (ex-type CBS 142110). Scale bars: F = 100 μm; others = 10 μm; I applies to I–L. Pictures A, C, D, G, H,
M taken from Crous et al. (2016b); B, E, F, N, P from Crous et al. (2016a); I–L, O from Quaedvlieg et al. (2013).
pathogens that had been previously been placed in the Stagonospora mainly in the sexual morph, being
genera Septoria/Stagonospora or Leptosphaeria/Phaeos- phaeosphaeria-like in Parastagonospora and didymella-like in
phaeria (Quaedvlieg et al. 2013). This genus differs from Stagonospora.
56
In the phylogenetic analysis based only on the ITS and LSU Quaedvlieg et al. 2013, Li et al. 2015, Thambugala et al. 2017
sequences of representative members of the family Phaeos- (morphology and phylogeny).
phaeriaceae (Fig. 28), all the species of Parastagonospora were
Parastagonospora dactylidis W.J. Li et al., Mycosphere 6: 691.
located in a well-supported clade (0.98 PP), except for
2015.
P. phoenicicola. The ex-type strain of this latter species clustered
Synonyms: Parastagonospora minima W.J. Li, et al., Myco-
in well-supported clade (84 % BS) representing the genus
sphere 6: 691. 2015.
Phaeosphaeria. Therefore, a new combination is proposed for
Parastagonospora cumpignensis Tibpromma et al., Fungal Di-
this taxon.
versity 78: 48. 2016.
In our phylogenetic analysis (Fig. 39), P. cumpignensis,
P. dactylidis and P. minima grouped in the same well-supported Typus: Italy, Province of Arezzo, Passo della Consuma, on dead
clade (79 % BS / 0.96 PP) without significant phylogenetic dis- stem of Dactylis sp. (Poaceae), 19 Jun. 2012, Erio Camporesi
tance. The three species all have been isolated from Thailand on (holotype MFLU 15-0693, culture ex-type MFLUCC 13-
dead stems of Dactylis (Li et al. 2015, 2016a). Only ITS se- 0375 = ICMP 20774 = KUMCC15-0131).
quences are available for P. cumpignensis and P. dactylidis, and
Additional materials: Italy, Province of Arezzo, Passo della Consuma, on
nucleotide similarity for this locus for all three species is 100 %.
dead stem of Dactylis sp. (Poaceae), 19 Jun. 2012, Erio Camporesi,
Therefore, these three species are reduced to synonymy. The
MFLUCC 13-0376 = ICMP 20776 = KUMCC15-0132; ibid., Campigna, Santa
same problem is found in P. forlicesenica, which is one of the
Sofia, ForlìCesena Province, on dead stem of Dactylis glomerata (Poaceae),
most recently described species in the genus (Thambugala et al.
23 Jun. 2012, Erio Camporesi, MFLUCC 13-0573.
2017). Based on ITS, P. forlicesenica shares a nucleotide simi-
larity of 99.8 % with P. avenae. Therefore, further studies should Notes: In our phylogenetic analysis (Fig. 39), the ex-type strains
be done to confirm if P. forlicesenica represents a separate of P. cumpignensis (MFLUCC 13- 0573), P. dactylidis (MFLUCC
species or should be synonymised with P. avenae. 13-0375) and P. minima (MFLUCC 13-0376) grouped in the
Species of Parastagonospora are pathogens or saprophytes same well-supported clade (79 % BS / 0.96 PP) without signif-
of grasses, being directly or indirectly responsible for significant icant phylogenetic distance. The ITS sequences of the three
annual crop losses worldwide. Parastagonospora avenae cau- species showed a nucleotide similarity of 100 %. Moreover, all of
ses minor leaf blotch of barley and rye, while it is considered an them were isolated from from Thailand on dead stems of Dactylis
important pathogen of oats (Cunfer 2000). Parastagonospora (Li et al. 2015, 2016a). Therefore, the three species are herewith
nodorum is known primarily as a major necrotrophic pathogen of reduced to synonymy.
wheat that causes leaf and glume blotch, but also infects barley,
on which it is considered as not economically important (Cunfer Parastagonospora novozelandica Crous, Thangavel & Y.
2000, Oliver et al. 2012). Marín, sp. nov. MycoBank MB829668. Fig. 40.
References: Cunfer 2000 (pathology and morphology), Oliver Etymology: Name refers to New Zealand, the country where this
et al. 2012 (pathology, genomics and host resistence), fungus was isolated.
Table 10. DNA barcodes of accepted Phaeosphaeriopsis spp.

ITS rpb2 tef1 tub2
Phaeosphaeriopsis agapanthi CBS 141287T KX228260 MK540094 MK540157a MK540173 Crous et al. (2016b), present study
T a
Phs. agavacearum CBS 142110 KY173430 KY173591 MK540158 KY173610 Crous et al. (2016a), present study
Phs. agavensis CBS 102206 KY090635 KY090685 – – Ahmed et al. (2017)
Phs. aloes CBS 145367 T
MK539959 MK540090 MK540153 a
– Present study
Phs. aloicola CBS 145368T MK539960 MK540091 MK540154a MK540170 Present study
Phs. amblyospora CBS 110131T AY188993 – – – Câmara et al. (2003)
Phs. dracaenicola MFLUCC 11-0157T KM434273 KM434309 KM434301b – Phookamsak et al. (2014b)
Phs. glaucopunctata MFLUCC 13-0265ET KJ522473 – MG520918b – Thambugala et al. (2014), Phookamsak et al. (2017)
Phs. grevilleae CBS 145369T MK539961 MK540092 MK540155a MK540171 Present study
Phs. nolinae CBS 102205 KY090637 KY090686 – – Ahmed et al. (2017)
Phs. obtusispora CBS 102204 KY090636 KY090687 – – Ahmed et al. (2017)
Phs. phacidiomorpha T111 FJ462742 – – – Zhang et al. (unpubl. data)
Phs. pseudoagavacearum CBS 145370T MK539962 MK540093 MK540156a MK540172 Present study
Phs. triseptata MFLUCC 13-0271T KJ522475 KJ522485 MG520919b – Thambugala et al. (2014), Phookamsak et al. (2017)
Phs. yuccae MFLUCC 16-0558 T
KY554482 – MG520920b – Phookamsak et al. (2017)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand; T: isolate housed
in China. T and ET indicate ex-type and ex-epitype strains.
2
elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used in sequencing: a: EF1-728F / EF-2, b: EF1-983F / EF1-2218R.
MARIN-FELIX ET AL.
Fig. 43. RAxML phylogram obtained from the combined ITS (587 bp), LSU (849 bp), rpb2 (838 bp), tef1 (601 bp) and tub2 (519 bp) sequence alignment of all accepted species
of Phaeosphaeriopsis. The tree was rooted to Neostagonospora caricis CBS 135092 and Neostagonospora elegiae CBS 135101. The novelties proposed in this study are
indicated in bold. RAxML bootstrap support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession
numbers of LSU are listed in Fig. 28, and of the other loci in Tables 7 and 9. T and ET indicate ex-type and ex-epitype strains, respectively. TreeBASE: S23834.
Culture nearly sterile, with only a few conidiomata observed. Typus: New Zealand, Browns Bay, on unidentified grass
Conidiomata solitary, pycnidial, dark brown, globose, (Poaceae), Nov. 2015, R. Thangavel (holotype CBS H-
180–200 μm diam, with central ostiole; conidiomatal wall with 23903, culture ex-type T15–06960B = CPC 29613 = CBS
3–6 layers of pale brown textura angularis. Conidiophores 145416).
reduced to conidiogenous cells lining the inner cavity. Con-
Notes: Parastagonospora novozelandica is related to
idiogenous cells hyaline, smooth, ampulliform to subcylindrical,
P. allouniseptata. Both species produce 1-septate conidia, but
6–8 × 2.5–5 μm, proliferating percurrently at apex. Conidia
these can be easily distinguished based on their conidial di-
solitary, hyaline to pale olivaceous, smooth, guttulate, sub-
mensions [(9–)11–13(–16) × (2–)2.5(–3) μm in
cylindrical, straight, apex subobtuse, base truncate, 1-septate,
P. novozelandica vs. 16–22 × 2.5–3.5 μm in P. allouniseptata].
(9–)11–13(–16) × (2–)2.5(–3) μm.
Parastagonospora phragmitis Crous & Y. Marín, sp. nov.
Culture characteristics: Colonies flat, spreading, reaching
60 mm diam after 2 wk, with moderate aerial mycelium, and even,
smooth margins. On MEA surface greenish olivaceous to umber, Etymology: Name reflects the genus Phragmites from which this
reverse olivaceous to umber; on OA surface brown vinaceous. fungus was isolated.
58
Fig. 44. Phaeosphaeriopsis aloes (ex-type CBS 145367). A. Conidiomata sporulating on MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 180 μm; all
others = 10 μm.
Conidiomata solitary, pycnidial, brown, globose, 250–300 μm Basionym: Parastagonospora phoenicicola Crous & Thangavel,
diam, with central ostiole; conidiomatal wall with 3–6 layers of Persoonia 37: 349. 2016.
pale brown cells of textura angularis. Conidiophores reduced to
conidiogenous cells lining the inner cavity. Conidiogenous cells Description and illustration: Crous et al. (2016a).
hyaline, smooth, ampulliform to doliiform, 7–10 × 8–9 μm, Typus: New Zealand, Auckland, Botany road, on leaves of
proliferating percurrently at apex. Conidia solitary, hyaline to pale Phoenix canariensis (Arecaceae), 2015, R. Thangavel (holotype
olivaceous, smooth, guttulate, subcylindrical-fusoid, straight to CBS H-22892, culture ex-type CPC 28711 = CBS 142107).
slightly curved, with prominent taper in upper third to subobtuse
apex, widest in middle to lower third, base truncate, 3-septate, Notes: In the phylogenetic analysis based on ITS and LSU se-
(18–)23–25(–27) × (3–)4 μm. quences (Fig. 28), the ex-type strain of this species was located
in the well-supported clade (84 % BS) representing the genus
Culture characteristics: Colonies flat, spreading, covering dish Phaeosphaeria. Morphologically, this species produces conidia
after 2 wk, with moderate aerial mycelium, and smooth, lobate more similar to Phaeosphaeria than to Parastagonospora, since
margins. On MEA surface saffron, reverse sienna; on PDA these are subcylindrical, mostly straight, while in Para-
surface saffron to sienna, reverse sienna; on OA surface pale stagonospora the conidia tend to be sigmoid and longer than in
luteous to saffron. P. phoenicicola. Based on morphology and molecular data, the
Typus: Australia, New South Wales, Sussex Inlet, on Phrag- new combination, Phaeosphaeria phoenicicola, is herewith
mites sp. (Poaceae), 27 Nov. 2015, P.W. Crous, HPC 1785 proposed.
(holotype CBS H-23902, culture ex-type CPC 32075 = CBS Authors: Y. Marin-Felix, R. Thangavel & P.W. Crous
143446).
Phaeosphaeriopsis M.P.S. Câmara et al., Mycol. Res. 107: 519.
Notes: Parastagonospora phragmitis is related to P. fusiformis. 2003. Fig. 42.
However, P. phragmitis produces an asexual morph, while in
P. fusiformis only the sexual morph has been observed. More- Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
over, P. phragmitis is the first species of the genus reported on sporales, Phaeosphaeriaceae.
Phragmites. Type species: Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S.
Phaeosphaeria phoenicicola (Crous & Thangavel) Y. Marín & Câmara et al., basionym: Cryptosphaeria glaucopunctata Grev.
Crous, comb. nov. MycoBank MB829700. [as “glauco-punctata”]. Epitype and ex-epitype strain designated
Fig. 45. Phaeosphaeriopsis aloicola (ex-type CBS 145368). A. Ascomata sporulating on SNA. B. Asci. C. Pseudoparaphyses. D. Ascospores. Scale bars: A = 200 μm; all
others = 10 μm.
MARIN-FELIX ET AL.
Fig. 46. Phaeosphaeriopsis grevilleae (ex-type CBS 145369). A. Conidiomata sporulating on PDA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.
by Thambugala et al. (2014): MFLU 14-0029, MFLUCC 13- cream at the margins, brown to orange-brown in the middle and
0265 = ICMP 20199. pale yellowish at the centre.
DNA barcode (genus): LSU. Fig. 28. Optimal media and cultivation conditions: On MEA, PDA or OA at
25 °C.
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 10. Fig. 43.
Ascomata solitary or aggregated, immersed, subepidermal to
erumpent, pushing up flaps of the epidermis, globose to pyriform, Hosts: Pathogens or saprophytes on Agapanthus praecox
often papillate, solitary or gregarious in a stroma of scle- (Alliaceae), Agave spp. and Yucca spp. (Agavaceae), Aloe
roplectenchyma or dark brown cells of textura angularis, often sp. (Aloaceae), Dracaena lourieri, Dracaena sp. and Nolina
surrounded by septate, brown hyphae extending into the host erumpens (Dracaenaceae), Grevillea sp. (Proteaceae),
tissues. Asci 8-spored, bitunicate, cylindrical to broadly fusoid, Phormium spp. (Phormiaceae) and Ruscus spp.
short stipitate, with visible apical chamber. Ascospores uni- to (Ruscaceae).
triseriate, cylindrical, broadly rounded at apex, tapering to Disease symptoms: Leaf spots and leaf blight.
narrowly rounded base, 4–5-septate, first septum submedian,
often constricted, medium brown, echinulate, punctate or Notes: The genus Phaeosphaeriopsis was introduced by
verrucose. Asexual morph coniothyrium-like or Câmara et al. (2003) to accommodate some species of
phaeostagonospora-like. Conidiomata pseudoparenchymatous, Paraphaeosphaeria that were not congeneric based on
sometimes of scleroplectenchyma. Conidiogenous cells lining phylogenetic data. Phaeosphaeriopsis is characterised by
locule, ampulliform, hyaline, proliferating percurrently, resulting in having uni- or multi-loculate stromata and 4–5-septate
inconspicuous annellations. Conidia cylindrical, with bluntly ascospores, and coniothyrium-like and phaeostagonospora-
rounded ends, 0–3-septate, yellowish brown, punctate like asexual morphs, while Paraphaeosphaeria produces
(Quaedvlieg et al. 2013). 2-septate ascospores and has a microsphaeropsis-like
asexual morph (Câmara et al. 2003, Quaedvlieg et al.
Culture characteristics: Colonies flat or rarely slightly raised, 2013). Phaeosphaeriopsis is related to Acericola, which is
spreading, feathery, velvety or floccose, with sparse to moderate a genus recently introduced to accommodate a saprobic
aerial mycelium, circular or lobate, margins smooth or rarely fungus found on dead twigs of Acer campestre (Hyde et al.
slightly radiating. On PDA, surface white, dirty white, pinkish 2017). Unfortunately, it appears that the LSU sequence of
white, primrose, pale grey or pale luteous; reverse dirty white, Acericola is incorrect (Crous et al. 2019a).
light to dark grey, luteous or olivaceous buff. On MEA, surface In our phylogenetic analysis based on ITS and LSU se-
dirty white, pale luteous, or white to cream at the margins, pale quences, 11 species are accepted in the genus Phaeosphaer-
yellowish to yellowish brown in the middle and pale brown to iopsis, and four strains located in independent branches are
brown or orange-brown at the centre, with small white to grey introduced as new species.
droplets; reverse luteous, umber with patches of dirty white, Species included in Phaeosphaeriopsis are saprobes or
isabelline in the middle and cinnamon in outer region, or white to presumed pathogens. The type species, Phaeosphaeriopsis
Fig. 47. Phaeosphaeriopsis pseudoagavacearum (ex-type CBS 145370). A. Conidiomata sporulating on MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.
60
Fig. 48. Pleiocarpon strelitziae (ex-type CBS 142251). A–F. Disease symptoms. A, B. Wilting and dying Strelitzia reginae plants in the nursery. C–F. Basal rot and wilting of
plant induced during the pathogenicity test. G–L. Asexual morph. G, H. Simple conidiophores. I, J. Sporodochia. K. Microconidia. L. Macroconidia. Scale bars: 10 μm; G applies
to G–L. Pictures taken from Aiello et al. (2017).
glaucopunctata, is associated with leaf spot and necrosis of solitary, aseptate, straight, verruculose, golden-brown, sub-
Ruscus aculeatus (Câmara et al. 2003, Golzar & Wang cylindrical, apex obtuse, base bluntly rounded, (4–)
2012). Phaeosphaeriopsis agapanthi and Phs. dracaeni- 5(–6) × 3(–3.5) μm.
cola are also associated with necrotic leaf spots of
Culture characteristics: Colonies flat, spreading, with sparse to
Agapanthus precox and Dracaena lourieri, respectively
moderate aerial mycelium, covering dish in 2 wk. On MEA sur-
(Phookamsak et al. 2014b, Crous et al. 2016b).
face honey, reverse cinnamon; on PDA surface and reverse
References: Câmara et al. 2003, Quaedvlieg et al. 2013, sepia; on OA surface saffron.
Thambugala et al. 2014 (morphology and phylogeny). Typus: USA, California, on leaves of Aloe sp. (Aloaceae), 6 Aug.
Phaeosphaeriopsis aloes Crous & Y. Marín, sp. nov. Myco- 2016, P.W. Crous, HPC 1326 (holotype CBS H-23870, culture
Bank MB829642. Fig. 44. ex-type CBS 145367 = CPC 31480).
Notes: Phaeosphaeriopsis aloes is related to Phs. obtusispora.
Etymology: Name refers to Aloe, the host from which this fungus
This latter species only produces the sexual morph, while only
was collected.
the asexual morph has been observed in our new species.
Conidiomata solitary, brown, pycnidial, globose, 150–180 μm Phaesphaeriopsis aloes and Phs. aloicola, both described
diam, with central ostiole, 30–40 μm diam; conidiomatal wall of here, are the first species of the genus found on a member of the
3–4 layers of brown cells of textura angularis. Conidiophores family Aloaceae, an Aloe sp. However, the two species are not
reduced to conidiogenous cells lining the inner cavity, hyaline, related. Phaeosphaeriopsis aloes produces an asexual morph,
smooth-walled, ellipsoid, phialidic, 4–6 × 3–4 μm. Conidia while in Phs. aloicola only the sexual morph has been observed.
MARIN-FELIX ET AL.
Table 11. DNA barcodes of accepted Pleiocarpon spp.

ITS his3 rpb2 tef1 tub2
Pleiocarpon livistonae CBS 145030T MK539963 MK540234 MK540095 MK540165 MK540179 Present study
Pl. strelitziae CBS 142251T KY304644 KY304616 KY304697 KY304722 KY304750 Aiello et al. (2017)
1 T
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; his3: partial histone H3 gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene;
tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
Phaeosphaeriopsis aloicola Crous & Y. Marín, sp. nov. not been observed in Phs. aloicola. Phaeosphaeriopsis aga-
MycoBank MB829643. Fig. 45. panthi only produces an asexual morph. The three species
were isolated from different hosts in different families.
Etymology: Name refers to the host genus Aloe from which this
Phaeosphaeriopsis agapanthi was isolated from Agapanthus
fungus was collected.
precox (Amaryllidaceae), Phs. aloicola from Aloe sp. (Aloa-
Ascomata solitary, aggregated, erumpent, brown, globose, ceae), and Phs triseptata from Ruscus aculeatus (Aspar-
150–200 μm diam, with papillate neck and central ostiole, agaceae). Moreover, Phs. aloicola was found in the USA
40–50 μm diam; ascomatal wall of 4–6 layers of brown cells of while the other species have been reported from Europe.
textura angularis. Pseudoparaphyses hyaline, smooth-walled, Phaeosphaeriopsis aloes was also isolated from Aloe in
hyphae-like, 2–3 μm diam, anastomosing, branched, inter- California. For comparison see notes of Phaeosphaeriopsis
mingled among asci. Asci bitunicate, subcylindrical, apex obtuse aloes.
with well-defined apical chamber, 2 μm diam, fasciculate, short
stipitate, 70–100 × 10–12 μm. Ascospores bi- to triseriate, Phaeosphaeriopsis grevilleae Crous & Y. Marín, sp. nov.
subcylindrical, straight to slightly curved, (2–)3-septate, at times MycoBank MB829644. Fig. 46.
slightly swollen in second cell from apex, medium brown, ver- Etymology: Name refers to the host genus Grevillea from which
ruculose, ends obtuse, (19–)22–25(–26) × (4–)5(–6) μm.
this fungus was collected.
Culture characteristics: Colonies flat, spreading, with sparse to
Conidiomata solitary, pycnidial, scattered, globose, 180–250 μm
moderate aerial mycelium and even, lobate margins, reaching diam, with central ostiole, exuding black mucoid conidial mass;
30 mm diam on PDA, covering dish on MEA and OA. On MEA
conidiomatal wall of 2–3 layers of brown cells of textura angu-
surface buff, outer region sienna, reverse sienna; on PDA sur-
laris. Conidiophores reduced to conidiogenous cells lining inner
face and reverse buff to sienna; on OA surface scarlet.
cavity, hyaline, smooth-walled, ampulliform, phialidic,
Typus: USA, California, on leaves of Aloe sp. (Aloaceae), 6 Aug. 5–7 × 3–5 μm. Conidia solitary, aseptate, medium brown, ver-
2016, P.W. Crous, HPC 1306 (holotype CBS H-23871, culture ruculose, ellipsoid to ovoid, (4–)5(–6) × (3–)3.5(–4) μm.
ex-type CBS 145368 = CPC 31454).
Culture characteristics: Colonies flat, spreading, with moderate
Notes: Phaeosphaeriopsis aloicola is related to Phs. aga- aerial mycelium and feathery margins, reaching 35 mm diam on
panthi and Phs. triseptata. Morphologically, Phs. aloicola is MEA, covering dish on PDA and OA. On MEA surface and
similar to Phs. triseptata since both produce verruculose, 3- reverse scarlet; on PDA surface ochreous, reverse sienna; on
septate ascospores. However, both species differ in the OA surface scarlet with patches of ochreous.
size of ascomata (up to 200 μm in Phs. aloicola vs. up to
Typus: Australia, Queensland, leaves of Grevillea sp. (Protea-
110 μm in Phs. triseptata), asci (70–100 × 10–12 μm in
ceae), 14 Jul. 2009, P.W. Crous, (holotype CBS H-23872, cul-
Phs. aloicola vs. 56−70 × 7.5−9 μm in Phs. triseptata) and
ture ex-type CBS 145369 = CPC 17003).
ascospores [(19–)22–25(–26) × (4–)5(–6) μm in Phs. aloi-
cola vs. 14.5−18 × 3−4 μm in Phs. triseptata]. Moreover, Notes: In the phylogenetic analysis based on ITS, LSU, rpb2,
Phs. triseptata also produces an asexual morph, which has tef1 and tub2, this species was located in an independent
Fig. 49. Pleiocarpon livistonae (ex-type CBS 145030). A. Sporodochium on SNA. B. Conidiophores with conidiogenous cells. C. Conidia. D. Chlamydospores. Scale
bars = 10 μm.
62
Fig. 50. Pyrenophora spp. A–D. Sexual morph. A, B. Sterile ascomata of Pyrenophora campanulata (CBS 127927). C. Protoascomata of Pyrenophora erythrospila on PDA
(CBS 312.69). D. Protoascoma of Pyrenophora erythrospila (CBS 108941). E–K. Asexual morph. E–H. Conidiophores. E. Pyrenophora fugax (CBS 509.77). F. Pyrenophora
novozelandica (CBS 127934). G. Pyrenophora erythrospila (CBS 312.69). H. Pyrenophora fugax (CBS 509.77). I, J. Conidia. I. Pyrenophora erythrospila (CBS 312.69).
J. Pyrenophora fugax (CBS 509.77). K. Chlamydospores of Pyrenophora tetrarrhenae (CBS 127924). Scale bars: A = 50 μm; D = 20 μm; G–K = 10 μm.
branch. This is the first species reported on Grevillea, which is a of 2–3 layers of brown cells of textura angularis. Co-
member of the Proteaceae. nidiophores reduced to conidiogenous cells lining the inner
cavity, hyaline, smooth-walled, ampulliform, phialidic,
Phaeosphaeriopsis pseudoagavacearum Crous & Y. Marín,
5–7 × 3–4 μm. Conidia solitary, golden-brown, verruculose,
sp. nov. MycoBank MB829645. Fig. 47.
thick-walled, straight to slightly curved, 1-septate, sub-
Etymology: Named after its similarity to Phaeosphaeriopsis cylindrical, apex obtuse, base bluntly rounded, (6–)
agavacearum. 8–9(–10) × 4(–4.5) μm.
Conidiomata solitary, globose, brown, pycnidial, 200–250 μm Culture characteristics: Colonies flat, spreading, with moderate
diam, with central ostiole, 15–20 μm diam; conidiomatal wall aerial mycelium and even, smooth margins, covering dish in
MARIN-FELIX ET AL.
Table 12. DNA barcodes of accepted Pyrenophora spp.

ITS LSU gapdh rpb2
Pyrenophora avenicola CBS 307.84T MK539972 MK540042 MK540180 – Present study
Py. biseptata CBS 307.69 MK539973 MK540043 MK540181 – Present study
CBS 319.69 MK539974 MK540044 MK540182 MK540102 Present study
CBS 108963 MK539975 JN712532 MK540183 – Crous et al. (2011), present study
Py. bromi CBS 311.68 MK539976 MH870851 MK540184 – Vu et al. (2019), present study
DAOMC 127414 JN943666 JN940074 AY004839 – Zhang & Berbee (2001), Hambleton (unpubl.
data)
Py. chaetomioides CBS 279.31A MK539977 MK540045 MK540185 MK540103 Present study
CBS 195.31 MK539978 MH866633 MK540186 MK540104 Vu et al. (2019), present study
Py. cynosuri CBS 127918T MK539980 MK540047 MK540188 MK540106 Present study
Py. dactylidis DAOMC 92161 JN943667 JN940087 AY004812 – Zhang & Berbee (2001), Hambleton (unpubl.
data)
Py. dictyoides DAOMC 63666 JN943653 JN940080 AY004836 – Zhang & Berbee (2001), Hambleton (unpubl.
data)
CBS 127933 MH877971 MK540050 MK540191 MK540109 Vu et al. (2019), present study
Py. erythrospila CBS 312.69 MK539983 MK540051 MK540192 – Present study
CBS 108941 MK539984 MK540052 MK540193 MK540110 Present study
Py. fugax CBS 509.77 MK539985 MK540053 MK540194 MK540111 Present study
Py. grahamii CBS 315.69 MK539986 MK540054 MK540195 – Present study
CBS 128043 MK539987 MH876230 MK540196 MK540112 Vu et al. (2019), present study
Py. leucospermi CBS 111083T JN712467 JN712533 MK540198 MK540114 Crous et al. (2011), present study
CBS 111505 MK539989 JN712542 MK540199 MK540115 Crous et al. (2011), present study
CBS 114493 MK539990 JN712545 MK540200 MK540116 Crous et al. (2011), present study
Py. lolii CBS 240.48 MK539991 MK540055 MK540201 MK540117 Present study
Py. nisikadoi CBS 190.29ET KM257054 KM243296 KM257057 – Manamgoda et al. (2014)
CBS 119213 EU552124 MK540056 MK540204 MK540120 Marincowitz et al. (2008), present study
CBS 127912 MH877963 MK540057 MK540205 MK540121 Vu et al. (2019), present study
Py. nobleae CBS 259.80 MK539994 MK540058 MK540206 MK540122 Present study
Py. novozelandica CBS 127934T MK539997 MK540061 MK540209 MK540125 Present study
Py. phaeocomes DAOMC 222769 JN943649 JN940093 – DQ497614 Hambleton (unpubl. data), James et al.
(unpubl. data)
Py. poae CBS 319.68A MK539998 MK540062 MK540210 MK540126 Present study
DAOMC 145373 JN943650 JN940083 AY004832 JN993632 Zhang & Berbee (2001), Schoch et al. (2012),
Hambleton (unpubl. data)
Py. pseudoerythrospila CBS 127931T MK540000 MK540063 MK540212 – Present study
Py. semeniperda DAOMC 213153 JN943665 JN940088 AY004826 – Zhang & Berbee (2001), Hambleton (unpubl.
data)
BRIP 10941 KJ415564 KJ415518 KJ415382 – Tan et al. (2014)
Py. sieglingiae CBS 127930 MK540002 MK540065 MK540214 MK540129 Present study
T of Py. teres f. maculata
Py. teres CBS 228.76 MK540003 MK540066 MK540215 MK540130 Present study
CBS 281.31A of Py. japonica
MK540004 MK540067 MK540216 MK540131 Present study
64

ITS LSU gapdh rpb2
Py. tetrarrhenae DAOMC 171966 JN943663 JN940090 – JN993620 Schoch et al. (2012), Hambleton (unpubl. data)
Py. trichostoma CBS 328.53 MK540012 MK540072 MK540224 MK540139 Present study
CBS 391.54 MK540013 MK540073 MK540225 – Present study
Py. triseptata CBS 128047 MK540015 MH877983 MK540227 MK540141 Vu et al. (2019), present study
Py. tritici-repentis CBS 259.59SynT of Py. tritici-vulgaris
MK540017 MK540075 AM884276 MK540143 Lepoint et al. (2010), present study
Py. variabilis CBS 127920T MK540020 MK540078 MK540231 MK540146 Present study
Py. wirreganensis CBS 109896 MK540021 MK540079 MK540232 MK540147 Present study
1
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; DAOMC: Plant Research
Institute, Department of Agriculture (Mycology), Ottawa, Canada. A, ET, SynT and T indicate authentic, ex-epitype, ex-syntype and ex-type strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase
gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene.
2 wk. On MEA surface ochreous, reverse bay; on PDA surface branched, septate, bearing up to two conidiogenous cells.
sienna with bay outer region, reverse bay; on OA surface sienna Conidiogenous cells monophialidic, cylindrical, tapering slightly
with scarlet margins. towards the apex. Macroconidia cylindrical to subcylindrical,
hyaline, straight to curved, 1−6-septate, apex or apical cell
Typus: France, Domaine la Fraysse, Valgorge, on leaves of
typically slightly bent to one side and minutely beaked, base with
Agave sp. (Asparagaceae), 15 Jul. 2010, P.W. Crous (holotype
sometimes visible, centrally located or laterally displaced hilum.
CBS H-23873, culture ex-type CBS 145370 = CPC 18383).
Microconidia absent or abundant, aseptate, hyaline, ellipsoid to
Notes: Phaeosphaeriopsis pseudoagavacearum is closely ovoid or subcylindrical, straight to slightly curved, with clearly
related to Phs. agavacearum. Morphologically these species are laterally displaced hilum. Chlamydospores absent or solitary,
also similar in producing verruculose, aseptate conidia. However, globose, brown, thich-walled, guttulate. Sexual morph not
they differ in the size of their conidiomata (up to 180 μm diam in observed (adapted from Aiello et al. 2017).
Phs. agavacearum vs. up to 250 μm diam in Phs. pseudoaga-
vacearum) and conidia [(5–)6–7(–9) × 3(–4) μm in Phs. aga- Culture characteristics: Colonies on PDA with sparse to mod-
vacearum vs. (6–)8–9(–10) × 4(–4.5) μm in Phs. erate aerial mycelium, even, smooth, with lobate margins; sur-
pseudoagavacearum]. face and reverse umber or cinnamon to honey.
Authors: Y. Marin-Felix & P.W. Crous Optimal media and cultivation conditions: On PDA, MEA, OA or
SNA with sterile filter paper and carnation leaf pieces at 25 °C.
Pleiocarpon L. Lombard & D. Aiello, IMA Fungus 8: 73. 2017.
Fig. 48. Distribution: Italy and Sri Lanka.
Classification: Sordariomycetes, Hypocreomycetidae, Hypo- Host: Livistona rotundifolia (Arecaceae) and Strelitzia reginae
creales, Nectriaceae. (Strelitziaceae).
Type species: Pleiocarpon strelitziae L. Lombard & D. Aiello. Disease symptoms: Basal rot and wilt.
Holotype and ex-type strain: CBS H-22967, CBS 142251.
Notes: Pleiocarpon was recently introduced by Aiello et al.
DNA barcode (genus): LSU. (2017) to accommodate a new species isolated from potted
plants of Strelitzia reginae in an ornamental nursery located in
DNA barcodes (species): ITS, his3, rpb2, tef1, tub2. Table 11.
eastern Sicily that had a new basal rot disease. This basal stem
Conidiophores simple or aggregated, forming sporodochia; rot disease resulted in the detachment of the roots from the stem.
simple conidiophores arising laterally or terminally from aerial Moreover, the diseased plants displayed symptoms of general
mycelium, solitary to loosely aggregated, unbranched or sparsely wilting and rot of the internal foliage. Pathogenicity tests indicated
MARIN-FELIX ET AL.
66
that Pl. strelitziae, was highly aggressive, killing all inoculated Authors: Y. Marin-Felix, W. Quaedvlieg & P.W. Crous
test plants within 2 mo (Aiello et al. 2017).
Pyrenophora Fr., Summa veg. Scand. 2: 397. 1849. Fig. 50.
The phylogenetic analysis based on ITS, LSU, tef1 and tub2
Synonyms: Polytrichia Sacc., Syll. fung. (Abellini) 1: 451. 1882.
demonstrated that Pleiocarpon is closely related to the genus
Pleospora subgen. Scleroplea Sacc., Syll. fung. (Abellini) 2: 277.
Thelonectria, with both genera being characterised by
1883.
cylindrocarpon-like asexual morphs (Aiello et al. 2017). Recently,
Neilreichina Kuntze, Revis. gen. pl. (Leipzig) 2: 862. 1891.
Thelonectria was segregated by introducing three new genera,
Scleroplea (Sacc.) Oudem., Verslag. Meded. K. Akad.
Cinnamomeonectria, Macronectria and Tumenectria (Salgado-
Wetensch., Afd. Natuurk., ser. 3 9: 152. 1900.
Salazar et al. 2016). These four related genera are mostly
Drechslera S. Ito, Proc. Imp. Acad. Japan 6: 355. 1930.
found on bark of exposed wood of dead, dying or diseased trees,
Marielliottia Shoemaker, Canad. J. Bot. 76: 1559. 1999.
and are rarely associated with small cankers and root rots
(Chaverri et al. 2011, Salgado-Salazar et al. 2016). Moreover, Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
Pleiocarpon can be distinguished from Thelonectria and these sporales, Pleosporaceae.
three new genera by the absence of a sexual morph.
Type species: Pyrenophora phaeocomes (Rebent.) Fr., basio-
Hitherto, Pleiocarpon was monospecific. Here we introduce a
nym: Sphaeria phaeocomes Rebent. Neotype specimen desig-
new species isolated from Livistona rotundifolia (Arecaceae) in
nated by Shoemaker (1961): UPS 170980. Representative
Sri Lanka, causing root and corm rot.
strain: DAOM 222769.
References: Aiello et al. 2017 (morphology, pathogenicity and
DNA barcodes (genus): LSU, ITS.
phylogeny).
DNA barcodes (species): ITS, gapdh, tef1. Table 12. Fig. 51.
Pleiocarpon livistonae Crous & Quaedvl., sp. nov. MycoBank
MB829613. Fig. 49. Ascomata perithecial, immersed, becoming erumpent to near
superficial, solitary or scattered, globose to subglobose, broadly
Etymology: Name refers to Livistona, the host from which this
or narrowly conical, smooth-walled, with central ostiole; necks
fungus was collected.
papillate, covered with brown to reddish brown setae, which are
Conidiophores simple, solitary or aggregated, forming spor- darkened at the base; ascomatal wall comprising 2–4 layers of
odochia, arising from superficial hyphae, branched, 2–4-septate, brown, thick-walled cells of textura angularis. Pseudoparaphyses
50–120 × 6–8 μm. Conidiogenous cells monophialidic, cylin- not observed. Asci 8-spored, bitunicate, fissitunicate, clavate to
drical, tapering slightly towards apex, 10–30 × 4–5 μm, forming subcylindrical, with a short, broad pedicel, with a distinct ocular
conidia in false chains that eventually aggregate in a mucoid chamber surrounded by a large apical ring. Ascospores 2–3-
mass. Macroconidia hyaline, smooth-walled, subcylindrical, 1–6- seriate, muriform, constricted at the septum, smooth-walled,
septate, curved, apex subobtuse, with base sometimes visible as surrounded by a mucilaginous sheath. Conidiophores semi- to
lateral hilum; 1–3 septate conidia (21–)28–34(–45) × (5–)6 μm; macronematous, mononematous, sometimes caespitose,
4–6-septate conidia (45–)55–65(–70) × (5–)6 μm. Chlamydo- straight or flexuous, often geniculate, usually unbranched,
spores solitary, globose, brown, thick-walled, guttulate, sometimes branched, pale brown to brown, rarely subhyaline to
15–20 μm diam. pale brown. Conidiogenous cells polytretic, integrated, terminal,
frequently becoming intercalary, sympodial, cylindrical, smooth-
walled, or less frequently verruculose, cicatrised. Conidia soli-
aerial mycelium and smooth, lobate margins, reaching
tary, in certain species also sometimes catenate or forming
60 mm diam after 2 wk at 25 °C. On MEA surface ochreous,
secondary conidiophores which bear conidia, acropleurogenous,
reverse umber; on PDA surface and reverse umber; on OA
simple, straight or curved, cylindrical, ellipsoidal or obclavate,
surface umber.
less frequently subglobose, obpyriform or fusiform, tapering to-
Typus: Sri Lanka, on Livistona rotundifolia (Arecaceae), W. wards apex, straw-coloured or pale to dark brown or olivaceous
Quaedvlieg, NAK Tuinbouw, INS-17-20656D (holotype CBS H- brown, sometimes the end cells are paler than the intermediate
23849, culture ex-type CBS 145030 = CPC 34576). ones, smooth-walled or verruculose, pseudoseptate; hila protu-
berant or flat, darkened, thickened (adapted from Ellis 1971,
Notes: Pleiocarpon livistonae is distinguished from Pl. strelitziae
Ariyawansa et al. 2014).
by the absence of microconidia, the production of chlamydo-
spores, and the septation and size of its macroconidia (1–6- Culture characteristics: Colonies flat or umbonate, cottony,
septate, up to 70 μm in Pl. livistonae vs. 1–5-septate, up to sometimes granular or powdery, with moderate to abundant
50 μm in Pl. strelitziae). Pleiocarpon livistonae is phylogenetically aerial mycelium, sometimes with sparse aerial mycelium, mar-
close but clearly differentiated from Pl. strelitziae based on ITS, gins fringed, sometimes arachnoid. On PDA smoke-grey to
his3, rpb2, tef1 and tub2 sequence similarity (96 %, 85 %, 92 %, olivaceous or olivaceous grey, primrose to greyish yellow-green,
94 %, and 91 %, respectively). Moreover, Pl. livistonae was greenish grey to olivaceous black, olivaceous black with patches
isolated from Livistona rotundifolia (Arecaceae) in Sri Lanka, white for the aerial mycelium, honey to isabelline, orange to
while Pl. strelitziae was found on Strelitzia reginae (Strelitzia- umber, cinnamon with centre white due to the aerial mycelium,
ceae) in Italy. greyish sepia to fuscous black, or fuscous grey with margins buff;
Fig. 51. RAxML phylogram obtained from the combined ITS (788 bp), LSU (862 bp), gapdh (694 bp) and tef1 (860 bp) sequences of all the accepted species of Pyrenophora.
Bipolaris panici-miliacei CBS 199.29 and Bipolaris yamadae CBS 202.29 were used as outgroup. The novelties proposed in this study are indicated in bold. RAxML bootstrap
support (BS) values above 70 % and Bayesian posterior probability scores 0.95 are shown in the nodes. GenBank accession numbers were indicated in Table 12 and
Manamgoda et al. (2014). A, ET, LT, SynT and T indicate authentic, ex-epitype, ex-lectotype, ex-syntype and ex-type strains, respectively. TreeBASE: S23834.
MARIN-FELIX ET AL.
Fig. 52. Pyrenophora avenicola (ex-type CBS 307.84). A, B. Conidiophores and conidia. C–I. Conidia. Scale bars: A, B = 10 μm; C–I = 5 μm; C applies to C–I.
reverse olivaceous, olivaceous black with or without margins, with centre fuscous black and margins olivaceous, isabelline with
primrose or luteous, fuscous black with or without margins centre olivaceous, orange to umber or greyish sepia to fuscous
transparent or buff, honey to isabelline, sienna to umber with black with margins transparent, olivaceous grey to olivaceous
margins luteous, or cinnamon with centre brick to dark brick. On black with margins transparent, or fuscous black with margins
MEA white to pale greenish glaucous or buff, greyish sepia to brick, or transparent with centre brick to dark brick.
pale mouse grey or mouse grey, smoke-grey to pale olivaceous
grey, glaucous grey to greenish grey, pale vinaceous to vina- Optimal media and cultivation conditions: On PDA, PNA, OA and
ceous buff, purplish grey with margins vinaceous buff, luteous MEA to induce sporulation of the asexual morph, while for the
with margins white to pale smoke-grey, vinaceous buff to hazel sexual morph Sach's agar with sterilised rice or wheat straw at
with margins white and saffron, pale greenish grey to greenish 25 °C is used.
grey, or fuscous black with margins luteous; reverse olivaceous Distribution: Worldwide, mainly in Australia, Europe, New Zea-
black, smoke-grey to olivaceous grey with middle white due to
land and North America.
the aerial mycelium, fuscous black with or without margins
luteous or olivaceous or buff to cinnamon or honey to isabelline, Hosts: Wide host range, occurring as pathogens, saprobes or
chestnut with margins luteous to rust, blood colour with margins endophytes. Mainly found in members of Poaceae, being path-
luteous or scarlet, chestnut with margins luteous, or orange to ogens of cereals and grasses, including barley, oats and wheat.
sienna. On OA smoke-grey, olivaceous, olivaceous black, hazel, The most common genera belonging to Poaceae which this
buff, cinnamon, olivaceous grey with margins luteous, greyish genus is associated to are Agropyron, Agrostis, Avena, Bromus,
sepia to fuscous black with margins brick, or orange to umber Dactylis, Festuca, Hordeum, Lolium, Poa and Triticum, among
with margins transparent; reverse olivaceous to olivaceous others. Pyrenophora species are also reported from other genera
black, smoke-grey to olivaceous grey, olivaceous black with outside this family, such as Protea and Leucospermum in the
margins transparent or luteous, leaden grey to leaden black, buff Proteaceae.
68
Fig. 53. Pyrenophora cynosuri (ex-type CBS 127918). A–C. Conidiophores and conidia. D. Conidium forming secondary conidium. E–M. Conidia. Scale bars: 10 μm; E applies
to E–M.
Disease symptoms: Leaf spots, leaf blight, leaf blotch, net blotch, Ariyawansa et al. (2014). However, there is still a large number of
melting out, head rot, foot rot, seed-borne diseases, among species which await treatment. Three new combinations are
others. introduced here, i.e. Py. nisikadoi, Py. poae and Py. wirrega-
nensis. The main problem encountered is the lack of type ma-
Notes: Pyrenophora is characterised by immersed to semi-
terial of the already known species, and this resulted in few
immersed ascomata with necks covered with brown to reddish
molecular studies being performed in the past.
brown setae, lack of pseudoparaphyses, clavate to saccate asci,
usually with a large apical ring, and muriform terete (cylindrical, Species delimitation in Pyrenophora based on morphology
frequently circular in section but narrowing to one end) asco- alone is complicated since many species have overlapping
spores (Ariyawansa et al. 2014). The asexual morph was known characters, similar to what is observed in Bipolaris and Curvu-
as Drechslera and it is characterised by brown, transversely laria (Marin-Felix et al. 2017). Therefore, molecular data (ITS,
septate conidia similar to those found in Bipolaris and Curvularia. gapdh and rpb2) are essential for an accurate identification of
In order to properly delineate these three genera, phylogenetic species of Pyrenophora. In our phylogenetic analysis, 21 species
analyses using sequence data of different loci (i.e. LSU, SSU, are accepted and an additional six are newly described.
ITS, gapdh and rpb2) were performed (Zhang & Berbee 2001, Pyrenophora includes saprobic and plant pathogenic species
Ariyawansa et al. 2014, Manamgoda et al. 2014). The synon- with a worldwide distribution, commonly associated with mem-
ymy of Drechslera with Pyrenophora was recently discussed by bers of the family Poaceae. Some species are serious plant
MARIN-FELIX ET AL.
pathogens, e.g. Py. teres, which is a necrotrophic pathogen thickened, 3–4.5 μm. Chlamydospores, microconidiation and
causing net blotch in barley (Crous et al. 1995, Louw et al. 1995, sexual morph not observed.
Campbell et al. 1999, 2002), and Py. tritici-repentis, which cau-
Culture characteristics: Colonies on MEA covering dish after
ses tan spot of wheat (Lamari & Bernier 1989, Balance et al.
1 wk at 25 °C, with abundant aerial mycelium, umbonate; surface
1996, Abdullah et al. 2017) in all the major wheat growing
fuscous black, margins luteous; reverse smoke-grey to oliva-
areas of the world resulting in 3–50 % yield losses (Lamari &
ceous grey, centre white due to the aerial mycelium. Colonies on
Bernier 1989).
PDA covering dish, with abundant aerial mycelium; surface
References: Sivanesan 1987 (morphology and pathogenicity), smoke-grey to olivaceous grey; reverse fuscous black to black,
Zhang & Berbee 2001, Ariyawansa et al. 2014 (morphology and margins transparent. Colonies on OA covering all dish, with
phylogeny). moderate to abundant aerial mycelium; surface smoke-grey to
olivaceous grey; reverse olivaceous grey to olivaceous black,
Pyrenophora avenicola Y. Marín & Crous, sp. nov. MycoBank margins transparent.
MB829614. Fig. 52.
Typus: Sweden, Uppsala, on seed of Avena sp. (Poaceae),
Etymology: Name refers to the host genus Avena, from which it unknown date, C. Svensson (holotype CBS H-23840, culture
was isolated. ex-type CBS 307.84).
Hyphae hyaline, branched, septate, verrucose, 2–6(–7.5) μm. Notes: Pyrenophora avenicola is closely related to Py. chaeto-
Conidiophores arising in groups, septate, straight or flexuous, mioides. Moreover, both species have been found on the same
sometimes geniculate at upper part, usually cells decrease in host, Avena. However, Py. avenicola can be easily distinguished
size towards apex, sometimes branched, cell walls thicker than by its shorter conidiophores (up to 350 μm in Py. avenicola vs.
those of vegetative hyphae, semi- to macronematous, subhya- 1 mm in Py. chaetomioides) and smaller conidia
line to pale brown, usually paler towards apex, not swollen at the (21.5–71.5 × 9.5–15 μm in Py. avenicola vs.
base, up to 350 μm long, 4.5–7 μm wide. Conidiogenous cells 25–140 × 12–22 μm in Py. chaetomioides) with less septa
verruculose, terminal or intercalary, proliferating sympodially, [1–4(–5) in Py. avenicola vs. 2–9 in Py. chaetomioides]. The
subhyaline to pale brown, subcylindrical to swollen, sexual morph has only been observed in Py. chaetomioides.
14–33.5(–43.5) × 6.5–9.5 μm. Conidia verruculose, straight,
Pyrenophora cynosuri Y. Marín & Crous, sp. nov. MycoBank
middle cells enlarged, cylindrical to obclavate, tapering towards
MB829615. Fig. 53.
apex, pale brown to brown, end cells rarely slightly paler,
1–4(–5)-distoseptate, 21.5–71.5 × 9.5–15 μm, forming sec- Etymology: Name refers to the host genus Cynosurus, from
ondary conidiophores or conidia; hila protuberant, darkened, which it was isolated.
Fig. 54. Pyrenophora nisikadoi (CBS 119213). A, B. Conidiophores and conidia. C. Chlamydospores. D–K. Conidia. Scale bars: A, B = 10 μm; C = 20 μm; D = 5 μm; D applies
to D–K.
70
Hyphae hyaline to pale brown, branched, septate, smooth- Colonies on PDA reaching 60–65 mm diam, with sparse aerial
walled or verrucose, 1.5 – 5 μm. Conidiophores arising in mycelium, flat, margins arachnoid; surface greyish sepia to
groups, septate, straight or flexuous, usually geniculate in fuscous black; reverse fuscous black. Colonies on OA covering
upper part, size of cells rarely decrease towards apex, rarely dish, with moderate aerial mycelium; surface greyish sepia to
branched, cell walls thicker than those of vegetative hyphae, fuscous black, margins brick; reverse fuscous black, margins
macronematous, rarely micronematous, pale brown to brown, brick.
slightly paler towards apex, not swollen at the base, (70 – )
95 –700 × 4.5 – 8 μm. Conidiogenous cells smooth-walled to Typus: New Zealand, on seeds of Cynosurus cristatus (Poa-
slightly verruculose, terminal or intercalary, proliferating sym- ceae), 1975, E.H.C. McKenzie (holotype CBS H-23841, culture
podially, pale brown to brown, subcylindrical to swollen, ex-type CBS 127918 = BRIP 12355 a = NZ 14880).
10 –30( – 37) × 5.5 – 10 μm. Conidia verruculose, mostly
Notes: Pyrenophora cynosuri was isolated from seeds of
curved, middle cells sometimes enlarged, cylindrical to
Cynosurus cristatum, a member of the Poaceae, which in-
obclavate, tapering towards apex, subhyaline to pale brown,
cludes host genera commonly infected by species of Pyr-
end cells rarely paler, 2 – 5-distoseptate, (25 – )
enophora. However, Cynosurus represents a new host genus
28 –80( – 83) × 9 –16.5 μm, forming secondary conidiophores
for Pyrenophora.
or conidia; hila protuberant, darkened, thickened, (3 – )
Pyrenophora cynosuri is closely related and morphologically
3.5 – 6 μm. Chlamydospores, microconidiation and sexual
similar to Py. dictyoides. However, both species differ in the size
morph not observed.
of their conidiophore length (up to 700 μm in Py. cynosuri vs. up
Culture characteristics: Colonies on MEA reaching to 250 μm in Py. dictyoides) and conidial length (up to 83 μm in
55–60 mm diam after 1 wk at 25 °C, with moderate aerial Py. cynosuri vs. up to 250 μm in Py. dictyoides), as well as
mycelium, flat to umbonate, margins arachnoid; surface greyish conidial septation (up to 5 in Py. cynosuri vs. up to 15 in Py.
sepia to mouse grey; reverse blood colour, margins luteous. dictyoides).
Fig. 55. Pyrenophora novozelandica (ex-type CBS 127934). A–D. Conidiophores and conidia. E–M. Conidia. Scale bars: A–D = 10 μm; E = 5 μm; E applies to E–M.
MARIN-FELIX ET AL.
Pyrenophora dictyoides A.R. Paul & Parbery, Trans. Brit. brown to brown, sometimes paler towards apex, rarely swollen at
Mycol. Soc. 51: 708. 1968. the base, 50–330 × 3.5–6.5(–8.5) μm. Conidiogenous cells
Synonyms: Helminthosporium dictyoides Drechsler, J. Agric. smooth-walled to verruculose, terminal or intercalary, prolifer-
Res. 24: 679. 1923. ating sympodially, pale brown to brown, subcylindrical to swollen,
Helminthosporium dictyoides var. dictyoides Drechsler, J. Agric. 7.5–20.5(–25) × 5–7.5 μm. Conidia verruculose, straight or
Res. 24: 679. 1923. curved, middle cells enlarged, cylindrical to obclavate, tapering
Helminthosporium dictyoides f. dictyoides Drechsler, J. Agric. towards apex, pale brown to brown, basal cell sometimes paler,
Res. 24: 679. 1923. less frequently apical cell also paler, (1–)2–4(–5)-distoseptate,
Drechslera dictyoides (Drechsler) Shoemaker, Canad. J. Bot. 37: (15–)17.5–42.5 × 8.5–12 μm, not forming secondary co-
881. 1959. nidiophores or conidia; hila flat, darkened, thickened, 2–4 μm.
Helminthosporium dictyoides f. perenne Braverman & J.H. Chlamydospores immersed in all media tested (MEA, OA and
Graham, Phytopathology 50: 695. 1960. PDA), brown to dark brown, lineally or irregularly disposed,
Drechslera andersenii Scharif, Studies on Graminicolous Spe- verrucose, globose to subglobose, up to 30 μm. Microconidiation
cies of Helminthosporium (Tehran): 29. 1963. (nom. inval., Art. and sexual morph not observed.
36.1).
Culture characteristics: Colonies on MEA reaching
Drechslera andersenii A. Lam, Trans. Brit. Mycol. Soc. 85: 601.
80–90 mm diam after 1 wk at 25 °C, with abundant aerial
1986.
mycelium, cottony, lobate; surface pale greenish grey to greenish
Notes: Isolates belonging to Py. dictyoides (CBS 127933 and grey; reverse olivaceous black. Colonies on PDA covering dish,
DAOM 63666) and Drechslera andersenii (CBS 258.80 and CBS with moderate aerial mycelium, cottony, powdery at margins, flat;
967.87) clustered together in a well-supported clade (100 % BS / surface greenish grey to olivaceous black; reverse olivaceous
1 PP). Moreover, the morphology of the asexual morph is similar black. Colonies on OA covering the dish, with sparse aerial
in both species, differing only in the production of conidia with mycelium, powdery to granular, flat; surface grey olivaceous to
much less tapered apices in D. andersenii (Sivanesan 1987). olivaceous black; reverse leaden grey to leaden black.
Therefore, we reduce these species to synonymy.
Typus: Japan, from Briza minor (Poaceae), Y. Nisikado [epitype
Pyrenophora nisikadoi Y. Marín & Crous, nom. nov. MycoBank designated by Manamgoda et al. (2014) CBS H-7218, culture ex-
MB829616. Fig. 54. epitype CBS 190.29 = MUCL 9613].
Replaced synonym: Helminthosporium brizae Y. Nisik., Ber. Additional materials examined: South Africa, Western Cape Province, J.S.
Ohara Inst. Landw. Biol.: 121, 133. 1928, non Pyrenophora bri- Marais Nature Reserve, from Protea burchellii senescent flowerheads (Protea-
zae C. Massal. ex Sacc. 1911. ceae), 6 Jun. 2000, S. Lee, CBS 119213. New Zealand, Auckland, Waitakere
Additional synonyms: Bipolaris brizae (Y. Nisik.) Shoemaker, Ranges, from Briza minor (Poaceae), 1 Nov. 1975, E.H.C. McKenzie, CBS
Canad. J. Bot. 37: 882. 1959. 127912 = ICMP 6183.
Drechslera brizae (Y. Nisik.) Subram. & B.L. Jain, Curr. Sci. 35: Notes: This species was originally described in Helminthospo-
354. 1966. rium as Hel. brizae (Nisikado 1928), then transferred to Bipolaris
(Shoemaker 1959), and finally placed in Drechslera
Etymology: Named after the Japanese plant pathologist and
(Subramanian & Jain 1966). Type material was not available,
mycologist, Y. Nisikado, who first described and named this
thus Manamgoda et al. (2014) designated CBS 190.29 as ex-
fungus.
epitype since this strain was isolated by the original author
Hyphae hyaline to pale brown, branched, septate, verrucose, from the same host and location. In our phylogenetic analysis
2–5.5 μm. Conidiophores arising in groups, septate, mostly based on ITS, gapdh and rpb2, CBS 190.29 together with other
flexuous, rarely straight, geniculate at upper part, sometimes size two strains were located in a well-supported clade (100 % BS / 1
of cells decrease towards apex, frequently branched, cell walls PP) within the main clade representing the genus Pyrenophora.
thicker than those of vegetative hyphae, macronematous, pale Therefore, we propose to transfer this species to Pyrenophora,
Fig. 56. Pyrenophora pseudoerythrospila (ex-type CBS 127931). A. Protoascomata on OA. B, C. Protoascomata. Scale bars = 10 μm.
72
Fig. 57. Pyrenophora sieglingiae (ex-type CBS 127930). A–C. Sterile ascomata. D. Neck of ascoma. E, F. Conidiophores and conidia. G–O. Conidia. Scale bars: C = 100 μm;
D–F = 20 μm; G = 10 μm; G applies to G–O.
changing the epithet to Py. nisikadoi since Py. brizae already walls thicker than those of vegetative hyphae, macronematous,
exists. pale brown to brown, paler towards apex, not swollen at the
Isolate CBS 119213 sporulated, enabling us to conduct a base, 35–700 × 4.5–7.5 μm. Conidiogenous cells smooth-
morphological comparison and provide a modern description of walled, terminal or intercalary, proliferating sympodially,
this species in PDA. Moreover, Py. nisikadoi was formerly only brown, terminal conidiogenous cells hyaline, cylindrical to
recorded from Briza minor, a member of the Poaceae, while it is subcylindrical, 11–22 × 5–8.5 μm. Conidia smooth-walled,
here also recorded from Protea birchellii, which belongs to the straight, rarely slightly curved, sometimes middle cells
Proteaceae. slightly enlarged, cylindrical to obclavate, tapering towards
apex, pale brown to brown, sometimes basal cell slightly paler,
Pyrenophora novozelandica Y. Marín & Crous, sp. nov.
(2–)3–5(–6)-distoseptate, 20.5–58 × 9.5–14 μm, not forming
secondary conidiophores or conidia; hila usually inconspic-
Etymology: Name refers to New Zealand, the country from where uous, flat, slightly darkened, slightly thickened, 2–4 μm.
it was isolated. Chlamydospores, microconidiation and sexual morph not
observed.
Hyphae hyaline to pale brown, branched, septate, verrucose,
1–6( –8.5) μm. Conidiophores arising in groups, septate, Culture characteristics: Colonies on MEA reaching
straight or flexuous, sometimes geniculate at upper part, size 27–30 mm diam after 1 wk at 25 °C, with sparse aerial
of cells not decreasing towards apex, rarely branched, cell mycelium, raised, margins fringed; surface grey olivaceous;
MARIN-FELIX ET AL.
reverse dark mouse grey, margins buff. Colonies on PDA margins fringed; surface grey olivaceous; reverse olivaceous
reaching 38–40 mm diam, with sparse aerial mycelium, flat, black.
margins fringed; surface smoke grey to olivaceous grey;
reverse olivaceous black. Colonies on OA reaching Typus: New Zealand, Wanganui, Palmerston North, Seed
37–40 mm diam, with moderate aerial mycelium, flat, Testing Station, on seed of Triticum sp. (Poaceae), 5 Oct. 1976,
Fig. 58. Pyenophora variabilis (ex-type CBS 127920). A–C. Conidiophores and conidia. D–N. Conidia. Scale bars = 10 μm; C applies to A–C; N applies to D–N.
74
G.F. Laundon (holotype CBS H-23843, culture ex-type CBS Drechslera poae (Baudys) Shoemaker, Canad. J. Bot. 40: 827.
127934 = LEV 11079b = PDD 50697). 1962.
Notes: Pyrenophora novozelandica is similar and closely related Description and illustration: Sivanesan (1987).
to Py. fugax. However, both species can be easily distinguished Materials examined: Canada, Saskatchewan, Saskatoon, from Poa pratensis
based on the size of their conidiophores (up to 250 μm in Py. (Poaceae), Oct. 1973, J.D. Smith, DAOMC 145373. Germany, Husum, from
fugax vs. up to 700 μm in Py. novozelandica) and conidia P. pratensis, Aug. 1966, U.G. Schlösser, CBS 319.68. USA, Maryland, Beltsville,
(50–170 × 14–24 μm in Py. fugax vs. 20.5–58 × 9.5–14 μm in from P. pratensis, Apr. 1979, A. Hagan, CBS 128045 = BRIP 12969a.
Py. novozelandica), as well as conidial septation [4–8(–10) in Notes: Pyrenophora poae was introduced as Helminthosporium
Py. fugax vs. (2–)3–5(–6) in Py. novozelandica]. Pyrenophora poae by Baudys (1916), then transferred to Drechslera
novozelandica is known to occur on Triticum in New Zealand, (Shoemaker 1959). The original description was based on a
which is a common host of species belonging to Pyrenophora, specimen isolated from Poa trivialis in the Czech Republic. Type
including Py. fugax. material is not available, but CBS 319.68 is considered here as
an authentic strain since was isolated from the same host genus
Pyrenophora poae (Baudys) Y. Marín & Crous, comb. nov.
and continent. Unfortunately, it did not sporulate and thus we
MycoBank MB829617.
chose to not designate it as epitype. All the strains identified as
Basionym: Helminthosporium poae Baudys, Lotos 63: 104. 1916.
D. poae were located in the main clade belonging to Pyr-
Synonyms: Helminthosporium vagans Drechsler, J. Agric. Res.,
enophora, and a new combination is proposed here.
Washington 24: 688. 1923.
Drechslera vagans (Drechsler) Shoemaker, Canad. J. Bot. 37: Pyrenophora pseudoerythrospila Y. Marín & Crous, sp. nov.
881. 1959. MycoBank MB829675. Fig. 56.
Fig. 59. Ramichloridium spp. A. Ramichloridium luteum on apple. B. Sporulating colonies of Ramichloridium luteum (ex-type CBS 132088) on PDA. C–G. Macronematous
conidiophores with sympodially proliferating conidiogenous cells, which give rise to a conidium-bearing rachis with crowded and prominent scars. C. Ramichloridium apiculatum
(ex-type CBS 156.59). D. Ramichloridium cucurbitae (ex-type CBS 132087). E, F. Ramichloridium luteum (ex-type CBS 132088). G. Ramichloridium punctatum (ex-type CBS
132090). H, I. Scanning electron micrographs of Ramichloridium luteum (ex-type CBS 132088) showing sympodial proliferation with scars on conidiogenous cells. J, K.
Conidiophores reduced to conidiogenous cells. J. Ramichloridium cucurbitae (ex-type CBS 132087). K. Ramichloridium luteum (ex-type CBS 132088). L, M. Conidia. L.
Ramichloridium apiculatum (ex-type CBS 156.59). M. Ramichloridium punctatum (ex-type CBS 132090). Scale bars: H = 2 μm; I = 1 μm; all others = 10 μm. Pictures C, L taken
from Li et al. (2012); all others from Arzanlou et al. (2007).
MARIN-FELIX ET AL.
Etymology: Named after its close phylogenetic relation to Py. Agrostis spp. in Australia and North America, but has also been
erythrospila. reported on Lolium in Germany (Farr & Rossman 2019), having
the same host and distribution as Py. pseudoerythrospila.
Only forming protoascomata in OA. Protoascomata composed of
pale brown to brown cells of up to 100 μm diam, of textura Pyrenophora sieglingiae Y. Marín & Crous, sp. nov. MycoBank
angularis to textura globulosa. Pyrenophora pseudoerythrospila MB829618. Fig. 57.
differs from its closest phylogenetic neighbour, Py. erythrospila
Etymology: Name refers to Sieglingia, the host genus from which
by unique fixed alleles in three loci based on alignments of the
this fungus was collected.
separate loci deposited in TreeBASE (S23834): LSU positions 70
(G), 395 (C), 396 (T), 397 (T), 500 (G), 536 (C), 537 (T); ITS Hyphae hyaline to pale brown, branched, septate, verru-
positions 141 (A), 146 (A), 147 (T), 148 (A), 149 (G), 152 (G), cose, (1 – )1.5 – 5.5 μm. Sterile ascomata solitary or arising
153 (A), 154 (G), 155 (T), 164 (T), 168 (T), 169 (G), 173 (G), 174 in groups, brown to dark brown or black, sometimes apical
(C), 176 (A), 178 (T), 179 (G), 184 (T), 187 (T), 189 (T), 197 (C), part of neck yellowish brown, composed of cells of textura
198 (C), 199 (C), 205 (C), 206 (T), 208 (C), 223 (T), 224 (T), 225 intricata, up to 1200 μm long, up to 300 μm wide, co-
(T), 226 (T), 234 (A), 252 (T), 264 (T), 268 (C), 279 (G), 288 (C), nidiophores arising from the body and neck; inside con-
289 (A), 293 (A), 302–304 (indels), 327 (T), 330–332 and 341 sisting of angular to globose, hyaline cells. Conidiophores
(indels),511 (T), 536–540 (indels), 558 (C), 591 (A), 594 (G), 601 arising in groups, septate, straight or flexuous, rarely
(T), 621 (A), 630 (C), 641 (G), 643 (A), 644 (G), 645 and 646 geniculate in the upper part, cell size rarely decreases
(indels), 659 (T), 660 (G), 664 (T), 666 (A), 667 (T); gapdh towards the apex, unbranched, cell walls thicker than those
positions 30 (C), 49 (A), 54 and 55 (indels), 59 (G), 60 (A), 69 of vegetative hyphae, macronematous, brown, mostly paler
(C), 71 (C), 74 (A), 85 (T), 87 (A), 177 (G), 178 (G), 179 (C), 180 towards apex, not swollen at the base, 100 – 700 × (5 – )
(C), 182 (C), 187 (G), 189 (C), 190 (T), 191 (A), 192 (T), 193 (C), 7 – 9( – 11) μm. Conidiogenous cells verruculose, terminal or
194 (A), 195 (G), 196 (A), 197 (C), 201 (G), 203 (A), 204 (G), 205 intercalary, proliferating sympodially, pale brown to brown,
(A), 256 (G), 319 (G), 373 (T), 406 (C), 487 (T), 493 (T), 496 (T), subcylindrical to slightly swollen, (12 – )
523 (G), 526 (C), 532 (C), 538 (A). 15 – 33.5( – 36.5) × 9 – 12( – 13.5) μm. Conidia verruculose,
Culture characteristics: Colonies on MEA reaching straight or curved, sometimes with middle cells enlarged,
55–60 mm diam after 1 wk at 25 °C, with sparse aerial cylindrical to obclavate, tapering towards apex, pale brown
mycelium, flat, margins fringed; surface pale vinaceous to to brown, end cells usually paler, 4 – 8-distoseptate,
vinaceous buff; reverse orange to sienna, margins transparent. 56 – 108( – 120) × 15 – 23( – 25.5) μm, forming secondary
Colonies on PDA reaching 56–59 mm diam, with sparse aerial conidiophores or conidia; hila not protuberant or flat,
mycelium, flat, margins fringed; surface cinnamon, white darkened, thickened, (4– )4.5 – 6.5( – 7) μm. Chlamydo-
mycelium in the centre; reverse cinnamon, centre brick to dark spores, microconidiation and sexual morph not observed.
brick. Colonies on OA reaching 50–55 mm diam, without aerial Culture characteristics: Colonies on MEA reaching 26 – 29
mycelium, flat; surface cinnamon; reverse transparent, centre diam after 1 wk at 25 °C, with abundant aerial mycelium,
brick to dark brick. raised, slightly lobate; surface white to buff; reverse fuscous
Typus: Germany, West Germany, on Lolium sp. (Poaceae), 9 black. Colonies on PDA reaching 27 – 30 diam, with abundant
Sep. 1968, U.G. Schlosser (holotype CBS H-23844, culture ex- aerial mycelium, lobate; surface olivaceous black with patches
type CBS 127931 = DAOMC 126772). of white due to aerial mycelium; reverse olivaceous black,
margins luteous. Colonies on OA reaching 54– 57 diam, with
Notes: The ex-type strain of Pyrenophora pseudoerythrospila did moderate to abundant aerial mycelium, flat; surface smoke-
not sporulate on any of the media tested, producing only few grey to olivaceous grey; reverse smoke-grey to olivaceous
protoascomata in OA. However, these remained sterile after grey.
several months of incubation. Pyrenophora pseudoerythrospila is
closely related to Py. erythrospila, which produces both sexual Typus: New Zealand, Auckland, Waikumete, from leaf of Sie-
and asexual morphs. The protoascomata were also reported in glingia decumbens (Poaceae), E.H.C. McKenzie (holotype CBS
Py. erythrospila, but these finally developed mature ascospores H-23842, culture ex-type CBS 127930 = ICMP 6170 = PDDCC
after 25 wk. Pyrenophora erythrospila is commonly found on 6170).
Table 13. DNA barcodes of accepted Ramichloridium spp.

ITS LSU rpb2 tef1
Ramichloridium apiculatum CBS 156.59T EU041791 EU041848 MF951416 – Arzanlou et al. (2007), Videira et al. (2017)
R. cucurbitae CBS 132087T JQ622087 JQ622095 – JQ622112 Li et al. (2012)
R. luteum CBS 132088T EU329730 JQ622099 MF951417 JQ622116 Li et al. (2012), Videira et al. (2017)
R. malus LQ73T EF627452 – – – Zhang et al. (2007)
R. punctatum CBS 132090 T
JQ622086 JQ622094 – JQ622111 Li et al. (2012)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; LQ: collection not specified in the work in which it was introduced. T indicates ex-type strains.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S)nrDNA; rpb2: partial DNA-directed RNA polymerase II second largest
subunit gene; tef1: partial elongation factor 1-alpha gene.
76
Notes: Pyrenophora sieglingiae is closely related to Py. seme- Moreover, Py. semeniperda produces conidia with more septa
niperda and Py. wirreganensis. Morphologically, these species (up to 12 in Py. semeniperda vs. up to 8 in Py. sieglingiae vs. up
are similar, producing sterile ascomata with long necks. How- to 9 in Py. wirreganensis), and it is the only one that produces a
ever, they can be distinguished by the size of their conidiophores sexual morph. Pyrenophora sieglingiae has been isolated from
(up to 700 μm in Py. sieglingiae vs. up to 180 μm in Py. seme- Sieglingia from New Zealand while Py. wirreganensis occurs on
niperda vs. up to 1 000 μm in Py. wirreganensis) and conidia Hordeum in Australia. Pyrenophora semeniperda has been
[56–108(–120) × 15–23(–25.5) μm in Py. sieglingiae vs. isolated from both hosts in both locations, apart from other hosts
70–160 × 13–17 μm in Py. semeniperda vs. (30–) that are widely distributed, i.e. Agropyron, Avena, Bromus,
40–80(–100) × (10–)12–19(–22) μm in Py. wirreganensis). Cortaderia, Ehrhartia, Pennisetum and Triticum.
Fig. 60. RAxML phylogram obtained from the combined ITS (594 bp), LSU (761 bp), rpb2 (819 bp) and tef1 (470 bp) sequence alignment of all accepted species of
Ramichloridium and related taxa. The tree was rooted to Parapenidiella tasmaniensis CBS 124991 and Stenella araguata CBS 105.75. The novelties proposed in this study are
indicated in bold. RAxML bootstrap support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession
numbers are listed in Table 13, Li et al. (2012), Videira et al. (2017) and Chen & Kirschner (2018). T and NT indicate ex-type and ex-neotype strains, respectively. TreeBASE:
S23834.
MARIN-FELIX ET AL.
Pyrenophora teres Drechsler, J. Agric. Res., Washington 24: Helminthosporium tuberosum G.F. Atk., Bulletin of Cornell Uni-
656. 1923. versity 3: 47. 1897.
Synonyms: Helminthosporium secalis Fee, Mem. Soc. Mus. Hist. Drechslera tuberosa (G.F. Atk.) Shoemaker, Canad. J. Bot. 37:
Nat. Strassbourg 3: 36. 1843. 881. 1959.
Alternaria secalis (Fee) Sacc. & Traverso, Syll. fung. (Abellini) Helminthosporium fragosoi Bubak, Hedwigia 57: 13. 1915.
20: 1184. 1911.
Pyrenophora teres f. teres Drechsler, J. Agric. Res., Washington
Helminthosporium gramineum Rabenh., Klotzschii Herb. Viv.
24: 656. 1923.
Mycol., Edn Nov, Ser. Sec., Cent. 4: no. 332. 1857.
Pyrenophora teres subsp. teres Drechsler, J. Agric. Res.,
Brachysporium gracile var. gramineum (Rabenh. ex Schltdl.)
Washington 24: 656. 1923.
Sacc., Syll. fung. (Abellini) 4: 430. 1886.
Drechslera graminea (Rabenh. ex Schltdl.) S. Ito, Proc. Imper. Helminthosporium japonicum S. Ito & Kurib., Proc. Imper. Acad.
Acad. Tokyo 6: 355. 1930. Tokyo 6: 353. 1930.
Drechslera teres subsp. graminea (Rabenh. ex Schltdl.) Simay, Pyrenophora japonica S. Ito & Kurib., Proc. Imper. Acad. Tokyo
Barley Newsletter 36: 174. 1992. 6: 353. 1930.
Helminthosporium teres Sacc., Syll. fung. (Abellini) 4: 412. 1886. Drechslera japonica (S. Ito & Kurib.) Shoemaker, Canad. J. Bot.
Drechslera teres (Sacc.) Shoemaker, Can. J. Bot. 37: 881. 1959. 37: 881. 1959.
Drechslera teres f. teres (Sacc.) Shoemaker, Canad. J. Bot. 37: Pyrenophora graminea S. Ito & Kurib., Proc. Imp. Acad. Japan 6:
881. 1959. 353. 1930.
Drechslera teres subsp. teres (Sacc.) Shoemaker, Canad. J. Bot. Pyrenophora teres subsp. graminea (S. Ito & Kurib.) Simay,
37: 881. 1959. Barley Newsletter 36: 174. 1992.
Helminthosporium hordei Eidam, Der Landw. (Schles. Landw. Pyrenophora secalis M.D. Whitehead & J. Dicks., Mycologia 44:
Ztg), Breslau 27: 509. 1891. 752. 1952.
Fig. 61. Seifertia azalae. A, B. Disease symptoms caused on Rhododendron. C–G. Synnemata. H, I. Conidiogenous cells and conidia. J, K. Conidia. Scale bars: F = 100 μm;
G = 50 μm; H–K = 10 μm.
78
Drechslera teres f. maculata Smed.-Pet., Arb. Tiflis Bot. Gard.: cell walls thicker than those of vegetative hyphae, semi- to
139. 1971. macronematous, brown, not swollen at the base, up to 321 μm
Pyrenophora teres f. maculata Smed.-Pet., The Royal Veterinary long, 5–10 μm wide. Conidiogenous cells smooth-walled, ter-
and Agricultural University Yearbook: 139. 1971. minal or intercalary, proliferating sympodially, brown, sub-
cylindrical, 18–27 × 8–10 μm. Conidia smooth-walled, straight to
Description and illustrations: Sivanesan (1987).
curved, cylindrical, subcylindrical, obclavate, obpyriform to sub-
Materials examined: Denmark, from Hordeum vulgare (Poaceae), unknown date, globose, pale brown to brown, 1–3-distoseptate,
V. Smedegaard-Petersen (ex-type culture of Py. teres f. maculata CBS 228.76). 20–75 × 13–19.5 μm; hila flat, darkened, thickened, 4–7 μm.
Japan, from H. vulgare (Poaceae), unknown date, S. Ito, CBS 281.31; unkown
substrate and date, S. Ito, CBS 282.31; from H. vulgare (Poaceae), unknown
Chlamydospores and sexual morph not observed.
date, Y. Nisikado, CBS 336.29 = MUCL 9687. Germany, Niedersachsen, Typus: Canada, British Columbia, Agassiz Research Station,
Rotenburg, from H. vulgare (Poaceae), Jul. 1968, U.G. Schlösser, CBS 314.69.
Hungary, Eszteragpuszta, from Hordeum vulgare leaf (Poaceae), unknown date,
from leaves of Poa trivialis (Poaceae), Jul. 1972, J.D. Smith
col. M. Csosz, dep. J. Bakonyi, CBS 123929; Taplanszentkereszt, from Hordeum (holotype CBS H-23843, culture ex-type CBS
vulgare leaf (Poaceae), unknown date, col. A. Tomcsanyi, dep. J. Bakonyi, CBS 127920 = DAOMC 139513).
123932.
Notes: Pyrenophora variabilis was located on an independent
Notes: In our phylogenetic analysis, isolates identified as Py. branch far removed from the other species in the genus. It can be
teres and Py. graminea were located in the same well-supported easily distinguished from all the species of the genus by its highly
clade (100 % BS / 1 PP), suggesting that these represent the variable conidial morphology in size and shape, from cylindrical,
same species. In fact, Py. graminea has been recently consid- subcylindrical or obclavate to subglobose or obpyriform. It was
ered as a subspecies of Py. teres (Simay 1992) since its isolated from Poa trivialis (Poaceae) leaves in Canada, a com-
morphology is similar and both species share the same host, mon host of Pyrenophora spp.
Hordeum. Therefore, we propose the synonymy of both species
under the name of Py. teres, which is well established and the Pyrenophora wirreganensis (Wallwork et al.) Y. Marín & Crous,
comb. nov. MycoBank MB829621.
most commonly used name for this taxon. Moreover, an
Basionym: Drechslera wirreganensis Wallwork et al., Mycol. Res.
authentic strain of Py. japonica was also located in this clade,
96. 888. 1992.
supporting the synonymy of Py. japonica with Py. teres proposed
by Crous et al. (1995), which was based on their morphological, Description and illustration: Wallwork et al. (1992).
molecular and pathological similarity.
Material examined: Australia, South Australia, from Hordeum sp. (Poaceae),
Pyrenophora teres produces net blotch on barley worldwide, unknown date, J. Bakonyi, CBS 109896.
causing cell death and feeding off the nutrients released
(Sivanesan 1987, Louw et al. 1995, Campbell et al. 1999, Notes: Pyrenophora wirreganensis was introduced as Drech-
Ellwood et al. 2012). Two different forms of Py. teres were rec- slera wirreganensis by Wallwork et al. (1992) to accommodate a
ognised depending on the disease symptoms produced, i.e. Py. specimen isolated from Hordeum in Australia. In the phylogenetic
teres f. teres producing the net form of net blotch, characterised analysis, the strain CBS 109896 identified as Py. wirreganensis
by elongated lesions where necrosis develops along leaf veins and isolated from the same host and location than the ex-type
with occasional transverse striations, while Py. teres f. maculata strain (IMI 348323), was located in an independent branch
produces the spot form of net blotch, typified by more ovoid within the main clade representing the genus Pyrenophora.
lesions, often surrounded by a chlorotic zone (Campbell et al. Therefore, this species is here transferred to the latter genus. For
1999, Ellwood et al. 2012). However, both forms are consid- comparison with close species see notes of Py. sieglingiae.
ered the same species, Py. teres. This disease becomes sys- Authors: Y. Marin-Felix, M. Hernandez-Restrepo, P.W. Crous
temic in plants infected from seed (Sivanesan 1987).
Ramichloridium Stahel ex de Hoog, Stud. Mycol. 15: 59. 1977.
Pyrenophora variabilis Hern.-Restr. & Y. Marín, sp. nov. Fig. 59.
Classification: Dothideomycetes, Dothideomycetidae, Capno-
Etymology: Name refers to the highly variable conidial diales, Dissoconiaceae.
morphology.
Type species: Ramichloridium apiculatum (J.H. Mill. et al.) de
Hyphae hyaline to brown, branched, septate, smooth-walled to Hoog, basionym: Chloridium apiculatum J.H. Mill. et al. Ex-type
verrucose, 2.5–7 μm. Conidiophores arising in groups, septate, strain: CBS 156.59 = ATCC 13211 = IMI 100716 = JCM
straight or flexuous, sometimes geniculate in upper part, simple, 6972 = MUCL 15753 = MUCL 7991 = QM 7716.
Table 14. DNA barcodes of accepted Seifertia spp.

LSU ITS tef1
Seifertia azaleae DAOM 239136 EU030276 – – Seifert et al. (2007)
CPC 35017 MK540034 MK539964 MK540166 Present study
Sei. shangrilaensis MFLUCC 16-0238T KU954100 – KU954101 Li et al. (2016b)
1
CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute; DAOM: Plant Research Institute, Department of Agriculture (Mycology),
Ottawa, Canada; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand. T indicates ex-type strains.
2
LSU: partial large subunit (28S) nrRNA gene; ITS: internal transcribed spacers and intervening 5.8S nrDNA; tef1: partial translation elongation factor 1-alpha gene.
MARIN-FELIX ET AL.
80
DNA barcode (genus): LSU. present study, only five species have been demonstrated to
belong to Ramichloridium, which is characterised by aseptate,
DNA barcodes (species): ITS, rpb2, tef1. Table 13. Fig. 60.
pale brown, smooth-walled to finely verrucose, clavate or
Mycelium consisting of hyaline, subhyaline, pale brown, or oblong to ellipsoid, or obovate to obconical conidia. Based on
brown, smooth-walled, septate, branched, submerged and molecular data, R. indicum is proposed here as a new genus,
aerial hyphae. Conidiophores straight to gently curved, Globoramichloridium indicum, and R. ducassei as a new
subcylindrical, unbranched, smooth- and thick-walled, brown combination in Zasmidium (see Zasmidium below). Moreover,
to dark brown, 0 – 3-septate, or sometimes reduced to R. apiculatum, R. cucurbitae and R. mali were located in a
intercalary conidiogenous cells. Conidiogenous cells inte- well-supported clade (100 % BS / 1 PP) without phylogenetic
grated, terminal, pale to medium brown, or golden-brown, distance. Unfortunately, the only loci available and common in
apical part subhyaline to pale brown, subcylindrical, rarely the three species are the ITS and LSU. Therefore, other loci
somewhat clavate, sometimes with irregular, nodulose should be sequenced to verify their status as separate species.
swellings along the length of the conidiogenous cells, Species of Ramichloridium cause sooty blotch and flyspeck
tapering towards apex, with sympodial proliferation, forming disease (SBFS) on members of the family Rosaceae, which
a rachis with slightly thickened and darkened, circular, produces blemishes on the epicuticular wax layer and is regar-
somewhat protruding scars. Conidia solitary, aseptate, pale ded as an economically serious disease (Wang et al. 2014).
brown, smooth-walled to finely verrucose, clavate or oblong Ramichloridium cucurbitae and R. punctatum have been found
to ellipsoid, or obovate to obconical, apex obtuse or sub- as SBFS pathogens only in the USA (Li et al. 2012), while
obtuse, base truncate, with a conspicuous, slightly dark- R. apiculatum, R. luteum and R. mali have been reported as
ened and thickened, not refractive hilum. causal agents of SBFS in China (Zhang et al. 2007, Li et al.
2012, Wang et al. 2014).
Culture characteristics: Colonies after 1 mo at 25 °C in the
dark flat, spreading, with sparse to moderate aerial mycelium, References: Arzanlou et al. 2007, Li et al. 2012 (morphology and
margins smooth and even, or lobate and feathery. On SNA phylogeny).
grey olivaceous, pale mouse grey, or smoke-grey. On PDA
olivaceous grey or smoke-grey; reverse olivaceous grey or Globoramichloridium Y. Marín & Crous, gen. nov. MycoBank
iron-grey. On OA olivaceous grey or iron-grey; reverse iron- MB829622.
grey, with pale luteous pigment diffusing into agar. On MEA Etymology: Name reflects the characteristic globose conidia
olivaceous green; reverse olivaceous black, often with a produced by this genus.
diffusing citron-yellow pigment.
Illustration: Arzanlou et al. (2007).
Optimal media and cultivation conditions: MEA, OA, PDA and
SNA at 25 °C under near-ultraviolet light. Mycelium consisting of submerged and aerial hyphae; sub-
merged hyphae smooth- and thin-walled, hyaline, with thin septa;
Distribution: Africa, America and Asia. aerial hyphae coarsely verrucose, olivaceous green, rather thick-
Hosts: Aloe sp. (Aloaceae), Cucumis sativus, Cucurbita maxima walled, with thin septa. Conidiophores arising vertically from
(Cucurbitaceae), Malus domestica, Malus pumila and Pyrus creeping hyphae at right angles, straight, unbranched, thick-
pyrifolia (Rosaceae). Also isolated from soil. walled, smooth-walled, dark brown, with up to 10 thin septa,
often with inflated basal cells. Conidiogenous cells terminally
Disease symptoms: Sooty blotch and flyspeck diseases. integrated, smooth-walled, dark brown, sympodially proliferating,
Notes: Ramichloridium was initially introduced by Stahel rachis straight or flexuose, geniculate or nodose, subhyaline;
(1937), designating R. musae as type species. However, the scars thickened and darkened, clustered at nodes. Microcyclic
name was invalid due to the lack of Latin description or conidiation observed in culture. Conidia solitary, (0–)1-septate,
diagnosis. Subsequently, de Hoog (1977) validated this genus not constricted at the septum, subhyaline to pale brown, smooth-
to include species with erect, dark conidiophores and pre- walled or coarsely verrucose, rather thin-walled, broadly ellip-
dominantly aseptate conidia, designating as new type species soidal to globose, with truncate base; hila conspicuous, slightly
R. apiculatum. In that study, 13 species were recognised in darkened, not thickened.
Ramichloridium, and subsequently more species were Culture characteristics: Colonies on MEA reaching 35 mm diam
included. However, several molecular studies demonstrated after 2 wks at 24 °C. Colonies velvety, rather compact, slightly
that some of them belonged to different genera, i.e. Myrme- elevated, with entire, smooth, whitish margins, dark olivaceous
cridium, Pachyramichloridium, Pleurothecium, Radulidium, green in the central part.
Rhinocladiella and Zasmidium, and were subsequently trans-
ferred (Arzanlou et al. 2007, Cheewangkoon et al. 2009, Type species: Globoramichloridium indicum (Subram.) Y. Marín
Videira et al. 2017). Some species still need to be molecu- & Crous. Holotype: IMI 114625. Representative strain: CBS
larly studied to confirm their phylogenetic position. In the 171.96.
Fig. 62. Seiridium spp. A–F. Disease symptoms on Cupressaceae hosts. A–C. Flagging of branches. D. Trunk canker with gummosis. E. Branch canker. F. Conidiomata. G–I.
Seiridium pezizoides (CBS 145115). G, H. Acervuli on Cupressaceae sp. I. Conidial masses on artificial media. J, K. Conidiophores and conidiogenous cells. J. Seiridium
neocupressi (CBS 142625). K. Seiridium eucalypti (CBS 343.97). L–R. Conidia. L. Seiridium cardinale (CBS 909.85). M. Seiridium spyridicola (CBS 142628). N. Seiridium
unicorne (CBS 538.82). O. Seiridium neocupressi (CBS 142625). P. Seiridium eucalypti (CBS 343.97). Q. Seiridium kartense (CBS 142629). R. Seiridium pezizoides (CBS
145115). S–U. Seiridium cupressi (IMI 40096). S, T. Ascomata. U. Ascospores. Scale bars: F = 2 mm; G, H = 50 μm; J–U = 10 μm. Pictures J–U taken from Bonthond et al.
(2018).
MARIN-FELIX ET AL.
Table 15. DNA barcodes of accepted Seiridium spp.

Species Isolates.1 GenBank accession numbers2 References
ITS rpb2 tef1 tub2
Seiridium camelliae MFLUCC 12-0647T JQ683725 – JQ683741 JQ683709 Maharachchikumbura et al. (2015)
S. cancrinum CBS 226.55T LT853089 LT853137 LT853186 LT853236 Bonthond et al. (2018)
R
S. cardinale CBS 909.85 LT853064 LT853113 LT853161 LT853211 Bonthond et al. (2018)
S. ceratosporum PHSI2001Pathcw07 AY687314 – – DQ534043 Liu et al. (2007)
S. cupressi CBS 224.55ET LT853083 LT853131 LT853180 LT853230 Bonthond et al. (2018)
S. eucalypti CBS 343.97ET LT853099 LT853146 LT853196 LT853246 Bonthond et al. (2018)
T
S. kartense CBS 142629 LT853100 LT853147 LT853197 LT853247 Bonthond et al. (2018)
S. kenyanum CBS 228.55T LT853098 LT853145 LT853195 LT853245 Bonthond et al. (2018)
GT
S. marginatum CBS 140403 KT949914 LT853149 LT853199 LT853249 Jaklitsch et al. (2016), Bonthond et al. (2018)
S. neocupressi CBS 142625T LT853079 LT853127 LT853176 LT853226 Bonthond et al. (2018)
T
S. papillatum CBS 340.97 LT853102 LT853150 LT853200 LT853250 Bonthond et al. (2018)
S. persooniae CBS 143445T MG386033 – – MG386163 Crous et al. (2017b)
S. pezizoides CBS 145115 MK079342 MK058475 MK058480 MK058485 Present study
S. phylicae CBS 133587T LT853091 LT853139 LT853188 LT853238 Bonthond et al. (2018)
T
S. podocarpi CBS 137995 LT853101 LT853148 LT853198 LT853248 Bonthond et al. (2018)
S. pseudocardinale MFLUCC 13-0525T KU848210 – – – Wijayawardene et al. (2016)
S. rosarum MFLUCC 17-0654T MG828961 – – – Wanasinghe et al. (2018)
S. spyridicola CBS 142628T LT853095 LT853142 LT853192 LT853242 Bonthond et al. (2018)
S. unicorne CBS 143871ET MK079339 MK058477 MK058482 MK058487 Present study
CBS 538.82R LT853088 LT853136 LT853185 LT853235 Bonthond et al. (2018)
S. venetum MFLU 15-0369 R
KT438836 – – KT438837 Maharachchikumbura et al. (2015)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLU and MFLUCC: Herbarium and culture collection of Mae Fah Luang University, Chiang Rai,
Thailand, respectively; PHSI: from Liu et al. (2007). T, ET, GT and R indicate type or ex-type, ex-epitype, ex-generic type and reference strains, respectively.
2
elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
Notes: This genus is introduced to accommodate R. indicum, Notes: The strain examined and included in the phylogenetic
which differs from Ramichloridium spp. by its broadly ellipsoidal analysis, CBS 171.96, was not derived from type material.
to globose, mostly 1-septate, smooth-walled or coarsely verru- However, the morphology of this strain fits perfectly with the
cose conidia, being clavate or oblong to ellipsoid, or obovate to morphology of the holotype IMI 114625 (de Hoog 1977).
obconical, aseptate, smooth-walled to finely verrucose in Ram- Therefore, CBS 171.96 is considered here as a representative
ichloridium. This genus is related to Dissoconium, but the latter strain, and we propose the new genus and combination based on
can be easily distinguished by its percurrent and sympodial the phylogenetic data derived from this isolate, as well as on the
proliferation, and the ellipsoid to obclavate, smooth-walled morphological differences observed.
conidia.
Authors: Y. Marin-Felix, J.Z. Groenewald & P.W. Crous
Globoramichloridium indicum (Subram.) Y. Marín & Crous,
Seifertia Partr. & Morgan-Jones, Mycotaxon 83: 348. 2002.
Fig. 61.
Basionym: Chloridium indicum Subram., Proc. Indian Acad. Sci.,
Sect. B 42: 286. 1955. Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
Synonyms: Veronaea verrucosa Geeson, Trans. Brit. Mycol. sporales, Melanommataceae.
Soc. 64: 349. 1975.
Type species: Seifertia azaleae (Peck) Partr. & Morgan-Jones,
Veronaea indica (Subram.) M.B. Ellis, in Ellis, More Dematia-
basionym: Periconia azaleae Peck. Representative strain:
ceous Hyphomycetes: 209. 1976.
DAOM 239136.
Ramichloridium indicum (Subram.) de Hoog, Stud. Mycol. 15: 70.
1977. DNA barcode (genus): LSU.
Description and illustration: Arzanlou et al. (2007). DNA barcodes (species): LSU, tef1. Table 14.
Material examined: Unknown collection details, Feb. 1996, L. Marvanova, CBS Mycelium superficial or immersed, composed of branched,
171.96. septate, smooth- and thin-walled, pale white to white or brown
82
length width CBS 908.85 Cu uprreessus lusittanica South Africa

40 12 CBS 123911 Cu uprre essus sem mperv rvirens Morocco
ref. strain
ref. strain
epitype
holotype
epitype
holotype
CBS 523.82 Cuprre ess socy cypari ris sp. New Zealand
90/1 CPC 23798 Cu uppre
r ssus semp perv rvirens Italy
35
10 CBS 522.82 Cupre r ss sus semp perv rvirens New Zealand
78/.88 CBS 122612 Cu uprreess sus sp. Chile
R
30 CBS 909.85 Cup u ressus lus sitaanica South Africa
98/.98 CPC 28287 Cu upre
r ssu us semperrvirens Spain
8
CPC 28286 Cu upre
r ssu us semperrvirens Spain
25 CPC 23794 Greece
CBS 280.93 Cu upress sus s sp. Italy Seiridium
6
CPC 23795 Greece
cardinale
20
CPC 23787 Ju uniiperrus phoeniica a Greece
CPC 23788 Cu upress sus s sempe errvvirens s Italy
unicorne
cardinale
cardinale
unicorne
cancrinum
kenyanum
cancrinum
kenyanum
cupressi
neocupressi
cupressi
neocupressi
CPC 23791 Cup upress sus s semperv rvirens s Italy
CPC 23797 Cup upress sus s sempe errvvirens s Italy
CBS 172.56
A B 95/1 CPC 23785 Cu upress sus s sempe errvvirens s France
basal appendage apical appendage CPC 23796 Cu upress sus s sempe errv
virens s Italy
15 20 CPC 23790 Cu upress sus sempe ervirens s Italy
ref. strain
ref. strain
epitype
holotype
epitype
holotype
CBS 910.85 Cup upress sus s semp perrvvirens s South Africa
85/1 CBS 225.5 55 Cu upress sus s ffo
orbesi Kenya
15 68/1 CBS 320.5 51 Cuprre ess sus s sp p. Kenya Seiridium
10 98/1
CBBS 224.55 ET
5 Cu upprer ssu us macro r carp r a Kenya cupressi
95/1
100/1
CBS 227.55 5 Cuupprre
essu us ma acror carp r a Uganda comb. nov.
10
86/1 CBS 122616 Cup u ress sus sp. Greece
89/1 CBS 143872 Cuprre ess sus sp. Maryland (USA)
96/1
5
CBS 143871ET Cu up
prre essus sp p. Maryland (USA)
5
100/1
CBS 143873 Cuprre ess sus sp. Maryland (USA)
Seiridium
64/1 unicorne
CBS 538.82R Cry rypto omeri ria jja
aponica a New Zealand
89/1 CBS 120306 Cup u ressus sem mperv rvirens South Africa
94/1 CBS 1426 625T Cu uppre r ssus macro r carp r a Italy
Seiridium
neocupressi
unicorne
cardinale
cancrinum
kenyanum
unicorne
cardinale
cancrinum
kenyanum
cupressi
neocupressi
cupressi
69/.98 CBS 142626 Cup uprre ess sus semperv rvirens Italy
CBS 142627 Cuprre essu us ley ylandidii Australia
neocupressi
100/1
100/1 CBS 907.85 Cup u ressus lusiita anica South Africa Seiridium
C D 87/1 CBS 226.55T Cu upprre essus ma acror carrpa Kenya cancrinum
CPC 1996 65 Phyl ylica arrb bore rea Tristan da Cunha
100/1 CBS 133587T Phy ylica a arbr orea Tristan da Cunha
Seiridium
phylicae
CPC 19962 2 Phy ylica arrb bore rea Tristan da Cunha
T
92/1 90/.99 CBS 142628 S p yr
y r i di
d i um glob b o sum Australia S. spyridicola
CBS 143 3445T Persr oonia i sp. Australia Seiridium persooniae
80/.99 CBS 122614 Cup upre
r ssu us sp. Portugal
65/.97 CBS 122613 Cu upress sus sp. Portugal
Seiridium
kenyanum
100/1 95/.97 CBS 288.55T Ju uniiperrus pro r cera ra Kenya
95/1 CBS S 145114 Cu upre
r ssus s mac crocarp r a California (USA)
59/.87 MFL LUCC 13-05 525T Cu uprre ess sus s aririzonica Italy S. pseudocardinale
80/1
CBS 343.97ET Eu ucaaly
lyptus s delega ate ensis i Australia S. eucalypti
CBS 142629T Eucalyptus cladocalyx Australia S. kartense
65/.90 100/1 MFLUCC 12-0647T Camellia reticulata China S. camelliae
T
CBS 137995 Podocarpus latifolus South Africa S. podocarpi
55/.87
MFLUCC 17-0654T Rosa canina Italy Seiridium rosarum
0.02
64/.99 90/.70 CBS 145115 Vitis vinifera Virginia (USA) Seiridium pezizoides comb. nov.
MFLUCC 15-0396R Cornus mas Italy Seiridium venetum
99/1
CBBS 14 40403GT Ros sa canin
i a France Seiridium marginatum
ET
100/1 CBS 340.97 Eucalyptus delegatensis Tasmania Seiridium papillatum
PHSI2001Pathcw07 Vitis vinifera China Seiridium ceratosporum
4x
81/1 CBS 114178GT Neopestalotiopsis protearum
4x
99/1 CBS 122705GT Bartalinia robillardoides
4x
E CBS 139823GT Seimatosporium rosae
Fig. 63. A–D. Boxplots of conidial measurement data in μm from S. unicorne and other Cupressaceae pathogens. The boxes show the lower and upper quantiles and whiskers
extend to 1.5x the interquartile range. Except for the new epitype all measurements are adapted from Bonthond et al. (2018). The ex-epitype strain (CBS 143871), holotype (IMI
5816) and reference strain (CBS 538.82) of S. unicorne are highlighted in blue. E. The best Maximum Likelihood (ML) tree based on four loci (ITS: 616 bp, rpb2: 802 bp, tef1:
633 bp, tub2: 809 bp). Nodes are labelled with ML bootstrap values (BS > 49 %)/Bayesian posterior probabilities (PP > 0.49) using the same model selection, settings and
software as in Bonthond et al. (2018). Strains are displayed by number, host and country of collection. GenBank accession numbers are listed in Bonthond et al. (2018) and in
Table 15. T, ET, GT and R indicate ex-type, ex-epitype, ex-generic type and reference strains, respectively. TreeBASE: S23390.
hyphae. Synnemata erect, simple, unbranched or very rarely slimy masses at the apex of the synnema, aseptate or very rarely
branched, greenish when young, becoming dark brown to black 1-septate, smooth- and thin-walled, oblong, ellipsoidal, sub-
at maturity, capitate at the apex. Conidiophores macronematous, globose or fusiform, hyaline to subhyaline, pale brown or oliva-
synnematous, straight or slightly flexuous, unbranched or ceous (adapted from Partridge & Morgan-Jones 2002).
branched toward the upper region, cylindrical, septate, smooth- Culture characteristics: Colonies effuse, powdery or cottony to
and thin-walled, hyaline to pale brown or olivaceous brown to fairy fluffy, grey to dark brown.
brown. Conidiogenous cells mono- or polyblastic, integrated,
terminal, determinate, cylindrical to subclavate or doliiform, Optimal media and cultivation conditions: MEA, OA and PDA at
smooth-walled. Conidia acropleurogenous, holoblastic, in simple 25 °C.
or branched acropetal chains, dry, sometimes aggregated into Distribution: North America, Europe and Asia.
MARIN-FELIX ET AL.
Fig. 64. Seiridium pezizoides (CBS 145115). A–D. Colony morphology in 90-mm-diam Petri dishes after 10 d at 22 °C on MEA, SNA, PDA and CMA, respectively.
E–K. Conidiomata on Vitis vinifera. L, M. Conidiophores. N. Conidia. Scale bars: E = 1 mm; F–K = 100 μm; L–N = 10 μm.
84
Fig. 65. Seiridium unicorne (ex-epitype CBS 143871). A–D. Colony morphology in 90-mm-diam Petri dishes after 10 d at 22 °C on MEA, SNA, PDA and CMA, respectively.
E. Symptoms on naturally infected host. F, G. Conidiomata on artificially infected Cupressaceae sp. H. Sporulation on PDA. I. Conidioma on SNA partially immersed in agar.
J–O. Conidiophores and conidia. P. Conidia. Scale bars: F–H = 100 μm; I–P = 10 μm.
MARIN-FELIX ET AL.
Fig. 66. Septoriella spp. A. Conidiomata on OA of Septoriella hirta (ex-neotype CBS 536.77). B. Conidiomata in vivo of Septoriella phragmitis (ex-epitype CBS 140065).
C. Conidial cirrhus of Septoriella phragmitis (ex-epitype CBS 140065). D. Conidioma of Septoriella hirta (ex-neotype CBS 536.77). E, F. Section through conidiomata of
Septoriella hirta (ex-neotype CBS 536.77). G–I. Conidiogenous cells. G. Septoriella oudemansii (ex-type CBS 138012). H, I. Septoriella phragmitis (ex-epitype CBS 140065).
J. Developing conidia of Septoriella hirta (ex-neotype CBS 536.77). K–O. Conidia. K. Septoriella hirta (ex-neotype CBS 536.77). L. Septoriella oudemansii (ex-type CBS
138012). M. Septoriella poae (ex-type CBS 136766). N, O. Septoriella phragmitis (ex-epitype CBS 140065). Scale bars: D, E = 100 μm; F = 50 μm; all others = 10 μm. Pictures
A–F, H–K, N, O taken from Crous et al. (2015a,b,c); G, L from Crous et al. (2014b).
86
Hosts: Species of Rhododendron. Rhododendron bud blight disease, in which the flower buds die,
and twig blight occurs. Infected buds are easily recognisable by
Disease symptoms: Bud and twig blight; Rhododendron bud the blackening of the bud and the development of numerous
blight disease. synnemata which appear as tiny black spines over the entire
surface (Partridge & Morgan-Jones 2002, Glawe & Hummel
Notes: Seifertia was introduced by Partridge & Morgan-Jones
2006).
(2002) to accommodate Pycnostysanus azaleae based on
This genus, which is relatively poorly studied, was recently
morphological differences. Seifertia azaleae is morphologically
placed in Melanommataceae by Li et al. (2016b), when they
similar to Sorocybe resinae. However, Partridge & Morgan-Jones
introduced the second species belonging to Seifertia, Sei.
(2002) decided to erect the new genus Seifertia since Sei.
azaleae produces much narrower conidia and has minute den- shangrilaensis. However, the relation of Seifertia with Myco-
pappus and its synasexual morph Xenostigmina, which are foliar
ticles on the conidiogenous cells. Seifertia is characterised by
pathogens belonging to Melanommataceae, was demonstrated
erect, simple, and dark synnemata, macronematous co-
previously by Crous et al. (2009a).
nidiophores, holoblastic, integrated, terminal and determinate
conidiogenous cells, and unicellular or very rarely 1-septate, pale References: Partridge & Morgan-Jones 2002 (morphology),
brown or olivaceous conidia. This cosmopolitan genus occurs on Glawe & Hummel 2006 (pathogenicity), Seifert et al. 2007, Li
azaleas and rhododendrons causing a disease known as et al. 2016b (morphology and phylogeny).
Table 16. DNA barcodes of accepted Septoriella spp.

ITS rpb2 tef1 tub2
Septoriella agrostina MFLU 18-0113HT MG828945 – MG829227a – Wanasinghe et al. (2018)
Sep. allojunci MFLUCC 15-0701 T
KU058718 – MG520935a – Li et al. (2015), Phookamsak et al. (2017)
Sep. artemisiae MFLUCC 17-0693 T
MG828929 – – – Wanasinghe et al. (2018)
Sep. arundinicola MFLU 16-0225HT MG828946 MG829261 MG829228a – Wanasinghe et al. (2018)
Sep. arundinis MFLUCC 15-0702T KU058716 – MG520921a – Li et al. (2015), Phookamsak et al. (2017)
Sep. bromi MFLUCC 13-0739 T
KU058717 – – – Li et al. (2015)
Sep. chlamydospora MFLUCC 15-0177 T
KU163658 – – – Hyde et al. (2018)
Sep. dactylidicola3 MFLUCC 14-0002T – – – – –
Sep. dactylidis MFLU 15-2720 HT
KU163657 – – – Jayasiri et al. (2015)
Sep. elongata MFLUCC 12-4444T KM491546 – – – Li et al. (2015)
Sep. forlicesenica MFLUCC 15-0470 T
KX926422 KY131966 MG520922 a
– Phookamsak et al. (2017), Thambugala et al. (2017)
Sep. garethjonesii MFLUCC 15-0469T KX926425 KX898363 MG520923a – Phookamsak et al. (2017), Thambugala et al. (2017)
T b
Sep. germanica CBS 145372 MK539965 MK540096 MK540159 MK540174 Present study
Sep. hibernica CBS 145371T MK539966 MK540097 – – Present study
Sep. hirta CBS 536.77 ET
KR873249 KR873324 – – Crous et al. (2015a)
Sep. hollandica CBS 145374T MK539967 MK540098 MK540160b MK540175 Present study
Sep. hubertusii CBS 338.86T KF251230 KF252235 – KF252717 Quaedvlieg et al. (2013)
Sep. italica MFLUCC 13-0267T KX926421 KX891169 MG520924a – Phookamsak et al. (2017), Thambugala et al. (2017)
Sep. leuchtmannii CBS 459.84IsoT KF251188 KF252195 KF253144b KF252682 Quaedvlieg et al. (2013)
Sep. muriformis MFLUCC 13-0277 T
KX926415 KX863710 – – Thambugala et al. (2017)
Sep. neoarundinis MFLUCC 15-0027T KY706139 – MG520936a – Phookamsak et al. (2017), Thambugala et al. (2017)
Sep. neodactylidis MFLUCC 13-0618T KP744432 – – – Liu et al. (2015)
Sep. oudemansii CBS 138012T KR873250 – – – Crous et al. (2015a)
Sep. phragmitis CBS 140065 ET
KR873251 – – – Crous et al. (2015a)
Sep. poae CBS 136766T KJ869111 KJ869233 – – Crous et al. (2014b)
T b
Sep. pseudophragmitis CBS 145417 MK560161 MK559450 MK559452 MK559451 Present study
Sep. rosae MFLU 18-0114HT MG828948 – MG829230a – Wanasinghe et al. (2018)
Sep. subcylindrospora MFLUCC 13-0380 T
KT314184 – – – Ariyawansa et al. (2015a)
Sep. tridentina MFLUCC 15-0475T KX926424 KX891171 – – Thambugala et al. (2017)
b
Sep. vagans CBS 604.86 KF251193 KF252200 KF253149 KF252687 Quaedvlieg et al. (2013)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; MFLU and MFLUCC: Herbarium and culture collection of Mae Fah Luang University, Chiang Rai,
Thailand, respectively. T, ET, HT and IsoT indicate ex-type, ex-epitype, holotype and ex-isotype strains, respectively.
2
elongation factor 1-alpha gene; tub2: partial β-tubulin gene. a and b in tef1 column indicate the primers used in sequencing: a: EF1-983F, EF1-2218R, b: EF1-728F, EF-2.
3
Only LSU and SSU sequences available: KY657264 and KY657265, respectively (Thambugala et al. 2017).
MARIN-FELIX ET AL.
Fig. 67. RAxML phylogram obtained from the combined ITS (580 bp), LSU (849 bp) and rpb2 (1083 bp) sequence alignment of all accepted species of Septoriella. The tree
was rooted to Neostagonospora caricis CBS 135092 and Neostagonospora elegiae CBS 135101. The novelties proposed in this study are indicated in bold. RAxML bootstrap
support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession numbers of LSU are listed in Fig. 28, and of
the other loci in Tables 7 and 16. T, ET, HT and IsoT indicate ex-type, ex-epitype, holotype and ex-isotype strains, respectively. TreeBASE: S23834.
88
Fig. 68. Septoriella germanica (ex-type CBS 145372). A. Conidiomata sporulating on PNA. B, C. Conidiogenous cells. D. Conidia with mucoid caps. Scale bars: A = 200 μm; all
others = 10 μm.
Authors: Y. Marin-Felix & P.W. Crous conidiogenous cells, hyaline, smooth-walled. Conidiogenous
cells discrete, integrated, ampulliform to lageniform or sub-
Seiridium Nees, Das System der Pilze und Schw€amme: 22.
cylindrical, hyaline, smooth-walled, proliferating percurrently at
1817. Fig. 62.
the apex. Conidia fusiform, euseptate (septal pores present or
Synonym: Pestalotia De Not., Mem. Reale Accad. Sci. Torino 3:
not), end cells hyaline, median cells dark brown to brown, wall
80. 1841.
thick, smooth or with striations, constricted at septa or not; apical
Additional synonyms in Bonthond et al. (2018).
cell with single or multiple, unbranched or branched appendages;
Classification: Sordariomycetes, Xylariomycetidae, Xylariales, basal cell with or without a centric, unbranched or sometimes
Sporocadaceae. branched appendage (adapted from Bonthond et al. 2018).
Type species: Seiridium marginatum Nees, Syst. Pilze (Würz- Culture characteristics: Colonies on PDA circular to irregular,
burg): 23. 1817. Neotype designated by Shoemaker et al. (1966): reaching 12–68 mm diam after 14 d at 22 °C, mostly flat, in
K 200376. Epitype and ex-epitype culture designated by some cultures elevated at margins, often white coloured or with
Jaklitsch et al. (2016): WU 33575, CBS 140403. other colours, sporulation rare, with pycnidioid conidiomata. On
SNA circular to irregular to rhizoid, reaching 20–54 mm diam
DNA barcode (genus): ITS.
after 14 d at 22 °C, mostly flat, white coloured, with moderate to
DNA barcodes (species): ITS, rpb2, tef1, tub2. Table 15. Fig. 63. abundant aerial mycelium, sporulation for some species within
and others after 2 wk, sporodochia often compact and scattered.
Ascomata perithecial, immersed to semi-erumpent, depressed,
globose to pyriform, scattered or confluent, with central ostiole; Optimal media and cultivation conditions: Colonies grow well on
necks slightly papillate, black, periphysate; ascomatal wall dark CMA, MEA, PDA and SNA at 22 °C. Most species sporulate on
brown, pseudoparenchymatous. Paraphyses hyaline, smooth- SNA and some species on CMA, MEA or PDA as well.
walled, filiform. Asci cylindrical, 8-spored, unitunicate, thin-
walled, stipitate, with an apical amyloid ring. Ascospores
cylindrical-oblong, euseptate, septa often thicker than the wall, Hosts: A diversity of gymnosperms and angiosperms. The genus
yellow- to dark brown, guttulate. Conidiomata acervuloid to is most well-known from members of Cupressaceae.
pycnidioid, semi-immersed to erumpent, uni- to plurilocular,
Disease symptoms: Cankers on stems and twigs.
brown or black, glabrous, dehiscing by irregular splits in the
upper wall. Conidiophores lining the cavity of the conidioma, Notes: Pestalotia was introduced in 1841 and is similar to
septate and sparsely branched at the base, or reduced to Seiridium (1817), one of the older names in the Sporocadaceae.
Fig. 69. Septoriella hollandica (ex-type CBS 145374). A. Ascomata sporulating on PNA. B, C. Asci. D. Ascospores. Scale bars = 10 μm.
MARIN-FELIX ET AL.
Fig. 70. Septoriella pseudophragmitis (ex-type CPC 24166). A. Conidiomata sporulating on MEA. B, C. Conidiogenous cells. D. Conidia. Scale bars: A = 250 μm; all
others = 10 μm.
The genus has been subjected to many rearrangements recently obtained (Table 1), which matches in all respects with
(reviewed in Sutton 1969) which eventually resulted in the genus the type of Pe. pezizoides. DNA sequence data generated here
accommodating only the type species, Pe. pezizoides. One of confirm that Pe. pezizoides clusters within Seiridium (Fig. 63).
the important characters used to separate Pestalotia from the Therefore, Pestalotia is reduced to synonymy with Seiridium. For
related genera Pestalotiopsis and Truncatella was the production Seiridium, the here synonymised Pestalotia and related genera,
of 5-septate (or 6-celled) conidia. However, this morphology is not only the number but also the type of conidial septation has
typical for Seiridium as well, and for this reason it was speculated been a commonly reported descriptor. Different authors have
that Pestalotia and Seiridium could be congeneric interpreted the appearance of the cell walls as either distoseptate
(Maharachchikumbura et al. 2014). A fresh collection of (e.g. Nag Raj 1993) or euseptate (e.g. Sutton 1980). Although
Pe. pezizoides from Vitis vinifera collected in the USA was when examined by light microscopy, conidia can appear as
Fig. 71. Setophoma spp. A, B. Conidioma forming in culture. A. Setophoma chromolaenae (ex-type CBS 135105). B. Setophoma vernoniae (ex-type CBS 137988). C–E.
Conidiomata with setae of Setophoma chromolaenae (ex-type CBS 135105). F–H. Conidiogenous cells. F, G. Setophoma chromolaenae (ex-type CBS 135105). H. Setophoma
vernoniae (ex-type CBS 137988). I, J. Conidia. I. Setophoma chromolaenae (ex-type CBS 135105). J. Setophoma vernoniae (ex-type CBS 137988). Scale bars: C–E = 20 μm;
all others = 10 μm; F applies to F and G.
90
Table 17. DNA barcodes of accepted Setophoma spp.

ITS rpb2 tef1 tub2
Setophoma brachypodii CBS 145418T MK539968 MK540099 MK540161 – Present study
Set. chromolaenae CBS 135105T KF251244 KF252249 KF253195 KF252728 Quaedvlieg et al. (2013)
Set. pseudosacchari CBS 145373 T
MK539969 MK540100 – MK540176 Present study
Set. sacchari CBS 333.39ET KF251245 KF252250 – – Quaedvlieg et al. (2013)
LT
Set. terrestris CBS 335.29 KF251246 KF252251 KF253196 KF252729 Quaedvlieg et al. (2013)
Set. vernoniae CBS 137988T KJ869141 – MK540162 MK540177 Crous et al. (2014b), present study
1 T, ET LT
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. and indicate ex-type strains, ex-epitype and ex-lectotype, respectively.
2
distoseptate, electron microscopic studies on S. cupressi Synonyms: Cryptostictis cupressi Guba, Monograph of Mono-
(Roberts & Swart 1980) and S. pezizoides (Griffiths & Swart chaetia and Pestalotia: 47. 1961. Nom. inval. Art. 40.3
1974) have shown that the conidial cell walls are differentiated (Shenzhen).
into multiple zones but arise from a single layered cell wall and Lepteutypa cupressi (Nattrass et al.) H.J. Swart, Trans. Brit.
are thus euseptate. Since the sexual morph is known for only a Mycol. Soc. 61: 79. 1973.
few species, the taxonomy in Seiridium has been based mainly Seiridium cupressi (Guba) Boesew, Trans. Brit. Mycol. Soc. 80:
on asexual morphology. However, the generic type 545. 1983. Nom. inval. Art. 40.3 (Shenzhen).
(S. marginatum) is one of the exceptions where both morphs
Description: Sexual morph Nattrass et al. (1963). Asexual morph
have been characterised. This species was re-described and
Bonthond et al. (2018).
epitypified by Jaklitsch et al. (2016), who also provided detailed
illustrations of sexual and asexual morphology. Known distribution: Africa (Kenya, Uganda) and Europe
(Greece).
References: Nag Raj 1993 (morphology), Danti & Della Rocca
2017 (pathogenicity), Bonthond et al. 2018 (morphology and Typus: Africa, Kenya, on Cupressus macrocarpa, July 1954,
phylogeny), R.M. Nattrass (holotype of Rhynchosphaeria cupressi IMI
56917); from cankers in branches of Cupressus macrocarpa,
Seiridium cupressi (Nattrass et al.) Bonthond, Sand.-Den. &
1949, D.R. Jones [epitype of Rhynchosphaeria cupressi
Crous, comb. nov. MycoBank MB830554.
designated here IMI 52254, MBT386544 (dried culture), culture
Basionym: Rhynchosphaeria cupressi Nattrass et al., Trans. Brit.
ex-epitype CBS 224.55].
Mycol. Soc. 46: 103. 1963.
Fig. 72. RAxML phylogram obtained from the combined ITS (589 bp), LSU (835 bp), tef1 (788 bp) and tub2 (532 bp) sequence alignment of all accepted species of
Setophoma. The tree was rooted to Neostagonospora caricis CBS 135092 and Neostagonospora elegiae CBS 135101. The novelties proposed in this study are indicated in
are listed in Fig. 28, and of the other loci in Tables 7, 17 and 18. T, ET and LT indicate ex-type strains, ex-epitype and ex-lectotype, respectively. TreeBASE: S23834.
MARIN-FELIX ET AL.
Fig. 73. Setophoma pseudosacchari (ex-type CBS 145373). A. Ascomata sporulating on OA. B. Asci with ascospores. C. Conidia. Scale bars: A = 300 μm; all others = 10 μm.
Additional materials examined: Europe, Greece, from Cupressus sp., A. Graniti, Seiridium pezizoides (De Not.) Crous, comb. nov. MycoBank
CBS 122616 = CMW 1646. Africa, Kenya, non-pathogenic isolate from MB828021. Fig. 64.
Cupressus sp., collection data unknown, CBS 320.51; on Cupressus macro-
carpa, July 1948, R.M. Nattrass, IMI 37158; on Cupressus macrocarpa, Dec.
Basionym: Pestalotia pezizoides De Not., Mem. Reale Accad.
1949, R.M. Nattrass, IMI 40096; from cankers in branches of Cupressus forbesii, Sci. Torino 2, 3: 80. 1839.
1949, D.R. Jones, IMI 52255 (dried culture); CBS 225.55.
Caulicolous. Isolated from branches of Vitis vinifera. On the host
Notes: This species has been a source of confusion since its (described in more detail by Nag Raj 1993): Conidiomata
introduction (Guba 1961). Bonthond et al. (2018) showed that irregularly scattered over the surface, gregarious to confluent,
Guba’s diagnosis included three different species (S. cancrinum, discoid to cupulate and occasionally globose, erumpent from
S. cupressi and S. kenyanum) and selected an epi- and lectotype tissue, acervular to sporodochial, occasionally with aerial
for Cryptosticis cupressi. The latter name, however, was invalidly mycelium, black to brown, (300–)350–500(–650) μm. On SNA:
published (article 40.1). Nattrass et al. (1963) re-examined the Conidiophores tightly aggregated in the conidioma, cylindrical,
material from Guba (1961) and synonymised C. cupressi with irregularly branched, hyaline or pale brown at the base, smooth-
Monochaetia unicornis. Despite noting small morphological dif- and thin-walled. Conidiogenous cells discrete, hyaline, cylindri-
ferences between the asexual stage of R. cupressi and the type cal, smooth- and thin-walled. Conidia lunate to falcate, often
of M. unicornis (i.e. smaller and slender conidia), they were not curved, 5-septate, not striate, bearing a basal and two or more
able to confirm the exogenous origin of the basal appendage, apical appendages, euseptate with pores sometimes visible,
which was the main argument for Guba (1961) to place the (24–)28–33.5(–38.5) × (6–)7–8(–9) μm, mean ± SD = 30.7 ±
species in Cryptostictis instead of Monochaetia. In the same 2.8 × 7.5 ± 0.4 μm; basal cell obconic with truncate base, hy-
study, Nattrass et al. (1963) described the sexual morph of aline, smooth-walled, bearing marginal frills, 4–7 μm; four me-
M. unicornis as Rhynchosphaeria cupressi based on three dian cells pale brown, smooth-walled, cylindrical to doliiform;
specimens: IMI 37158, IMI 40096 and the holotype IMI 56917. second cell from base 3.5–8 μm; third cell 3.5–7 μm; fourth cell
Bonthond et al. (2018) examined each of these specimens but 3.5–6.5 μm; fifth cell 3.5–7.5 μm; apical cell conical, hyaline,
incorrectly cited the holotype as IMI 37158. While only the sexual smooth-walled, 4.5–8.5 μm long; apical appendages single or
morph was found in these materials, the original description from multiple, centric, branched or unbranched, 8.5–27 μm; basal
Nattrass et al. (1963) includes drawings, photographs and appendage single, cylindrical, centric, occasionally branched,
measurements of the conidia. These measurements 5.5–14 μm.
(22–32 × 6–9.5 μm) fall perfectly within the range documented
Culture characteristics: Colonies on PDA circular, reaching
for the lineage currently assigned to S. cupressi (Guba) Boesew.
34–37 mm diam after 10 d at 22 °C, flat, olivaceous to luteous in
(18–36 × 5–11.5 μm) (Bonthond et al. 2018). Consequently,
the centre, white to brown at the margins, with abundant aerial
being the oldest valid name for this lineage and in accordance
mycelium at the margins, not sporulating within 10 d. On CMA
with the rule of priority Ry. cupressi is recombined in Seiridium,
circular, reaching 39–41 mm diam after 10 d at 22 °C, flat at
as S. cupressi (Nattrass et al.) Bonthond, Sand.-Den. & Crous,
centre and margins, dark brown to black, without aerial myce-
and an epitype is designated (IMI 52254). The similar species
lium, not sporulating within 10 d. On MEA circular to slightly
S. cancrinum and S. unicorne show smaller conidia (20–30.5 μm
irregular, reaching 25–27 mm diam after 10 d at 22 °C, flat,
and 19–28 μm, respectively), whereas conidia of S. kenyanum
olivaceous to pale green, with a white outer ring, with moderate
are considerably larger (24–39 μm).
aerial mycelium, massive spore production in the centre. On
SNA circular, reaching 31–33 mm diam after 10 d at 22 °C,
Table 18. DNA barcodes of the accepted Wingfieldomyces sp.

ITS rpb2 tef1 tub2
T
Wingfieldomyces cyperi CBS 141450 KX228286 MK540101 MK540163 MK540178 Crous et al. (2016b), present study
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. T indicates ex-type strain.
2
92
raised in the centre, flat at the margins, with moderate aerial the conclusion that Pestalotia and Seiridium are congeneric.
mycelium, no sporulation within 10 d. Consequently, the species is transferred to Seiridium as
S. pezizoides. The S. pezizoides strain (CBS
Distribution: Europe (France, Italy), USA.
145115 = CPC 35011) that was included in this analysis
Typus: Italy, near Mailand, twig of Vitis vinifera (Vitaceae), leg. conforms morphologically with the description and was
Oct. 1838, De Notaris, holotype, RO. isolated from the same host, Vitis. However, since the
Additional material examined: USA, Virginia, Charlottesville, from a complex
specimen was collected in Virginia (USA) and the holotype
hybrid of Vitis æstivalis × Vitis cinerea × Vitis vinifera, 2018, L. Morton, CBS is from Italy, it is not suitable for epitypification. The phy-
145115 = CPC 35011. logeny generated here suggests that S. pezizoides is most
Notes: After several rearrangements and the introduction of closely related to S. rosarum (Rosa canina, Italy).
the genera Pestalotiopsis and Truncatella (see Sutton 1969 Seiridium unicorne (Cooke & Ellis) B. Sutton, Mycol. Pap. 138:
and references therein), the generic type Pe. pezizoides 74. 1975. Fig. 65.
was the only remaining species in Pestalotia. A specimen Basionym: Pestalotia unicornis Cooke & Ellis, Grevillea 7: 6.
of Pe. pezizoides (IMI 83642, from branches of Vitis 1878, as "Pestalozzia".
vinifera, Italy, non-type) was examined and redescribed by Synonym: Monochaetia unicornis (Cooke & Ellis) Sacc. & D.
Sutton (1980). A more detailed description was provided by Sacc., Syll. Fung. 18: 485. 1906.
Nag Raj (1993), based on several materials, including this
specimen. The present study is the first to provide DNA Caulicolous. Most commonly isolated from cankers on
branches of species from Cupressaceae. Conidiomata on PDA
sequence data on this species. We sequenced four loci
numerous, sporodochial, globose or clavate, mostly solitary,
(ITS, rpb2, tef1 and tub2) and included Pe. pezizoides in
erumpent from agar, partially immersed in mycelium, producing
an updated phylogeny of Seiridium (Fig. 63) which supports
Fig. 74. Wingfieldomyces cyperi (ex-type CBS 141450). A. Symptomatic leaf of Cyperus sphaerocephala. B–D. Asci. E. Pseudoparaphyses. F–I. Ascospores. Scale bars:
10 μm; F applies to F–I. Pictures B, C, E–I taken from Crous et al. (2016b).
MARIN-FELIX ET AL.
Fig. 75. Stagonosporopsis spp. A, B. Disease symptoms of Stagonosporopsis tanaceti (ex-type CBS 131484). A. Leaf necrosis. B. Drooping flower heads. C–G. Sexual morph
of Stagonosporopsis inoxydabilis (ex-type CBS 425.90). C. Close-up of ascoma with darkened ostiolar area. D, E. Stipitate, bitunicate asci. F, G. Ascospores (arrows denote
sheath). H–T. Asexual morph. H, I. Colony sporulating on OA. H. Stagonosporopsis chrysanthemi (CBS 500.63). I. Stagonosporopsis tanaceti (ex-type CBS 131484). J. Close-
up of pycnidial conidiomata of Stagonosporopsis tanaceti (ex-type CBS 131484). K. Close-up of darkened ostiolar area of Stagonosporopsis chrysanthemi (CBS 500.63). L–Q.
Conidiogenous cells. L–N. Stagonosporopsis chrysanthemi (CBS 500.63). O–Q. Stagonosporopsis tanaceti (ex-type CBS 131484). R, S. Conidia. R. Stagonosporopsis
chrysanthemi (CBS 500.63). S. Stagonosporopsis tanaceti (ex-type CBS 131484). T. Chain of chlamydospores of Stagonosporopsis tanaceti (ex-type CBS 131484). Scale bars:
C = 35 μm; J = 150 μm; all others = 10 μm; D applies to D and E; L applies to L–N; O applies to O–Q. Pictures taken from Vaghefi et al. (2012).
94
Table 19. DNA barcodes of accepted Stagonosporopsis spp.

Species Strain1 GenBank accession numbers2 References
ITS LSU act rpb2 tub2
Stagonosporopsis actaeae CBS 106.96T GU237734 GU238166 JN251974 KT389672 GU237671 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. ailanthicola MFLUCC 16-1439T KY100872 KY100874 – KY100876 KY100878 Tibpromma et al. (2017)
Sta. ajacis CBS 177.93NT GU237791 GU238168 JN251962 KT389673 GU237673 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. andigena CBS 101.80 GU237714 GU238169 JN251958 – GU237674 Aveskamp et al. (2010), de Gruyter et al.
(2012)
Sta. artemisiicola CBS 102636 GU237728 GU238171 JN251971 KT389674 GU237676 Aveskamp et al. (2010), de Gruyter
et al. (2012), Chen et al. (2015)
Sta. astragali CBS 178.25 GU237792 GU238172 JN251963 – GU237677 Aveskamp et al. (2010), de Gruyter et al.
(2012)
Sta. bomiensis CGMCC 3.18366T KY742123 KY742277 – KY742189 KY742365 Chen et al. (2017)
Sta. centaureae MFLUCC 16-0787T KX611240 KX611238 – – – Hyde et al. (2016)
Sta. citrulli ATCC TSD-2 T
KJ855546 – – – KJ855602 Stewart et al. (2015)
Sta. caricae CBS 248.90 GU237807 GU238175 JN251969 – GU237680 Aveskamp et al. (2010), de Gruyter et al.
(2012)
DAR 28714 MK253236 MK253237 MK255065 MK255066 MK255067 Present study
Sta. chrysanthemi ATCC 10748 ET
JQ897484 JQ897460 JQ897508 – JQ897504 Vaghefi et al. (2012)
Sta. crystalliniformis CBS 713.85T GU237903 GU238178 JN251960 KT389675 GU237683 Aveskamp et al. (2010), de Gruyter
et al. (2012), Chen et al. (2015)
Sta. cucurbitacearum CBS 133.96 GU237780 GU238181 JN251968 KT389676 GU237686 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. dennisii CBS 631.68ET GU237899 GU238182 Q-bank3 KT389677 GU237687 Aveskamp et al. (2010), Chen et al.
(2015)
Sta. dorenboschii CBS 426.90T GU237862 GU238185 JN251980 KT389678 GU237690 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. helianthi CBS 200.87T KT389545 KT389761 – KT389683 KT389848 Chen et al. (2015)
Sta. heliopsidis CBS 109182 GU237747 GU238186 JN251980 KT389679 GU237691 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. hortensis CBS 572.85 GU237730 GU238198 JN251966 KT389680 GU237703 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. inoxydabilis CBS 425.90T GU237861 GU238188 JN251972 KT389682 GU237693 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. loticola CBS 562.81IsoT GU237890 GU238192 JN251978 KT389684 GU237697 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. lupini CBS 101494T GU237724 GU238194 JN251967 KT389685 GU237699 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. oculo-hominis CBS 634.92T GU237901 GU238196 JN251976 KT389686 GU237701 Aveskamp et al. (2010), de Gruyter
et al. (2012), Chen et al. (2015)
Sta. papillata CGMCC 3.18367T KY742125 KY742279 – KY742191 KY742367 Chen et al. (2017)
Sta. rudbeckiae CBS 109180 GU237745 GU238197 Q-bank3 – GU237702 Aveskamp et al. (2010)
Sta. tanaceti CBS 131484T JQ897481 JQ897461 JQ897512 – JQ897496 Vaghefi et al. (2012)
Sta. trachelii CBS 379.91 GU237850 GU238173 JN251977 KT389687 GU237678 Aveskamp et al. (2010), de Gruyter et al.
(2012), Chen et al. (2015)
Sta. valerianellae CBS 329.67IsoT GU237832 GU238201 JN251965 – GU237706 Aveskamp et al. (2010), de Gruyter et al.
(2012)
1
ATCC: American Type Culture Collection, Virginia, USA; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: Chinese General Microbi-
ological Culture Collection Center, Beijing, China; DAR: New South Wales Plant Pathology Herbarium, NSW, Australia; MFLUCC: Mae Fah Luang University Culture
Collection, Chiang Rai, Thailand. T, ET, IsoT and NT indicate ex-type, ex-epitype, ex-isotype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: partial large subunit (28S) nrRNA gene; act: partial actin gene; rpb2: partial DNA-directed RNA
polymerase II second largest subunit; tub2: partial β-tubulin gene.
3
Q-bank: Sequences retrieved from Q-bank Fungi database (http://www.q-bank.eu/fungi/).
MARIN-FELIX ET AL.
large black spore masses; on SNA, sporodochial, mostly 1.9 × 7.8 ± 0.4 μm; basal cell obconic with a truncate base,
aggregated, erumpent from agar, producing large black spore hyaline, walls smooth, bearing minute marginal frills,
masses. On SNA: Conidiophores septate, cylindrical, irregu- 2.5–9.5 μm, ex-epitype: 2.5–6 μm (n = 119); four median cells
larly branched, hyaline or brown, thin-walled, 22–68 μm long, colour varying from pale to dark brown, smooth-walled, cylin-
ex-epitype: 22–50 μm long. Conidiogenous cells discrete, hy- drical to doliiform; second cell from base 3.5–6 μm (n = 152);
aline, cylindrical, smooth- and thin-walled, third cell 3–5.5 μm; fourth cell 3–5.5 μm; fifth cell 3–5.5 μm,
3.5–29.5 × 1.5–3.5 μm, ex-epitype: ex-epitype: second cell from base 3.5–6 μm; third cell
16.2–28.9 × 1.7–3.5 μm, proliferating percurrently, with 3–5.5 μm; fourth cell 3–5 μm; fifth cell 3–5.5 μm; apical cell
visible collarettes and minute periclinal thickenings. Conidia conical, hyaline, smooth-walled, 2–5.5 μm long, ex-epitype:
lunate to falcate, curved, 5-septate, rarely 4- or 6-septate, not 2–5.5 μm long; apical appendage single, mostly centric,
striate, bearing two appendages, euseptate with no visible 5–10 μm, ex-epitype: 5–9.5 μm; basal appendage single, cy-
pores, (19–)22.5–26.5( –28) × (6.5–)7.5–8.5(–9.5) μm, mean lindrical, mostly excentric, 2.7 –7.1 μm, ex-epitype: 4–6.5 μm
± SD = 24.5 ± 1.8 × 7.9 ± 0.5 μm, ex-epitype: (19–) (adapted from Bonthond et al. 2018).
22.5–26.5( –28) × (6.5–)7.5–8.5( –9) μm, mean ± SD = 24.4 ±
Fig. 76. The majority rule consensus phylogram of Stagonosporopsis spp. inferred from the concatenated LSU (876 bp), ITS (459 bp), tub2 (299 bp) and rpb2 (596 bp)
sequence alignment using Bayesian Inference. The tree is rooted to Boeremia exigua var. exigua CBS 431.74. Bootstrap support values > 75 % and PP values > 0.90 are
shown above or below the branches. T, ET, IsoT and NT indicate ex-type, ex-epitype, ex-isotype and ex-neotype strains, respectively. Genbank accession numbers are indicated in
Table 19, Aveskamp et al. (2010) and Vaghefi et al. (2012). TreeBASE: S23800.
96
Fig. 77. Stemphylium spp. A. Disease symptoms caused by Stemphylium vesicarium (BRIP 65181) on pyrethrum leaves. B, C. Pseudothecial ascomata of Stemphylium
vesicarium (BRIP 65181) on pyrethrum flower stems in vivo and in vitro respectively. D–H. Conidia of Stemphylium spp. D, H. Stemphylium vesicarium (BRIP 65181). E.
Stemphylium truncatulae (holotype BRIP 14850). F, G. Stemphylium waikerieanum (ex-type VPRI 21969). I–K. Conidiophores of Stemphylium spp. I. 1-branched conidiophore
of Stemphylium vesicarium (BRIP 65181). J. Simple conidiophore of Stemphylium truncatuale (holotype BRIP 14850). K. Branched and immature conidiophores of Stemphylium
waikerieanum (ex-type VPRI 21969). L–N. Sexual morphs. L, M. Asci and ascospores of Stemphylium vesicarium isolated from the dead flower stems of pyrethrum. N.
Immature pseudothecial ascomata of Stemphylium truncatulae on SNA after 1 wk. Scale bars: D–K = 20 μm; L, M = 100 μm. Pictures A–D, H, I, M taken from Moslemi et al.
(2017).
MARIN-FELIX ET AL.
Table 20. DNA barcodes of accepted Stemphylium spp.

ITS gapdh cmdA
Stemphylium amaranthi CBS 124746T KU850505 KU850652 KU850793 Woudenberg et al. (2017)
Ste. armeriae CBS 338.73 KU850511 KU850658 KU850799 Woudenberg et al. (2017)
ET
Ste. astragali CBS 116583 KU850512 KU850659 KU850800 Woudenberg et al. (2017)
Ste. beticola CBS 141024T KU850520 KU850667 KU850808 Woudenberg et al. (2017)
VPRI 42502 MK336834 MK336880 MK336857 Present study
Ste. botryosum CBS 714.68T KC584238 AF443881 KU850826 Woudenberg et al. (2017)
T
Ste. callistephi CBS 527.50 KU850539 KU850686 KU850828 Woudenberg et al. (2017)
Ste. canadense CBS 116602T KU850641 KU850782 KU850932 Woudenberg et al. (2017)
T
Ste. chrysanthemicola CBS 117255 KU850640 KU850781 KU850931 Woudenberg et al. (2017)
Ste. drummondii CBS 346.83T GQ395365 KU850687 KU850829 Woudenberg et al. (2017)
Ste. eturmiunum CBS 109845T KU850541 KU850689 KU850831 Woudenberg et al. (2017)
BRIP 27557 MK336820 MK336866 MK336843 Present study
Ste. gracilariae CBS 482.90T KU850549 AF443883 KU850839 Woudenberg et al. (2017)
Ste. halophilum CBS 337.73T KU850553 KU850700 KU850843 Woudenberg et al. (2017)
Ste. ixeridis CBS 124748T KU850590 KU850737 KU850881 Woudenberg et al. (2017)
ET
Ste. lancipes CBS 133314 KU850596 KU850742 KU850887 Woudenberg et al. (2017)
Ste. loti CBS 407.54T KU850597 KU850743 KU850888 Woudenberg et al. (2017)
T
Ste. lucomagnoense CBS 116601 KU850629 KU850770 KU850920 Woudenberg et al. (2017)
Ste. lycii CBS 125241T KU850602 KU850748 KU850893 Woudenberg et al. (2017)
NT
Ste. lycopersici CBS 122639 KU850611 KU850756 KU850902 Woudenberg et al. (2017)
Ste. majusculum CBS 717.68T KU850618 AF443891 KU850909 Woudenberg et al. (2017)
T
Ste. novae-zelandiae CBS 138295 KU850631 KU850772 KU850922 Woudenberg et al. (2017)
Ste. paludiscirpi CBS 109842T KU850620 KU850762 KU850911 Woudenberg et al. (2017)
Ste. rombundicum BRIP 27486 HT
MK336819 MK336865 MK336842 Present study
Ste. sarciniforme CBS 110049 KU850591 KU850738 KU850882 Woudenberg et al. (2017)
T
Ste. simmonsii CBS 133518 KU850637 KU850778 KU850928 Woudenberg et al. (2017)
Ste. solani CBS 116586ET KU850627 KU850768 KU850918 Woudenberg et al. (2017)
T
Ste. symphyti CBS 115268 KU850643 KU850784 KU850934 Woudenberg et al. (2017)
Ste. trifolii CBS 116580T KU850647 KU850788 KU850938 Woudenberg et al. (2017)
T
Ste. triglochinicola CBS 718.68 KU850648 KU850789 KU850939 Woudenberg et al. (2017)
Ste. truncatulae BRIP 14850HT MK336815 MK336861 MK336838 Present study
Ste. vesicarium CBS 715.68 KU850565 KU850712 KU850855 Woudenberg et al. (2017)
Ste. waikerieanum VPRI 21969HT MK336832 MK336878 MK336855 Present study
1
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Queensland, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; VPRI: Victorian
Plant Pathology Herbarium, Bundoora, Victoria, Australia. T, ET, HT and NT indicate ex-type, ex-epitype, holotype and ex-neotype strains, respectively.
2
ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; cmdA: partial calmodulin gene.
98
Culture characteristics: Colonies on PDA irregular, reaching Distribution: New Zealand, South Africa and USA.
65–68 mm diam after 14 d at 22 °C, slightly umbonate, colour
Typus: USA, Maryland, Pocomoke City, 38.072952N,
citrine, olivaceous buff to olivaceous, with compact aerial
75.555852W, from branch canker of Cupressus sp., 2017, S.A.
mycelium on the surface, abundant sporulation surrounding
Krueger-Hadfield (epitype of Pestalotia unicornis designated
centre and at the margins of the colony. On CMA circular or
here CBS H-23739, MBT383596, cultures ex-epitype CBS
irregular, reaching 58–59 mm diam after 14 d at 22 °C, flat at the
143871 = CPC 34650, CBS 143872 = CPC 34649, CBS
centre and margins, citrine to olivaceous coloured, with moderate
143873 = CPC 34651); New Jersey, from Chamaecyparis
aerial mycelium on the surface, sporulating abundantly. On MEA
thyoides, 1878, J.B. Ellis (holotype of Pestalotia unicornis IMI
irregular, reaching 35–40 mm diam after 14 d at 22 °C, flat to
5816).
crateriform, slightly sunk into the agar, buff to olivaceous col-
oured at the centre becoming white at the margins, with dense Additional materials examined: New Zealand, from Cryptomeria japonica, 1981,
mycelium on the surface, sporulating near the centre. On SNA H.J. Boesewinkel (CBS H-23151 reference specimen, culture CBS 538.82 = CPC
23783 = NBRC 32684). South Africa, from Cupressus sempervirens, 1999, I.
circular to slightly irregular, reaching 20–21 mm diam after 14 d Barnes (culture CBS 120306 = CMW 5596).
at 22 °C, umbonate, with moderate aerial mycelium, sporulation
abundant between centre and margins. Notes: Seiridium unicorne (basionym: Pestalotia unicornis) is the
earliest described cypress pathogen (Cooke & Ellis 1878) of
Fig. 78. Bayesian phylogenetic tree inferred from ITS (565 bp), gapdh (613 bp) and cmdA (863) using partitioned analysis with N92+G+I substituition model for ITS and gapdh
and GTR+G+I for cmdA. Highest log likelihood -34098.05. The analysis involved 122 nucleotide sequences including 98 sequences obtained from GenBank and 24 sequences
obtained in the present study. Scale bar indicates expected changes per site. The tree was rooted to Alternaria alternata (GV14 634a1). The novel species described in this
study are shown in bold. Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession numbers are indicated in Table 20. T, ET, HT and NT
indicate ex-type, ex-epitype, ex-holotype and ex-neotype strains, respectively. TreeBASE:S23794.
MARIN-FELIX ET AL.
Seiridium and was isolated from “cedar wood” collected in New to consolidate a stable taxonomic concept for this taxon. Given
Jersey (USA). The host was later identified by W.W. Wagener as the occurrence of related species which are pathogenic on the
Chamaecyparis thyoides, as indicated on the holotype label (see same hosts (i.e. S. cancrinum, S. cardinale, S. cupressi and
Bonthond et al. 2018). The genus Seiridium accommodates S. neocupressi) the availability of ex-epitype DNA sequence data
multiple species infecting Cupressaceae and S. unicorne has provides a valuable reference for the identification of future
traditionally been regarded as a mild pathogen but capable of collections. The specimen we introduce here as epitype (CBS H-
infecting a broad range of hosts, including plant species beyond 23739) was collected from necrotic lesions of a Cupressus sp. in
the Cupressaceae (Guba 1961). The holotype (IMI 5816) was Maryland, USA and matches morphologically in all respects with
obtained and examined during a preceding study (Bonthond the holotype (IMI 5816) and the reference strain (CBS 538.82) of
et al. 2018) and found to be limited to two microscope slides. S. unicorne. Furthermore, in the four-locus phylogeny (Fig. 63E)
Therefore, the selection of an epitype for S. unicorne is important the ex-epitype strain clusters under a fully supported node in the
100
Fig. 79. Stemphylium rombundicum (ex-type BRIP 27486). A–J. Conidia. K–S. Verrucose conidiogenous cells and straight and simple conidiophores with conidia attached.
Scale bars: A–N, P = 20 μm; others 100 μm.
MARIN-FELIX ET AL.
Fig. 80. Stemphylium truncatulae (ex-type BRIP 14850). A–H. Asexual morph. A–D. Conidia on SNA. E–H. Straight and simple or multibranched conidiophores and
conidiogenous cells. I–J. Sexual morph. I. Immature pseudothecium. J. Ascomatal wall. Scale bars: I = 100 μm; others 20 μm.
clade that was assigned to S. unicorne based on morphology Septoriella Oudem., Ned. kruidk. Archf, ser. 2, 5: 504. 1889.
(Bonthond et al. 2018). Conidial measurements strongly overlap Fig. 66.
between the selected epitype, holotype and reference strain Synonyms: Allophaeosphaeria Ariyaw. et al., Fungal Diversity
(Fig. 63A–D), although the median width of the epitype being 72: 137. 2015.
slightly narrower in comparison to the holotype and reference Poaceicola W.J. Li et al., Mycosphere 6: 696. 2015.
strain. Measurements of basal and apical appendages and dis- Vagicola Chethana & K.D. Hyde, Fungal Diversity 75: 113. 2015.
tributions of those measurements are highly similar between
epitype and reference strain. For both appendages, the variation
in measurements is higher for the holotype compared to the
reference strain and the epitype, which, however, likely results Type species: Septoriella phragmitis Oudem. Epitype and ex-
from the age and condition of the material as we observed that epitype strain designated by Crous et al. (2015a): CBS H-
conidial appendages from the holotype were often damaged. 22281, CBS 140065.
Authors: G. Bonthond, M. Sandoval-Denis, S.A. Krueger- DNA barcode (genus): LSU. Fig. 28.
Hadfield, L. Morton, C. Ambers & P.W. Crous
102
Fig. 81. Stemphylium waikerieanum (ex-type VPRI 21969). A. Dried leaf of Allium cepa showing leaf lesions caused by the pathogen. B. Dried type culture. C. Revived colony
after 1 wk on PDA. D–K. Asexual morph. D–H. Simple or 1-branched conidiophores on PDA and SNA. I–K. Phaeodictyospores. L–N. Sexual morph. L, M. Immature
ascomata on SNA after 1 wk. N. Ascomatal wall. Scale bars: N = 100 μm; others 20 μm.
MARIN-FELIX ET AL.
Fig. 82. Tubakia spp. A–E. Disease symptoms. A. Tubakia californica on Tanoak tree (Notholithocarpus densiflorus). B. Tubakia californica on California black oak (Quercus
kelloggii). C. Tubakia sierrafriensis (holotype CFNL 2944) on Quercus eduardi. D. Tubakia japonica (epitype NBRC H-11611) on Castanea crenata. E. Tubakia melnikiana
(holotype HAL 3179 F) causing necrotic leaf lesion. F–N. Asexual morph. F. Scutellum of Tubakia paradryinoides (holotype TFM:FPH 3923). G. Central columella of Tubakia
oblongispora (holotype NBRC H-11881). H–J. Conidiophores. H. Tubakia japonica (epitype NBRC H-11611). I. Tubakia oblongispora (holotype NBRC H-11881). J. Tubakia
paradryinoides (holotype TFM:FPH 3923). K–M. Conidia. K. Tubakia dryinoides (holotype NBRC H-11618). L. Tubakia oblongispora (holotype NBRC H-11881). M. Tubakia
paradryinoides (holotype TFM:FPH 3923). N. Microconidia of Tubakia dryinoides (holotype NBRC H-11618). Scale bars: 10 μm. Pictures taken from Braun et al. (2018).
104
Ascomata ostiolate, solitary to gregarious, immersed to proliferations near apex. Conidia cylindrical to sub-
semi-immersed or superficial, broadly ellipsoidal to globose, cylindrical, fusiform, or subfusiform, apex obtuse to sub-
subglobose, or obpyriform, brown to dark brown or black, obtuse, base truncate, straight or curved, euseptate, pale
smooth-walled, coriaceous, uni- to biloculate; necks central, brown to brown, thin-walled, smooth-walled or minutely
flush to papillate, brown to dark brown or black, with or verruculose, bearing mucoid appendages at both ends
without periphyses, rarely comprising short, hyaline setae; (type H sensu Nag Raj 1993) (description of asexual morph
ascomatal wall thin-walled, outer layers composed of brown adapted from Crous et al. 2015a).
to dark brown or blackish cells of textura angularis, inner
Culture characteristics: Colonies with moderate to abundant
layers composed of brown cells of textura prismatica, or of
aerial mycelium, variable in colour, circulate or lobate. On
hyaline or brown cells of textura angularis, rarely composed
PDA surface white to iron-grey or grey olivaceous, greyish
of hyaline gelatinous cells. Hamathecium composed of
white, pale olivaceous grey, or dull green margins, mouse
numerous, 1 – 3 μm wide, filiform to broadly cylindrical,
grey in the middle and pale mouse grey at the center; reverse
septate, cellular pseudoparaphyses, or lacking pseudopar-
grey, olivaceous grey, dull green, buff to dark brown, oliva-
aphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical,
ceous to pale brown or black, brown or olivaceous brown to
broadly cylindrical, cylindrical-fusiform, cylindrical-clavate,
black. On MEA surface white, dirty white, or white to pale
clavate, or broadly clavate, pedicellate, apically rounded
yellow or iron-grey; reverse yellow, umber, buff, or dark grey
with an ocular chamber. Ascospores overlapping, 1 – 3-
to black.
seriate, hyaline to yellowish brown or pale brown, brown,
golden brown, or reddish brown, narrowly or broadly fusi- Optimal media and cultivation conditions: CMA, MEA, OA and
form or oblong to narrowly oblong, straight or slightly PDA at 25 °C.
curved, smooth-walled or echinulate, multi-septate, with
Distribution: Mostly Europe, but also reported in Asia and North
transverse septa, with or without longitudinal septa,
America.
sometimes with enlarged medium cells, constricted or not
at septa, conical or obtuse, rounded ends, with or without Hosts: Mostly saprophytes of grasses (Poaceae), including
sheath. Conidiomata pycnidial, solitary or aggregated, Arundo spp., Agrostis stolonifera, Bromus sterilis, Calama-
immersed to semi-immersed, globose to subglobose, uni- grostis spp., Dactylis glomerata, Elymus glaucus, E. repens,
locular, pale brown to brown or dark brown, with central, Poa sp., Phragmites spp. and Setaria verticillata. Also found
papillate, circular ostiole; conidiomatal wall of brown cells of on Rosa canina (Rosaceae) and Juncus sp. (Juncaceae),
textura angularis, rarely of textura oblita, inner layers and others hosts not molecularly corroborated. Septoriella
becoming hyaline. Conidiophores lining the inner cavity, hirta is considered and important secondary pathogen of
reduced to conidiogenous cells, invested in mucus. Con- grasses, including Agropyron spp., Bromus spp., Dactylis
idiogenous cells ampulliform to lageniform, hyaline, smooth- glomerata, Festuca spp., Poa spp., Stipa spp., and Triticum
walled, proliferating via inconspicuous percurrent spp., among others.
Table 21. DNA barcodes of accepted Tubakia spp.

Species Isolates1 GenBank accession number2 References
ITS rpb2 tef1 tub2
Tubakia americana CBS 129014 MG591873 MG976449 MG592058 MG592152 Braun et al. (2018)
T. braunii CBS 115011T MG591912 MG976488 MG592101 MG592193 Braun et al. (2018)
T. californica CBS 143670T MG591835 MG976451 MG592023 MG592117 Braun et al. (2018)
ET
T. dryina CBS 112097 MG591851 MG976455 MG592039 MG592133 Braun et al. (2018)
T. dryinoides NBRC 9267T MG591878 MG976461 MG592063 MG592157 Braun et al. (2018)
T
T. hallii CBS 129013 MG591880 MG976462 MG592065 MG592159 Braun et al. (2018)
T. iowensis CBS 129012T MG591879 – MG592064 MG592158 Braun et al. (2018)
ET
T. japonica NBRC 9268 MG591886 MG976465 MG592071 MG592165 Braun et al. (2018)
T. liquidambaris CBS 139744 MG605068 – MG603578 – Harrington & McNew (2018)
T. macnabbii CBS 137349 T
MG605069 – MG603579 – Harrington & McNew (2018)
T. melnikiana CPC 32255T MG591893 MG976472 MG592080 MG592174 Braun et al. (2018)
T
T. oblongispora NBRC 9885 MG591897 MG976474 MG592084 MG592178 Braun et al. (2018)
T. paradryinoides NBRC 9884T MG591898 MG976475 MG592085 MG592179 Braun et al. (2018)
IsoT
T. seoraksanensis CBS 127492 MG591908 MG976485 MG592096 MG592188 Braun et al. (2018)
T. sierrafriensis CPC 33020T MG591910 MG976486 MG592099 MG592191 Braun et al. (2018)
IsoT
T. suttoniana CBS 639.93 MG591921 MG976493 MG592110 MG592202 Braun et al. (2018)
T. tiffanyae CBS 137345T MG605081 – – – Harrington & McNew (2018)
1
CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at the Westerdijk Fungal Biodiversity Institute;
NBRC: Biological Resource Center, NITE, Chiba, Japan. T, ET and IsoT indicate ex-type, ex-epitype and ex-isotype strains, respectively.
2
MARIN-FELIX ET AL.
106
Disease symptoms: Secondary foot rot and rot of mature straw. to the two former species of the genus, while V. dactylidis pro-
Discoloured culms and predisposition of the plant to premature duces ascospores with transverse and longitudinal septa as
collapse. seen in species of Allophaeosphaeria, indicating again that the
longitudinal septation of ascospores is not phylogenetically
Notes: Septoriella was considered an asexual genus charac-
informative. Vagicola chlamydospora was recently transferred to
terised by pycnidial, unilocular conidiomata, and cylindrical to
Septoriella based on phylogenetic data (Jayasiri et al. 2015).
fusoid, euseptate conidia bearing mucoid appendages at both
Surprisingly, the asexual morph reported in that species does not
ends (Crous et al. 2015a). However, in our phylogenetic studies
fit with the morphology of Septoriella, since it produces micro- to
based on ITS and LSU, and on the combined dataset, the ex-
macronematous conidiophores and chlamydospore-like conidia.
type strains of the sexual genera Allophaeosphaeria, Poacei-
Recently, Thambugala et al. (2017) introduced the last species of
cola and Vagicola were located in the clade representing the
the genus, V. arundinis, which produces both morphs and is
genus Septoriella. Therefore, these genera are synonymised
characterised by ascospores with transverse septa and an
with Septoriella in the present study.
asexual morph similar to Septoriella, which demonstrates the link
Allosphaeosphaeria was recently introduced by Liu et al.
of Vagicola with Septoriella. This last species was invalid
(2015) to incorporate two new saprophytic species found on
because two holotypes were designated. Therefore, this taxon is
Dactylis glomerata from Italy, i.e. Al. dactylidis and Al. mur-
validated in the present study.
iformia, the latter designated as type species. These species
Moreover, in our phylogenetic analyses, the ex-type strain
only produce the sexual morph characterised by ascospores with
of the most recently described species of Neostagonospora
transverse and longitudinal septa, and a gelatinous sheath.
was located in the clade representing Septoriella. This spe-
Subsequently, three other new species were introduced in the
cies is characterised by the production of conidia that are
genus, i.e. Al. clematidis, Al. cytisi and Al. subcylindrospora.
subcylindrical or fusiform, euseptate, with a subobtuse apex
Allosphaeosphaeria clematidis and Al. cytisi only produce the
and truncate base. However, the presence of mucoid ap-
sexual morph, while Al. subcylindrospora only produces an
pendages at both ends, as the other species of Septoriella,
asexual morph. The morphology of this asexual morph fits
has not been reported. Septoriella artemisiae is saprobic or
perfectly in the description of Septoriella, corroborating the
weakly necrotrophic on dead and dying stems of Artemisia
synonymy proposed in the present study based on the phylo-
austriaca.
genetic data. Allosphaeosphaeria clematidis was recently
Most of the species now included in Septoriella are sapro-
excluded from the genus and transferred to the new genus
phytes, except for Sep. hirta, which is an important secondary
Embarria (Wanasinghe et al. 2018). Moreover, in our phyloge-
pathogen of grasses (Sprague 1950). This species is often found
netic studies, Al. cytisi formed an independent lineage in
in association with other fungi such as Gaeumannomyces gra-
Phaeosphaeriaceae far from the clade representing Septoriella.
minis (Johnston et al. 2014) and Oculimacula yallundae causing
Therefore, a new genus is proposed to accommodate this
foot rot of wheat (Crous et al. 2003, 2015a). Other disease
species.
symptoms observed in plants affected by Sep. hirta are dis-
The genus Poaceicola was introduced by Li et al. (2015) to
coloured culms and predisposition to premature collapse,
accommodate Phaeosphaeria elongata and two new species,
especially in rainy and windy seasons, since this species pro-
i.e. Po. arundinis and Po. bromi. The two latter species are
duces a weakness in the culms of plants with ripe grains. The
characterised by the production of an asexual morph similar to
result of all these symptoms resulted in the increasing of the cost
Septoriella. Poaceicola elongata produces a sexual morph
of harvesting and decreasing of the grain quality (Sprague 1950).
characterised by ascospores with transverse septa. The
presence of only transverse septa could be a morphologic References: Sprague 1950 (pathogenicity), Crous et al. 2014b,
difference from Allophaeosphaeria. However, seven more new Ariyawansa et al. 2015a, Li et al. 2015, Liu et al. 2015,
species have been included in the genus, including one spe- Thambugala et al. 2017 (morphology and phylogeny).
cies presenting ascospores with transverse and longitudinal
Septoriella agrostina (Mapook et al.) Y. Marín & Crous, comb.
septa, Po. arundinis, demonstrating that the longitudinal sep-
nov. MycoBank MB829676.
tation of the ascospores is not phylogenetically informative in
Basionym: Poaceicola agrostina Mapook et al., Fungal Diversity
these genera.
89: 132. 2018.
Vagicola was recently introduced by Ariyawansa et al.
(2015a), in the same year as the other two genera. Description and illustration: Wanasinghe et al. (2018).
Ariyawansa et al. (2015a) raised the subgenus Vagicola
Septoriella artemisiae (Wanas. et al.) Y. Marín & Crous, comb.
(Shoemaker & Babcock 1989) to generic rank to accommodate
Phaeosphaeria vagans, a species characterised by a sexual
Basionym: Neostagonospora artemisiae Wanas. et al, Fungal
morph similar to the species of Poaceicola, having ascospores
Diversity 89: 130. 2018.
with transverse septa only. Subsequently, Jayasiri et al. (2015)
introduced two new species: V. chlamydospora, which pre- Description and illustration: Wanasinghe et al. (2018).
sents both morphs, and V. dactylidis, which produces only the
Septoriella arundinicola (Wanas. et al.) Y. Marín & Crous,
sexual morph. The sexual morph of V. chlamydospora is similar
Fig. 83. RAxML phylogram obtained from the combined ITS (534 bp), LSU (741 bp) and rpb2 (739 bp) sequence alignment of all accepted species of Zasmidium. The tree was
rooted to Anellosympodiella juniperi CBS 137992 and Neopenidiella nectandrae CBS 734.78. The novelties proposed in this study are indicated in bold. RAxML bootstrap
support (BS) values above 70 % and Bayesian posterior probability scores above 0.95 are shown at the nodes. GenBank accession numbers are listed in Table 22 and
Videiraet al. (2017). T, ET, HT and NT indicate ex-type, ex-epitype, holotype and ex-neootype strains, respectively. TreeBASE: S23834.
MARIN-FELIX ET AL.
Table 22. DNA barcodes of accepted Zasmidium spp.

ITS LSU rpb2 act tef1 tub2
Zasmidium angulare CBS 132094T JQ622088 JQ622096 MF951690 – JQ622113 – Li et al. (2012), Videira et al. (2017)
Z. anthuriicola CBS 118742T FJ839626 FJ839662 MF951691 – KF253229 KF252763 Crous et al. (2009a), Quaedvlieg et al. (2013),
Videira et al. (2017)
Z. aporosae MFLU 12-2206HT KC677912 – – – – – Phengsintham et al. (2013)
Z. arcuatum CBS 113477 T
EU041779 EU041836 MF951692 – – – Arzanlou et al. (2007), Videira et al. (2017)
Z. aucklandicum CPC 13569 MF951409 MF951280 MF951733 – – – Videira et al. (2017)
Z. biverticillatum CBS 335.36 EU041796 EU041853 – – – – Arzanlou et al. (2007)
Z. cellare CBS 146.36NT EU041821 EU041878 MF951693 – – – Arzanlou et al. (2007), Videira et al. (2017)
Z. cerophillum CBS 103.59T EU041798 GU214485 MF951694 – – – Arzanlou et al. (2007), Crous et al. (2009b),
Z. citri-griseum CBS 139467ET KF901796 KF902155 KF902518 – KF903385 KF903079 Quaedvlieg et al. (2014)
Z. commune CBS 142530T KY979765 KY979820 – KY979860 – – Crous et al. (2017a)
Z. cyatheae CPC 24725 T
KT037530 KT037571 – KT037629 KT037490 – Guatimosim et al. (2016)
Z. dasypogonis CBS 143397T MG386048 MG386101 – – – – Crous et al. (2017b)
Z. daviesiae CBS 116002 FJ839633 FJ839669 MF951698 KF903477 KF903373 KF903069 Crous et al. (2009a), Quaedvlieg et al. (2014),
Z. ducassei BRIP 53367T HQ149687 – – – – – Shivas et al. (2011)
CPC 32929 MK539971 MK540041 – MK540233 MK540167 - Present study
Z. elaeocarpi CBS 142187T MF951398 MF951263 MF951699 – – – Videira et al. (2017)
Z. eucalypticola CBS 142186 T
MF951400 MF951265 MF951701 – – – Videira et al. (2017)
Z. eucalyptigenum CBS 138860T KP004458 KP004486 – KT037630 – – Crous et al. (2014c), Guatimosim et al. (2016)
Z. eucalyptorum CBS 118500T KF901652 MF951266 MF951702 KF903495 KF903101 – Quaedvlieg et al. (2014), Videira et al. (2017)
Z. fructicola CBS 139625T KP896052 KP895922 MF951703 KP896003 KP896099 – Huang et al. (2015), Videira et al. (2017)
T
Z. fructigenum CBS 139626 KP896056 KP895926 MF951704 KP896007 KP896103 KP896149 Huang et al. (2015), Videira et al. (2017)
Z. gahniicola CBS 143422T MG386050 MG386103 – – – – Crous et al. (2017b)
Z. grevilleae CBS 124107 T
FJ839634 FJ839670 MF951705 – – – Crous et al. (2009a), Videira et al. (2017)
Z. gupoyu CBS 122099 MF951401 MF951267 MF951706 – – – Videira et al. (2017)
Z. hakeae CBS 142185 T
Z. indonesianum CBS 139627T KF901739 KF902086 MF951710 – KF903377 – Quaedvlieg et al. (2014), Videira et al. (2017)
Z. iteae CBS 113094 T
Z. lonicericola CBS 125008ET KF251283 KF251787 MF951712 – KF253231 KF252765 Quaedvlieg et al. (2013), Videira et al. (2017)
Z. macluricola BRIP 52143T GU108499 – – – – – Shivas et al. (2009)
Z. musae CBS 122477T EU514291 – – EU514346 – – Arzanlou et al. (2008)
Z. musae-banksii CBS 121710T EU041795 EU041852 MF951716 – – – Arzanlou et al. (2007), Videira et al. (2017)
Z. musicola CBS 122479T EU514294 MF951275 MF951717 – – – Arzanlou et al. (2008), Videira et al. (2017)
Z. musigenum CBS 365.36 T
EU041801 EU041858 – – – – Arzanlou et al. (2007)
Z. nocoxi CBS 125009T KF251284 KF251788 MF951719 – KF253232 KF252766 Quaedvlieg et al. (2013), Videira et al. (2017)
Z. pitospori CBS 122274 MF951406 MF951276 MF951720 – – – Videira et al. (2017)
Z. podocarpi CBS 142529T KY979766 KY979821 – KY979861 – KY979930 Crous et al. (2017a)
Z. proteacearum CBS 116003 FJ839635 FJ839671 MF951721 KF903478 – KF903070 Crous et al. (2009a), Quaedvlieg et al. (2014),
Z. pseudoparkii CBS 110999T DQ303023 JF700965 MF951723 KF903419 KF903273 KF902977 Crous et al. (2006), Quaedvlieg et al. (2011,
2013), Videira et al. (2017)
Z. pseudotsugae rapssd EF114687 EF114704 – – – – Winton et al. (2007)
Z. pseudovespa CBS 121159T MF951407 KF901836 MF951724 KF902812 Quaedvlieg et al. (2013, 2014), Videira et al.
(2017)
Z. queenslandicum CBS 122475T EU514295 MF951277 MF951725 – – – Arzanlou et al. (2008), Videira et al. (2017)
Z. rothmanniae CBS 137983 T
KJ869135 MH878613 – – – – Crous et al. (2014c), Vu et al. (2019)
Z. scaevolicola CBS 127009T KF251285 KF251789 MF951726 – KF253233 KF252767 Quaedvlieg et al. (2013), Videira et al. (2017)
Z. schini CBS 142188T MF951408 MF951278 MF951727 – – – Videira et al. (2017)
108

ITS LSU rpb2 act tef1 tub2
Z. strelitziae CBS 121711T EU041803 EU041860 MF951729 – – – Arzanlou et al. (2007), Videira et al. (2017)
Z. suregadae MFLU 12-2212HT KC677914 KC677939 – – – – Phengsintham et al. (2013)
Z. syzygii CBS 133580 T
KC005777 KC005798 MF951730 – – – Crous et al. (2012), Videira et al. (2017)
Z. thailandicum CBS 145027T MK539970 MK540040 – – – – Present study
Z. tsugae ratstk EF114688 EF114705 – – – – Winton et al. (2007)
Z. velutinum CBS 101948ET EU041781 EU041838 MF951731 – – – Arzanlou et al. (2007), Videira et al. (2017)
T
Z. xenoparkii CBS 111185 DQ303028 JF700966 MF951732 KF903438 KF903274 KF902978 Crous et al. (2006), Quaedvlieg et al. (2011,
2014), Videira et al. (2017)
1
BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of
Pedro Crous, housed at the Westerdijk Fungal Biodiversity Institute; MFLU: Mae Fah Luang University herbarium, Chiang Rai, Thailand; rapssd and ratstk were not
specified in the original publications. T, ET, HT and NT indicate ex-type, ex-epitype, holotype and ex-neotype strains, respectively.
2
subunit gene; act: partial actin gene; tef1: partial translation elongation factor 1-alpha gene; tub2: partial β-tubulin gene.
Basionym: Poaceicola arundinicola Wanas. et al., Fungal Di- Septoriella garethjonesii (Thambug. et al.) Y. Marín & Crous,
versity 89: 135. 2018. comb. nov. MycoBank MB829685.
Basionym: Poaceicola garethjonesii Thambug et al., Mycosphere
Description and illustration: Wanasinghe et al. (2018).
8: 756. 2017.
Septoriella arundinis (W.J. Li et al.) Y. Marín & Crous, comb.
Description and illustration: Thambugala et al. (2017).
Basionym: Poaceicola arundinis W.J. Li et al., Mycosphere 6: Septoriella germanica Crous, R.K. Schumach. & Y. Marín, sp.
698. 2015. nov. MycoBank MB829701. Fig. 68.
Description: Li et al. (2015). Etymology: Name refers to Germany, from where this fungus
was isolated.
Septoriella bromi (Wijayaw. et al.) Y. Marín & Crous, comb.
nov. MycoBank MB829680. Conidiomata solitary, pycnidial, erumpent, globose, brown,
Basionym: Poaceicola bromi Wijayaw. et al., Mycosphere 6: 698. 180–220 μm diam, in vivo gregarious, caespitose or in rows,
2015. but also pseudostromatic, up to 300 μm diam, with central
ostiole, 30–40 μm diam; conidiomatal wall of 3–4 layers of
Description and illustration: Li et al. (2015).
brown cells of textura angularis. Conidiophores reduced to
Septoriella dactylidicola Y. Marín & Crous, nom. nov. Myco- conidiogenous cells lining the inner cavity, hyaline, smooth-
Bank MB829681. walled, globose to ampulliform, phialidic, 4–6 × 4–5 μm.
Replaced synonym: Poaceicola dactylidis Tibpromma et al., Conidia solitary, scolecosporous, fusoid to subcylindrical, apex
Mycosphere 8: 755. 2017, non Septoriella dactylidis (Wanas. subobtuse, base truncate, straight to sligthly curved, 3–6-
et al.) Y. Marín & Crous. 2019. septate, golden-brown, smooth-walled, granular with mucoid
caps at each end, (35–)37–42(–46) × 3(–3.5) μm, in vivo
29–46 × 3–4.5 μm.
Septoriella dactylidis (Wanas. et al.) Y. Marín & Crous, comb.
aerial mycelium, covering dish in 2 wk. On MEA, PDA and OA
Basionym: Vagicola dactylidis Wanas. et al., Phytotaxa 6: 725.
surface olivaceous grey, reverse iron-grey.
2015.
Typus: Germany, near Berlin, on dead culm of Phragmites
Description and illustration: Jayasiri et al. (2015).
australis (Poaceae), 16 Apr. 2016, R.K. Schumacher, HPC 1168
Septoriella elongata (Wehm.) Y. Marín & Crous, comb. nov. (holotype CBS H-23875, culture ex-type CBS 145372 = CPC
MycoBank MB829683. 30511).
Basionym: Leptosphaeria elongata Wehm., Mycologia 44: 633.
Notes: Septoriella germanica is related to Sep. artemisiae.
1952.
However, both species differ in the size of the conidia
Synonym: Poaceicola elongata (Wehm.) W.J. Li et al., Myco-
[15–25 × 2–2.5 μm in Sep. artemisiae vs. (35–)
sphere 6: 701. 2015.
37–42(–46) × 3(–3.5) μm in Sep. germanica], as well as in the
Description and illustration: Wehmeyer (1952). conidial septation (2–4 in Sep. artemisiae vs. 3–6 μm in Sep.
germanica). Septoriella germanica was isolated from Phragmites
Septoriella forlicesenica (Thambug. et al.) Y. Marín & Crous,
australis (Poaceae), while Sep. artemisiae was found on Arte-
misia austriaca (Asteraceae).
Basionym: Poaceicola forlicesenica Thambug et al., Mycosphere
8: 756. 2017. Septoriella hibernica Crous, Quaedvl. & Y. Marín, sp. nov.
MycoBank MB829703.
MARIN-FELIX ET AL.
Etymology: Name refers to Ireland, where this fungus was Basionym: Poaceicola italica Thambug. et al., Mycosphere 8:
collected. 759. 2017.
Culture sterile. Septoriella hibernica differs from its closest Description: Thambugala et al. (2017).
phylogenetic neighbour, Septoriella subcylindrispora by unique
Septoriella muriformis (Ariyaw. et al.) Y. Marín & Crous, comb.
fixed alleles in the ITS locus based on the alignment deposited in
TreeBASE (S23834): positions 5 (T), 33 (T), 34 (A), 46 (T), 61
Basionym: Allophaeosphaeria muriformis Ariyaw. et al., Fungal
(A), 89 (T), 477 (A), 479 (T), 480 (A), 512 (T), 528 (G), 534 (G).
Diversity 72: 137.
Culture characteristics: Colonies erumpent, spreading, covering
Descriptions and illustrations: Liu et al. (2015), Thambugala et al.
dish in 2 wk, with fluffy aerial mycelium and even margins. On
(2017).
MEA, PDA and OA surface and reverse olivaceous grey.
Septoriella neoarundinis Y. Marín & Crous, nom. nov. Myco-
Typus: Ireland, on unidentified grass species (Poaceae), Mar.
Bank MB829706.
2014, W. Quaedvlieg (holotype CBS H-23874, culture ex-type
Replaced synonym: Vagicola arundinis Phukhams., Camporesi
CBS 145371 = CPC 24290).
& K.D. Hyde, sp. nov. MycoBank MB831056, non Septoriella
Notes: Septoriella hibernica remained sterile on all media tested. arundinis (W.J. Li et al.) Y. Marín & Crous. 2019.
It is related to Sep. subcylindrispora, but the ITS sequences of Synonym: Vagicola arundinis Phukhams. et al. Mycosphere 8:
both species showed only 97.75 % of nucleotide similarity 763. 2017. (nom. inval., Art. 40).
(Identities = 522/538, 3 gaps).
Etymology: Name reflects the host genus Arundo from which it
Septoriella hollandica Crous, Quaedvl. & Y. Marín, sp. nov. was isolated.
Etymology: Name refers to the Netherlands, where this fungus
Typus: Italy, Province of Marsignano, Predappio, on a dead stem
was collected.
of Arundo plinii (Poaceae), 10 Nov. 2014, E. Camporesi IT 2223A
Ascomata solitary, erumpent, globose, brown, 150–180 μm diam (holotype MFLU 17-0016, ex-type living culture MFLUCC 15-
with central ostiole; conidiomatal wall of 3–4 layers of brown 0027).
cells of textura angularis. Pseudoparaphyses hyphae-like, hya-
Notes: This species was initially introduced by Thambugala et al.
line, smooth-walled, branched, septate, 1.5–2 μm diam. Asci
(2017) as Vagicola arundinis. However, it was invalid since two
subcylindrical, flexuous, bitunicate, with well-defined apical
different holotype numbers were cited. Therefore, this species is
chamber, 1–1.5 μm diam, fasciculate, short stipitate,
validated here and a new name in Septoriella is proposed, using
70–90 × 8–10 μm. Ascospores bi- to triseriate, fusoid-ellipsoid,
a new epithet since Sep. arundinis is occupied.
5-septate, constricted at median septum, medium brown,
smooth-walled, guttulate, widest above median septum, (27–) Septoriella neodactylidis Y. Marín & Crous, nom. nov.
28–30(–32) × (4–)4.5(–5) μm. MycoBank MB829707.
Replaced synonym: Allophaeosphaeria dactylidis Wanas. et al.,
Culture characteristics: Colonies flat, spreading, covering dish in
Fungal Diversity 72: 137. 2015, non Septoriella dactylidis
2 wk with moderate aerial mycelium. On MEA surface vinaceous
(Wanas. et al.) Y. Marín & Crous. 2019.
buff, reverse sienna; on PDA surface isabelline, reverse hazel;
on OA surface saffron. Description and illustration: Liu et al. (2015).
Typus: The Netherlands, Oosterbeek, on leaves of Phrag- Septoriella pseudophragmitis Crous, Quaedvl. & Y. Marín, sp.
mites australis (Poaceae), 24 Jan. 2014, W. Quaedvlieg (ho- nov. MycoBank MB829708. Fig. 70.
lotype CBS H-23877, culture ex-type CBS 145374 = CPC
Etymology: Name refers to its morphological similarity with Sep.
24109).
phragmitis, which occurs on the same host.
Notes: Septoriella hollandica is related to Sep. chlamydospora
Conidiomata solitary, pycnidial, erumpent, globose, brown-black,
and Sep. tridentina. Septoriella hollandica, as well as Sep.
200–250 μm diam with central ostiole; conidiomatal wall of 6–8
chlamydospora and Sep. tridentina, produce sexual morphs in
layers of brown textura angularis. Conidiophores reduced to
culture. Septoriella hollandica can be easily distinguished from
conidiogenous cells lining the inner cavity, hyaline, smooth-
Sep. chlamydospora by its 5-septate ascospores, being 9-
walled, globose to ampulliform, phialidic, 4–10 × 4 μm. Con-
septate in Sep. chlamydospora. Septoriella tridentina is the
idia solitary, subcylindrical, golden-brown, guttulate, smooth-
only species of this complex that produces ascospores sur-
walled, apex obtuse, base truncate, 3(–6)-septate, with
rounded by a mucilaginous sheath. The asexual morph was
mucoid caps at each end, (20–)24–28(–32) × (3–)3.5(–4) μm.
reported only for Sep. chlamydospora. However, as it was
mentioned above, the asexual morph described in Sep. chla- Culture characteristics: Colonies erumpent, spreading, covering
mydospora (Jayasiri et al. 2015) corresponds to chains of dish after 2 wk at 25 °C, with moderate aerial mycelium and feathery
chlamydospores instead of scolecosporous conidia typical of margins. On MEA surface olivaceous grey, reverse iron-grey.
Septoriella. Septoriella hollandica was isolated from Phragmites
Typus: The Netherlands, on leaves of Phragmites sp. (Poa-
australis, while the other two species were found on Dactylidis
ceae), 5 Mar. 2014, W. Quaedvlieg (holotype CBS-H 23904,
spp.
culture ex-type CBS 145417 = CPC 24166).
Septoriella italica (Thambug. et al.) Y. Marín & Crous, comb.
Notes: Septoriella pseudophragmitis is similar to Sep. phragmi-
tis, which is reported from the same host, Phragmites (Poaceae).
110
These species differ in the size of their conidiomata (up to located in an independent lineage distant to the clade repre-
250 μm diam in Sep. pseudophragmitis vs. 350 μm diam in Sep. senting the genus Septoriella. Moreover, based on a megablast
phragmitis) and conidia [(20–)24–28(–32) × (3–)3.5(–4) μm in search using the ITS sequence, the closest matches in NCBIs
Sep. pseudophragmitis vs. (29–)32–40(–46) × 3(–3.5) μm diam GenBank nucleotide database were Ophiosimulans tanaceti
in Sep. phragmitis], as well as in the conidial septation, being [GenBank KU738890; Identities = 534/586 (91 %), 11 gaps
mostly 3-septate in Sep. pseudophragmitis and 5-septate in Sep. (1 %)], Ophiobolus cirsii [GenBank KM014664; Identities = 514/
phramitis. Based on our phylogenetic analysis, Septoriella 566 (91 %), 22 gaps (1 %)], and Chaetosphaeronema hispidulum
pseudophragmitis is related to Sep. allojunci. However, Sep. [GenBank KX096655; Identities 535/588 (91 %), 22 gaps (3 %)].
allojunci produces smaller conidiomata (up to 150 μm) and larger Arezzomyces cytisi is a saprobe found on dead herbaceous
conidia (48–70 × 3–6.6 μm). Moreover, Sep. allojunci was branches of Cytisus.
isolated from Juncus (Juncaceae).
Arezzomyces cytisi (Wanas. et al.) Y. Marín & Crous, comb.
Septoriella rosae (Mapook et al.) Y. Marín & Crous, comb. nov. nov. MycoBank MB829712.
MycoBank MB829713. Basionym: Allophaeosphaeria cytisi Wanas. et al., Fungal Di-
Basionym: Poaceicola rosae Mapook et al., Fungal Diversity 89: versity 75: 97. 2015.
136. 2018.
Description and illustration: Ariyawansa et al. (2015a).
Description and illustration: Wanasinghe et al. (2018).
Typus: Italy, Arezzo Province, Casuccia di Micheli in Quota,
Septoriella subcylindrospora (W.J. Li et al.) Y. Marín & Crous, dead and hanging branches of Cytisus sp. (Fabaceae), 20 Jun.
comb. nov. MycoBank MB829709. 2012, E. Camporesi (holotype MFLU 15-1502, culture ex-type
Basionym: Allophaeosphaeria subcylindrospora W.J. Li et al., MFLUCC 15-0649).
Fungal Diversity 75: 100. 2015.
Authors: Y. Marin-Felix, W. Quaedvlieg, R.K. Schumacher & P.W.
Description and illustration: Ariyawansa et al. (2015a). Crous
Septoriella vagans (Niessl) Y. Marín & Crous, comb. nov. Setophoma Gruyter et al., Mycologia 10: 1077. 2010. Fig. 71.
MycoBank MB829710.
Basionym: Pleospora vagans Niessl, Verh. nat. Ver. Brünn 14:
174. 1876.
Synonym: Vagicola vagans (Niessl) O.E. Erikss. et al., Fungal Type species: Setophoma terrestris (H.N. Hansen) Gruyter et al.,
Diversity 75: 115. 2015. basionym: Phoma terrestris H.N. Hansen. Lectotype and ex-
lectotype strain designated by de Gruyter et al. (2010): CBS
Description and illustration: Jayasiri et al. (2015).
H-20311, CBS 335.29.
Arezzomyces Y. Marín & Crous, gen. nov. MycoBank
DNA barcode (genus): LSU. Fig. 28.
MB829711.
Etymology: Name reflects the Italian province Arezzo where it
was collected. Ascomata scattered, gregarious, immersed, visible as raised,
minute black dots on host surface, uniloculate, globose to sub-
Ascomata solitary, scattered, immersed to erumpent, obpyriform, globose, glabrous, brown to dark brown, ostiole central, with
dark brown to black, coriaceous, with ostiole filled with hyaline minute papilla; ascomatal wall thin, of equal thickness,
cells, appearing as a white ring around ostiole; necks papillate, composed of pseudoparenchymatous cells, arranged in flattened
black, smooth; ascomatal wall comprising 6–8 layers, outer layer textura angularis to textura prismatica. Hamathecium composed
heavily pigmented, comprising blackish to dark brown, thick-walled of numerous, filiform, broad cellular pseudoparaphyses, with
cells of textura angularis, inner layer composed of brown, thin- distinct septa, anastomosing at apex. Asci 8-spored, bitunicate,
walled cells of textura angularis. Hamathecium comprising fissitunicate, cylindrical to cylindric-clavate, short pedicellate,
numerous, filamentous, branched, septate pseudoparaphyses. apically rounded, with well-developed narrowly ocular chamber.
Asci 8-spored, bitunicate, fissitunicate, cylindrical, pedicel furcate, Ascospores overlapping, 2-seriate, phragmosporous, cylindrical
rounded and thick-walled at the apex, with an ocular chamber. to cylindrical-clavate, hyaline, 3-septate, usually enlarged at the
Ascospores mostly uniseriate, initially hyaline, becoming yellowish second cell from apex, smooth-walled with large guttules. Con-
brown at maturity, ellipsoidal, muriform, with 6–8 transverse septa, idiomata pycnidial, solitary to confluent, superficial or submerged
3–7 vertical septa, strongly constricted at the central septa, weakly in agar, globose to subglobose, setose, with papillate necks,
constricted at the other septa, with conical and narrowly rounded honey to olivaceous or olivaceous black, with 2–7(–11) layers of
ends, lacking a mucilaginous sheath. Asexual morph not observed. pseudoparenchymatal cells. Conidiogenous cells hyaline,
Culture characteristics: Colonies spreading, surface erumpent, monophialidic. Conidia aseptate, ellipsoidal to subcylindrical or
with moderate aerial mycelium, and feathery margins. On MEA, subfusoid, guttulate (adapted from Quaedvlieg et al. 2013,
PDA and OA surface dirty white; reverse dirty white to luteous. Phookamsak et al. 2014a).
Type species: Arezzomyces cytisi (Wanas. et al.) Y. Marín & Culture characteristics: Colonies spreading with sparse to
Crous. Holotype and ex-type cultures: MFLU 15-1502, MFLUCC moderate aerial mycelium, smooth or folded surface, even or
15-0649. lobate margins. On PDA surface iron-grey or grey olivaceous
with outer region iron-grey; reverse olivaceous grey or iron-grey.
Notes: Arezzomyces is introduced to accommodate Allo- On MEA surface olivaceous grey or umber with patches of
phaeosphaeria cytisi since, based on phylogenetic data, it is apricot and dirty white; reverse ochreous or cinnamon with
MARIN-FELIX ET AL.
patches of olivaceous grey. On OA surface isabelline or iron-grey 637 (T), 638 (A); ITS positions 32 (C), 34 and 35 (indels), 36 (T),
surrounded by orange to apricot diffuse pigment layer in agar. 37 (T), 38 (T), 42 (G), 43 (T), 44 (A), 54 (C), 56 (G), 57 (T), 58
(T), 59 (C), 60 (G), 61 (C), 62 (T), 63 (G), 64 (T), 66 (G), 67 (T),
Optimal media and cultivation conditions: OA and SNA at 25 °C
72 (T), 77 (G), 78 (T), 80 (T), 99 (T), 100 (G), 101 (A), 103 (C),
under continuous near-ultraviolet light to promote sporulation.
114 (C), 117 (G), 118 (T), 119 (A), 121 (C), 122 (T), 124 (C), 130
Sterile bamboo pieces on WA to induce sporulation of the sexual
(A) 138 (C), 140 (A), 143 (T), 146 (A), 148 (C), 172 (A), 176 (T),
morph.
178 (A), 180 (T), 182 (A), 186 (indel), 354 (T), 379 (T), 381
Distribution: Worldwide. (indel), 388 (T), 389 (G), 390 (G), 391 (T), 392 (C), 393 (C), 394
(T), 395 (C), 396 (T), 399 (G), 400 (A), 401 (C), 402 (C), 409 (A),
Hosts: Mainly on members of Poaceae, but also of Amar-
418 (A), 419 (T), 433 (G), 434 (T), 435 (A), 441 (G), 444 (T), 467
yllidaceae and Asteraceae, among others.
(A), 470 (indel), 473 (T), 475 (C), 477 (A), 478 (C), 479 (T), 482
Disease symptoms: Leaf spots and necrosis, leaf dieback, and (A), 485 (C), 486 (C), 490 (A), 495–498 (indels), 499 (C), 500
pink root. (C), 502 (T), 504 (A), 506 (T), 507 (A), 511 (C).
Notes: Setophoma was introduced by de Gruyter et al. (2010) to Culture characteristics: Colonies flat, spreading, with moderate
accommodate two species previously placed in Pyrenochaeta, aerial mycelium and even, lobate margins, reaching 60 mm diam
i.e. Pyr. sacchari and Pyr. terrestris. Setophoma is characterised after 2 wk. On MEA, PDA and OA, surface and reverse oliva-
by pycnidial conidiomata covered by setae, phialidic con- ceous grey.
idiogenous cells, and hyaline, ellipsoidal to subcylindrical,
Typus: Belgium, Dinant, 173 m a.s.l., on border of calcareous
aseptate, guttulate conidia (de Gruyter et al. 2010, Quaedvlieg
meadow, on a dead and attached leaf of Brachypodium syl-
et al. 2013). When Setophoma was introduced, the sexual
vaticum (Poaceae), 2 Nov. 2016, L. Bailly & R.K. Schumacher,
morph had not been observed. Subsequently, Phookamsak et al.
HPC 1503, RKS 1 (holotype CBS H-23905, culture ex-type CBS
(2014a) reported the sexual morph of this genus. It was found
145418 = CPC 32492).
causing leaf spots of sugarcane (Saccharum officinarum), and
based on the phylogenetic data it was shown to be the sexual Notes: Setophoma brachypodii remained sterile on all media
morph of Set. sacchari. This sexual morph is similar to tested, and the original specimen was depleted, hence we could
Phaeosphaeria species, producing ascospores with three septa not describe it based on morphology. This is the first species of
with the second cell from the apex being swollen; these cells Setophoma reported on Brachypodium.
differ in colour (hyaline in Setophoma vs. yellowish to brown in
Setophoma pseudosacchari Crous & Y. Marín, sp. nov.
Phaeosphaeria).
Setophoma encompasses pathogenic or saprobic species
associated with monocotyledonous plants (de Gruyter et al. Etymology: Named after its closely phylogenetic relation to
2010). The type species, Set. terrestris, causes pink root on Setophoma sacchari.
Allium spp., and also on Zea mays and Oryza sativa, but it is
asymptomatic on other hosts (Farr & Rossman 2019). Seto- Ascomata developing on OA, solitary, erumpent, brown,
phoma vernoniae produces leaf spots on Vernonia polyanthes 200–300 μm diam, globose, with large central ostiole, 30–40 μm
(Crous et al. 2014b), while Set. sacchari is considered a weak diam; ascomatal wall of 3–4 layers of brown cells of textura
pathogen of members of the Poaceae that is only noticeable angularis, ascomata setose; setae brown, flexuous, thick-walled,
when conditions are favourable for disease spread, and causes septate, base verruculose, with slight taper to obtuse apex, up to
leaf spots and necrosis and leaf dieback (Farr & Rossman 2019). 150 μm long. Pseudoparaphyses hyphae-like, anastomosing,
In our phylogenetic analysis based on ITS and LSU (Fig. 28), branched, septate, hyaline, occurring intermingled among asci.
the clade representing the genus Setophoma is well-supported Asci bitunicate, ellipsoid to subcylindrical, hyaline, curved to
(95 % BS / 1 PP). However, the most recently described spe- straight, fasciculate, apex obtuse, with well-defined ocular
cies, Set. cyperi, is not located in that clade, representing a new chamber, 2 μm diam, stipitate, 70–100 × 10–13 μm. Ascospores
genus in the family Phaeosphaeriaceae. bi- to triseriate, fusoid-ellipsoid with subobtuse ends, straight, 3-
At the proof stage of this paper, a new publication appeared septate, widest in second cell from apex, prominently guttulate,
on Setophoma (Liu et al. 2019), which contains four new species. hyaline, smooth-walled, (22–)25–30 × (5.5–)6 μm. Conidiomata
developing on SNA, solitary to aggregated, erumpent, brown,
References: de Gruyter et al. 2010, Quaedvlieg et al. 2013, globose, 200–300 μm diam, with 1–2 ostioles, lacking setae;
Phookamsak et al. 2014a (morphology and phylogeny). conidiomatal wall of 2–3 layers of brown textura angularis. Co-
Setophoma brachypodii Crous, R.K. Schumach. & Y. Marín, nidiophores reduced to conidiogenous cells lining inner cavity,
sp. nov. MycoBank MB829669. dissolving at maturity, hyaline, smooth-walled, globose to
ampulliform, phialidic, 4–6 × 5–6 μm. Conidia solitary, aseptate,
Etymology: Name reflects the host genus Brachypodium from straight to slightly curved, subcylindrical to fusoid-ellipsoid, apex
which it was isolated. obtuse, base truncate, hyaline, smooth-walled, guttulate, (8–)
Culture sterile. Setophoma brachypodii differs from its closest 11–12(–14) × (3–)4 μm.
phylogenetic neighbour, Setophoma terrestris by unique fixed Culture characteristics: Colonies erumpent, spreading, surface
alleles in two loci based on alignments of the separate loci folded, with moderate aerial mycelium and even, lobate margins,
deposited in TreeBASE (S23834): LSU positions 42 (G), 67 (C), reaching 55 mm diam. On MEA surface peach, outer region
75 (C), 77 (T), 79 (G), 81 (C), 82 (A), 89 (C), 133 (C), 144 (G), scarlet, reverse sienna; on PDA surface umber, outer region
145 (C), 146 (C), 147 (T), 150 (G), 302 (C), 348 (T), 380 (C), 392 saffron, reverse sienna with patches of saffron; on OA surface
(A), 437 (T), 445 (T), 446 (C), 473 (A), 477 (G), 601 (G), 636 (T), sienna with patches of saffron.
112
Typus: France, La Reunion Island, leaf spots on Saccharum Typus: South Africa, Eastern Cape Province, Haga Haga, on
officinarum (Poaceae), May 2015, P.W. Crous, HPC 296 (ho- leaves of Cyperus sphaerocephala (Cyperaceae), Dec. 2014,
lotype CBS H-23876, CBS 145373 = CPC 26421). M.J. Wingfield (holotype CBS H-22622, culture ex-type CPC
25702 = CBS 141450).
Notes: This species is closely related to Nph. sacchari, which is a
species also isolated from sugarcane. However, the ITS se- Authors: Y. Marin-Felix & P.W. Crous
quences of the type material of both species showed only a
Stagonosporopsis Died., Ann. Mycol. 10: 142. 1912. Emend.
97.68 % of nucleotide similarity. Unfortunately, tef1 and tub2
Aveskamp et al., Stud. Mycol. 65: 44. 2010. Fig. 75.
sequences of Nph. sacchari are not available in order to compare
both species. These species produce both the sexual and Classification: Dothideomycetes, Pleosporomycetidae, Pleo-
asexual morphs, with morphological differences most obvious in sporales, Didymellaceae.
the sexual morph. Neosetophoma pseudosacchari can be easily
distinguished by its larger ascomata (up to 300 μm diam in Nph. Type species: Stagonosporopsis boltshauseri (Sacc.) Died.,
pseudosacchari vs. up to 180 μm diam in Nph. sacchari), asci designated as lectotype by Clements & Shear (1931), basionym:
[70–100 × 10–13 μm in Nph. pseudosacchari vs. Ascochyta boltshauseri Sacc. = Stagonosporopsis hortensis
60–75(–85) × 12–15(–17) μm in Nph. sacchari] and asco- (Sacc. & Malbr.) Petr., basionym: Hendersonia hortensis Sacc. &
spores [(22–)25–30 × (5.5–)6 μm in Nph. pseudosacchari vs. Malbr. Representative strain of Sta. hortensis: CBS 572.85 = PD
20–23(–25) × 5–6 μm in Nph. sacchari]. 79/269.
Wingfieldomyces Y. Marín & Crous, gen. nov. MycoBank DNA barcode (genus): LSU.
MB829671. Table 18. Fig. 74. DNA barcodes (species): ITS, rpb2, tub2. Table 19. Fig. 76.
Etymology: Named in honour of its collector, Prof. dr M.J. Ascomata pseudothecial, globose to subglobose, sometimes
Wingfield, who contributed greatly to the elucidation of African with a somewhat conical neck. Asci cylindrical or subclavate, 8-
fungal biodiversity. spored, biseriate. Ascospores ellipsoid, fusoid or obovoid, uni-
Ascomata immersed on host, erumpent in culture, black, septate, guttulate, sometimes with a gelatinous sheath. Con-
globose, with central ostiole; ascomatal wall of 3–4 layers of dark idiomata pycnidial, globose to subglobose, glabrous or with
brown cells of textura angularis. Pseudoparaphyses intermingled hyphal outgrowths, superficial on agar surface or immersed,
among asci, hyaline, septate, branched prominently, constricted solitary or confluent, ostiolate or poroid, occasionally papillate;
at septa. Asci bitunicate with apical chamber, subcylindrical, conidiomatal wall pseudoparenchymatous, 2–6 layered, with an
hyaline, smooth, fasciculate, stipitate, 8-spored. Ascospores tri- outer wall composed of 1–3 layers of brown to olivaceous cells.
to multiseriate, fusoid with subobtusely rounded ends, finely Conidiogenous cells phialidic, hyaline, simple, smooth-walled,
verruculose, red-brown, guttulate, 2-septate, slightly constricted ampulliform or doliiform. Conidia often dimorphic: mainly asep-
at septa, with central cell somewhat swollen. tate, hyaline, ellipsoid to subglobose, thin-walled, smooth-walled,
eguttulate or with several polar or scattered guttules; second type
Culture characteristics: Colonies spreading, erumpent surface, of conidia larger in size, can be produced both in vivo and in vitro
with moderate aerial mycelium, margins feathery. On MEA, PDA in the same pycnidia as the other type of conidia, 0–3-septate.
and OA surface dirty white; reverse dirty white to luteous.
Culture characteristics: Colonies on OA regular to somewhat
Type species: Wingfieldomyces cyperi (Crous & M.J. Wingf.) Y. irregular, colourless, buff, luteous to ochraceous or amber, or
Marín & Crous. Holotype and ex-type cultures: CBS H-22622, olivaceous grey to greenish grey, with scarce or abundant floc-
CBS 141450 = CPC 25702. cose white to pale salmon, or olivaceous grey aerial mycelium.
Notes: Wingfieldomyces is introduced to accommodate Set. Optimal media and cultivation conditions: On OA at 20–24 °C
cyperi since, based on phylogenetic data, it is located in an in- under near-ultraviolet light (13 h light, 11 h dark) to induce
dependent lineage distant to the clade representing the genus sporulation of the asexual morph, while MEA stimulates
Setophoma. Moreover, based on a megablast search using the pigmentation and crystal formation. Changes in colour of the
ITS sequence, the closest matches in NCBIs GenBank nucleo- fungal cultures upon a sudden increase of pH (NaOH spot test),
tide database were Pringsheimia euphorbiae [GenBank which may be used for taxonomic characterisation, are best
NR_145344; Identities = 456/500 (91 %), 8 gaps (1 %)] and observed on OA.
Phaeosphaeria caricis [GenBank KY090633; Identities 439/485
(91 %), 12 gaps (2 %)]. It only produces a sexual morph in Distribution: Worldwide.
culture, characterised by tri- to multiseriate, 2-septate, red-brown Hosts: Associated with at least 30 plant genera in Asteraceae,
ascospores, while Setophoma produces both morphs and 2- Brassicaceae, Campanulaceae, Caricaceae, Cucurbitaceae,
seriate, 3-septate, hyaline ascospores with the second cell Fabaceae, Lamiaceae, Ranunculaceae, Solanaceae and Valer-
from the apex becoming swollen. Wingfieldomyces is associated ianaceae as saprobes or pathogens. Stagonosporopsis oculo-
with leaf scorch symptoms on Cyperus. hominis is the only species that is not associated with a plant
Wingfieldomyces cyperi (Crous & M.J. Wingf.) Y. Marín & host and was isolated from human corneal ulcer in the USA.
Crous, comb. nov. MycoBank MB829672. Fig. 74. Disease symptoms: Plant stunting, seedling damping-off, leaf spots
Basionym: Setophoma cyperi Crous & M.J. Wingf., Persoonia and dieback, crown rot, stem canker, flower blight, and fruit rot.
36: 385. 2016.
Notes: Many species of Stagonosporopsis are phytopathogens,
Description: Crous et al. (2016b). causing devastating diseases on plants from various families.
MARIN-FELIX ET AL.
Fig. 84. Zasmidium spp. A–C. Disease symptoms caused by Zasmidium cyatheae (ex-type CPC 24725). A, B. Frond spots on Cyathea delgadii. C. Erumpent subcuticular
ascomata, fruiting epiphyllous. D–F. Sexual morph of Zasmidium cyatheae (ex-type CPC 24725). D. Ascoma. E. Asci. F. Ascospores. G–R. Asexual morph. G–L. Co-
nidiophores. G. Zasmidium biverticillatum (CBS 335.36). H. Zasmidium citri-griseum (ex-epitype CBS 139467). I. Zasmidium fructigenum (ex-type CBS 139626). J. Zasmidium
indonesianum (ex-type CBS 139627). K. Zasmidium musigenum (ex-type CBS 365.36). L. Zasmidium strelitziae (ex-type CBS 121711). M–Q. Conidiophores. M. Zasmidium
biverticillatum (CBS 335.36). N. Zasmidium biverticillatum (CBS 335.36). O. Zasmidium citri-griseum (ex-epitype CBS 139467). P. Zasmidium fructicola (ex-type CBS 139625).
Q. Zasmidium musigenum (ex-type CBS 365.36). R. Primary and secondary conidia of Zasmidium cellare (ex-neotype CBS 146.36). Scale bars: 10 μm. Pictures A–F taken
from Guatimosim et al. (2016); G, K–N, Q, R from Arzanlou et al. (2007); H–J, O, P from Huang et al. (2015).
114
Some species have a worldwide distribution, e.g., Sta. cucur- Stagonosporopsis chrysanthemi (F. Stevens) Crous et al.,
bitacearum on Cucurbitaceae, and Sta. hortensis on Fabaceae, Australas. Pl. Pathol. 41: 681. 2012.
while others represent important quarantine plant pathogens Basionym: Ascochyta chrysanthemi F. Stevens, Bot. Gaz. 44:
limited to certain geographical areas. For example, Sta. andi- 246. 1907.
gena and Sta. chrysanthemi are classified as A1 and A2 Synonyms: Mycosphaerella ligulicola, K.F. Baker et al., Phyto-
quarantine pathogens, respectively, by the European and pathology 39: 799. 1949.
Mediterranean Plant Protection Organisation (EPPO; 2016). Didymella ligulicola (K.F. Baker et al.) Arx, Beitr. Kryptfl. Schweiz
Stagonosporopsis tanaceti is a destructive pathogen of pyre- 11: 364. 1962.
thrum (Tanacetum cinerariifolium) in Australia but has not been Didymella ligulicola var. ligulicola (K.F. Baker et al.) Arx, Stud.
reported elsewhere in the world (Vaghefi et al. 2012). Some Mycol. 32: 199. 1990.
Stagonosporopsis species have been isolated from plants but Phoma ligulicola var. ligulicola Boerema, Stud. Mycol. 32: 9.
their pathogenicity has not been established. For example, Sta. 1990.
dennisii has been reported from dead stems of Solidago spp. Stagonosporopsis ligulicola var. ligulicola (K.F. Baker et al.)
but no data are available on its pathogenicity (Boerema et al. Aveskamp et al., Stud. Mycol. 65: 46. 2010.
2004).
Typus: USA, North Carolina, West Raleigh, on Chrysanthemum
Species identification based on only morphology is unreliable.
indicum, Dec. 1906, F.L. Stevens (Bartholomew, Fungi Colum-
Stagonosporopsis was originally separated from Ascochyta
biani no. 2502, Field Museum of Natural History, C0004169F;
based on occasional formation of multi-septate (stagonospora-
designated here as lectotype, MBT385563); on Chrysan-
like) conidia (Diedicke 1912). However, later phylogenetic
themum morifolium, 1949, L.H. Davis [epitype of Ascochyta
studies revealed that some Stagonosporopsis spp. lack the
chrysanthemi designated here ATCC 10748, MBT385567 (pre-
stagonospora-like spores or any features except for globose
served in a metabolically inactive state)].
pycnidial conidiomata, and aseptate, hyaline conidia (Aveskamp
et al. 2010). Thus, multi-locus sequence typing is essential for Notes: Stevens (1907) described Ascochyta chrysanthemi on
identification of Stagonosporopsis species. The emended Chrysanthemum indicum from North Carolina; however, he did
description of the genus Stagonosporopsis by Aveskamp et al. not refer to a holotype specimen in the original description. A
(2010) states that the sexual morph of Stagonosporopsis, if specimen at Field Museum of Natural History (C0004169F) is
present, occurs only in vivo. However, some strains of Sta. chosen as lectotype, among numerous other duplicates depos-
chrysanthemi, Sta. caricae and Sta. inoxydabilis have been ited at BPI, CUP, NYBG, MSC, and various other herbaria that
shown to produce pseudothecial ascomata intermingled with include collections distributed as E. Bartholomew, Fungi
pycnidial conidiomata on agar media (Boerema et al. 2004, Columbiani 2502. Since no living cultures derived from these
Vaghefi et al. 2012). specimens are available, we designate ATCC 10748, isolated
Currently, more than 40 species are linked to the genus Sta- from Chrysanthemum morifolium from North Carolina, as ex-
gonosporopsis. However, only 27 species are recognised based on epitype culture of Ascochyta chrysanthemi here.
molecular data (Table 19). Previous phylogenetic studies have
Authors: N. Vaghefi, Y. Marin-Felix, P.W. Crous & P.W.J. Taylor
used one locus (act in De Gruyter et al. 2012), three loci (LSU, ITS
and tub2 in Aveskamp et al. 2010), four loci (LSU, ITS, tub2 and act Stemphylium Wallr., Flora Cryptogamica Germaniae 2: 300.
in Hyde et al. 2014; LSU, ITS, tub2 and rpb2 in Chen et al. 2015; 1833. Fig. 77.
ITS, tub2, chs and cal in Stewart et al. 2015) and five loci (LSU, ITS, Synonyms: Scutisporium Preuss, Linnaea 24: 112. 1821.
tub2, act and tef1 in Vaghefi et al. 2012) for phylogenetic species Epochniella Sacc., Michelia 2: 127. 1880.
recognition in Stagonosporopsis. However, in most cases, ITS and Soreymatosporium Sousa da Câmara, Proposta Stemphylium:
tub2 sequences are sufficient for achieving resolution to species 18. 1930.
level. While ITS sequences alone may be used to distinguish Thyrodochium Werderm., Annls mycol. 22: 188. 1942.
Stagonosporopsis as a monophyletic clade within Didymellaceae,
tub2 fails to distinguish Stagonosporopsis and, thus, needs to be
sporales, Pleosporaceae.
always combined with ITS (Chen et al. 2015). A phylogeny pro-
duced by rpb2 alone is highly similar to the combined four-locus Type species: Stemphylium botryosum Wallr. = Pleospora tarda
phylogeny based on LSU, ITS, tub2 and rpb2. However, amplifi- E.G. Simmons. Holotype of Ste. botryosum: "Ad sparagam" in
cation of rpb2 has not been successful for many Stagonosporopsis herb. Wallroth, STR. Ex-type strain of Ple. tarda: CBS 714.68.
spp. (Chen et al. 2015). Likewise, while partial cal sequences
DNA barcode (genus): ITS.
provide high resolution for Stagonosporopsis species delineation, it
has not been successfully amplified in some strains (Aveskamp DNA barcodes (species): cmdA, gapdh. Table 20. Fig. 78.
et al. 2010, Vaghefi et al. 2012). Thus, the use of ITS and tub2 is
Ascomata pseudothecial, globose or ovoid, membranous, dark
recommended as they will provide sufficient resolution for almost all
brown to black, sometimes with a slender neck. Asci oblong to
Stagonosporopsis species, are easier to amplify, and are available
clavate, with distinct outer and inner walls. Ascospores elongate
for the majority of Stagonosporopsis spp. described to date
to oval, with 7 horizontal and 3–5 longitudinal septa, yellowish to
(Table 19). The only two species that cannot be separated based
brown, muriform on maturity. Conidiophores dark due to per-
on the LSU-ITS-tub2 phylogeny are S. bomiensis and S. papillata,
current proliferation forming phaeodictyospores, mostly solitary,
for which sequencing of rpb2 was necessary (Chen et al. 2017).
straight or flexuous, short or long, branched or unbranched,
References: Boerema et al. 2004 (morphology and distribution); aseptate or septate. Conidiophores proliferate further after a
Aveskamp et al. 2010, Chen et al. 2015 (morphology and conidium is produced, producing new cells and new conidia.
phylogeny). Conidiogenous cells swollen at apex, single or in group. Conidia
MARIN-FELIX ET AL.
olive, dark or pale brown, verrucose, oblong or muriform, with 3 resolution; however, more significant support is obtained when all
or more constricted transverse and 1–2 longitudinal or oblique three loci are combined.
septa. Environmental factors such as temperature and moisture are
key in Stemphylium disease development. Plant debris and
Culture characteristics: Stemphylium colonies grow rapidly on a
seeds are primary sources of inoculum of Stemphylium in most
variety of media. On most media, the colonies are velvety to
host plant species. When environmental conditions are favour-
cottony in texture with a pale or dark olivaceous grey, brown or
able, the pathogen can cause significant loss to various agri-
brownish black colour and black pigmentation on the colony
cultural crops such as lupin and cotton (Boshuizen et al. 2004).
reverse. Conidial density is low in cultures produced under
laboratory conditions even when the isolate is grown under References: Simmons (1967), Ellis (1971), Bayaa & Erskine
alternate cycle of 12 h light and 12 h darkness on PDA. Aerial (1998), Câmara et al. (2002) (morphology); Bashi & Rotem
mycelia flat/effuse, woolly or compact. Margins smooth and (1975), Mwakutuya (2006) (culture characteristics); Boshuizen
sharp or crenate and lobate. et al. (2004) (biology and life cycle); Wang et al. (2010) (host
range); Woudenberg et al. (2017), Crous et al. (2019b) (optimal
Optimal media and cultivation conditions: SNA for pigmentation
media and growing conditions).
and morphological identification, PCA for morphological identi-
fication and alternatively PDA for pigmentation and morpholog- Stemphylium rombundicum Moslemi, Y.P. Tan & P.W.J. Taylor,
ical identification. Incubation for 1–2 wk at moderate sp. nov. MycoBank MB829291. Fig. 79.
temperatures from 23–27 °C (depending on the species) under
Etymology: Named after the famous beverage, Bundaberg Rum
cool white florescent light with an 8-hr or 12-hr photoperiod.
(Bundy Rum), produced in Bundaberg, Queensland, Australia,
Distribution: Worldwide. where the fungus was first isolated.
Hosts: Many Stemphylium species are saprophytes and grow on Conidiophores long, solitary, straight, septate, verrucose, light or
plant debris and cellulose material. However, plant pathogenic dark brown, (62–)111–258(–307) μm, bearing one thickened,
species, such as Ste. beticola, Ste. botryosum, Ste. loti, Ste. darkened, percurrent rejuvenation site. Conidiogenous cells
solani and Ste. vesicarium, can cause devastating damage and swollen at apex, darkened, (5–)6–9(–10) μm wide. Conidia
significant loss of agriculturally important crops annually. Stem- solitary, conidium body light brown to golden, turning into dark
phylium spp. are pathogenic to a wide range of hosts, such as brown around longitudinal and transverse septa, verrucose,
tomato, garlic, asparagus, alfalfa, lupin, lentil and cotton. The oblong or cylindrical, occasionally ovoid with curved apex,
ability of pathogens to infect a wide range of crops reflects its (27.5–)35–55.5(–61) × (10–)12.5–24(–26.5) μm, with 3–4
adaptability to wide range of climatic conditions and provides transverse septa, 1–2(–3) longitudinal or oblique septa per
better survival chances. transverse sector. Sexual morph not observed.
Disease symptoms: Leaf spot, defoliation, curling and bending of Culture characteristics: Colonies on SNA after 1 wk reaching
the leaf margins and stems. Lesion size differs in various hosts 35 mm diam, effuse, hairy or velvety, white, colourless, mycelia
and can grow to encompass the entire leaf and reduce mostly immersed in the agar. On PDA reaching 35 mm diam, fast
photosynthesis. growing, with compact, entire, aerial mycelium, fine, woolly on
the surface; reverse dark orange to dark brown in the centre with
Notes: Simmons (1967) established criteria for morphological
central pale brown rings growing towards the sides; thick yellow
identification of various Stemphylium spp. and introduced Pleo-
margins, and grey zones can also be seen.
spora herbarum as the sexual morph of Ste. botryosum. How-
ever, Simmons (1985) subsequently reclassified Stemphylium/ Typus: Australia, Queensland, Bundaberg (Burnett Heads), from
Pleospora holomorphs and reported P. tarda as the sexual morph fruit lesions of Solanum lycopersicum (Solanaceae), 9 Aug.
of Ste. botryosum and P. herbarum the sexual morph of Ste. 2000, J. Maltby (holotype BRIP 27486, culture ex-type BRIP
herbarum (Moslemi et al. 2017). The asexual morph Stemphy- 27486).
lium has been well studied, though the sexual morph Pleospora
Notes: Colonies of Ste. lycopersici produce a yellow dark red
is poorly defined. The number of Pleospora spp. identified may
pigmentation diffusing out in PDA and other media (Yamamoto
be as many as 1000 and they are reported to be polyphyletic.
1960). As Ste. rombundicum is closely related to species in
Stemphylium is morphologically similar to the closely related
the Ste. lycopersici complex, similar physiological characters can
genus Alternaria. However, unlike Alternaria in which its conidia
be observed on PDA. Colonies only sporulated at 23 °C under
remain attached and form a chain, Stemphylium conidia are
12 h photoperiod on PDA. Against Ste. lycopersici in which the
always solitary, arising from a conidiogenous cell with a swollen
conidia are mostly ovoid with a pointed apex, Ste. rombondicum
apex (Inderbitzin et al. 2009). Morphological features such as
mostly contains cylindrical or oblong conidia. It is difficult to
spore shape and size, conidiophores, ascospores and the size
observe the longitudinal septa in conidia of Ste. rombundicum.
and time of pseudothecial maturation are important character-
Conidiophores are significantly long compared to the type Ste.
istics in species identification. Other features such as variation in
lycopersici in which the conidiophores length do not exceed
conidial wall ornamentation and septum development are not
140 μm. Conidia of Ste. lycopersici are significantly longer
considered as important parameters in Stemphylium identifica-
(50–74 μm × 16–23 μm) than those of Ste. rombundicum.
tion (Câmara et al. 2002).
For phylogenetic analyses of Stemphylium species, cmdA and Stemphylium truncatulae Moslemi, Y.P. Tan & P.W.J. Taylor,
gapdh were identified as the most informative genes, and rpb2 sp. nov. MycoBank MB829282. Fig. 80.
and actA as the least informative (Woudenberg et al. 2017).
Etymology: Named after the host species, Medicago truncatula,
Among ITS, cmdA and gapdh, cmdA provides the highest
from which it was first collected.
116
Immature ascomata observed on SNA and PDA embedded in 235.5–337(–381) μm, with outgrowing mycelia; ascomatal wall
the agar. Ascomata pseudothecial, dark brown to black, globose thin, 10–15 μm thick. Conidiophores solitary, straight, simple
or flask shaped, solitary or aggregated in groups of 3–5, (83.5–) or occasionally 1-branched, septate, smooth-walled, pale
185.5–186(–304) × (351–)453–483(–603) μm, with outgrowing brown, (29–)42–85(–98) μm long, cylindrical, enlarging
dark mycelia; ascomatal wall 27–32 μm thick. Conidiophores apically to the site of conidium production. Conidiogenous cells
solitary, straight to flexuous, mostly branched, septate or occa- swollen at apex, darkened, (6–)6.5–7(–7.5) μm wide. Conidia
sionally aseptate, smooth-walled, light brown, mostly 7–11 μm solitary, dark reddish brown, verrucose, ovoid, oblong or cy-
length, some 17–39 μm in length, bearing 2–6 thickened, pale, lindrical, (18–)25–49(–52) × (10.5–)14–26(–30) μm, with
percurrent rejuvenation sites. Conidiogenous cells slender or 2–5 transverse septa, 1–2(–3) longitudinal or oblique septa
slightly swollen at apex, pale, (5.5–)6.5–7(–9.5) μm. Conidia per transverse sector, constricted at multiple darkened trans-
solitary, conidium body pale brown to golden or olivaceous verse septa.
brown, mostly smooth-walled, sometimes minutely verrucose,
Culture characteristics: Colonies on SNA reaching 28 mm diam
usually ovoid, occasionally with pointed apex, (8–)
after 1 wk with flat, entire and fluffy aerial mycelia in the centre,
9.5–18(–21) × (14–)15.5–29.5(–32) μm, with 2–4 transverse
sub-hyaline. On PDA reaching 20 mm diam after 1 wk, compact,
septa, one longitudinal or oblique septa per transverse sector.
entire, aerial mycelium white, woolly, with rings of light olivaceous
Culture characteristics: Colonies on SNA reaching grey in the centre and dark olivaceous grey to black on the
21–25 mm diam after 1 wk, with white and fluffy aerial mycelia in reverse side, margins regular, thick and white.
the centre; reverse colourless with pale olivaceous grey centre.
Typus: Australia, South Australia, Waikerie, from leaf spots on
Colonies on PDA reaching 25 mm diam after 1 wk, with white
Allium sativum (Alliaceae), 14 Nov. 1997, H. Suheri (holotype
fluffy mycelia in the centre; reverse dark green to grey with thin
VPRI 21969, culture ex-type VPRI 21969).
white margins.
Notes: Stemphylium waikerieanum is morphologically similar to
Typus: Australia, Victoria, from seeds of Medicago truncatula
species in the Ste. vesicarium species complex. However, the
(Fabaceae), 10 Sep. 1982, M. Mebalds (holotype BRIP 14850,
conidium length in Ste. vesicarium complex does not exceed
culture ex-type BRIP 14850).
45 μm (Simmons 1969), whereas conidia of Ste. waikerieanum
Notes: Differs from the type species Ste. botryosum described by were observed up to 49 μm long. The multigene phylogenetic
Simmons (1969) by producing significantly smaller conidia. Ac- analysis of ITS, gapdh and cmdA also support (PP value =1) this
cording to Simmons (1969), conidia of Ste. botryosum are species as a novel taxon.
24–26 μm wide and 33–35 μm long. Additionally, colonies of
Authors: A. Moslemi, J. Edwards, Y.P. Tan & P.W.J. Taylor
Ste. botryosum grow rapidly to 48 mm diam after 6 d of incu-
bation at 25 °C (Hosen et al. 2013), while Ste. truncatulae is slow Tubakia B. Sutton, Trans. Brit. Mycol. Soc. 60: 164. 1973.
growing. The morphological identifications along with sequence Fig. 82.
analyses support Ste. truncatulae as a unique taxon closely Synonyms: Actinopelte Sacc., Annls mycol. 11: 315. 1913.
related to the type species Ste. botryosum (CBS 714.68).
Classification: Dothideomycetes, Diaporthomycetidae, Dia-
Stemphylium waikerieanum Moslemi, Jacq. Edwards & P.W.J porthales, Tubakiaceae.
Taylor, sp. nov. MycoBank MB829283. Fig. 81.
Type species: Tubakia japonica (Sacc.) B. Sutton, basionym:
Etymology: Named after the location, Waikerie in South Actinopelte japonica Sacc. Epitype and ex-epitype strains
Australia, from where it was collected. designated by Braun et al. (2018): NBRC H-11611, NBRC
9268 = MUCC2296 = ATCC 22472.
Immature ascomata observed on SNA after 2 wk embedded in
the agar. Ascomata pseudothecial, dark brown to black, mostly DNA barcodes (genus): ITS, LSU, rpb2.
flask-shaped or occasionally globose, solitary or aggregated in
DNA barcodes (species): ITS, tef1, tub2. Table 21.
groups of 4–6, (267.5–)292–349.5(–374) × (191.5–)
Fig. 85. Zasmidium thailandicum (ex-type CBS 145027). A–C. Conidiophores sporulating on SNA. D–F. Conidiogenous cells with apical rachis giving rise to conidia. Scale
bars = 10 μm.
MARIN-FELIX ET AL.
Ascomata perithecial, pigmented, dark, on fallen overwintered revision of the genus Tubakia, were accommodated in the new
leaves, rostrate, beak short, usually lateral-eccentric, slightly family Tubakiaceae (Braun et al. 2018). Presently, 16 species are
protuberant, with periphysate ostiolate; ascomatal wall variable accepted in the genus based on molecular data (ITS, tef1 and
in thickness, paler than stromatic layers, polyascal. Paraphy- tub2 sequences). These species may form different types of
ses lacking. Asci unitunicate, 8-spored, oblong-ellipsoid, stalk asexual morphs, being the punctiform conidiomata (pycnothyria)
short to oblong, ascal apex with two refractive conoid struc- composed of convex scutella with radiating threads of cells fixed
tures, asci deliquescing at maturity. Ascospores more or less to the substratum by a central columella the most common and
uniseriate, becoming irregularly biseriate, one-celled, hyaline, characteristic structure formed. Other asexual morphs include
ellipsoid to fusiform, often inequilateral or slightly curved, wall sporodochial conidiomata and crustose or pustulate pycnidioid
finely ornamented, content granular-guttulate. Conidiomata conidiomata. The conidia are globose to broad ellipsoid-obovoid,
pycnothyrial, usually circular or subcircular when viewed from sometimes subcylindrical or somewhat irregular, aseptate, hya-
above, superficial, easily removable, scutellate, fixed to the line, subhyaline to pigmented. Tubakia suttoniana is the only
leaf by a central columella; scutellum composed of loose to species that produces sexual morph characterised by ostiolate
dense hyphal strands, mostly branched, thick-walled, pig- ascomata, unitunicate asci with two refractive conoid structures
mented, margin compact or outer portions of the radiating in the ascal apex, and one-celled, hyaline ascospores.
hyphal strands looser to free, tips rounded, truncate or Species of Tubakia are endophytes and/or pathogens in
pointed, margin usually not recurved. Conidiophores reduced leaves and twigs of many tree species, causing distinct leaf le-
to conidiogenous cells. Conidiogenous cells phialidic, usually sions in different hosts including oak (Quercus spp.), chestnut
subcylindrical-conical, lageniform, hyaline to pale brown, (Castanea spp.) and other hardwood species. Moreover,
arising from small, colourless fertile cells around the upper T. iowensis is also capable of causing petiole necrosis and death
part of the central pycnothyrial columella, percurrently prolif- of whole leaves on bur oak (Q. macrocarpa), sometimes killing
erating, sometimes forming indistinct periclinal thickenings or nearly every leaf on a susceptible tree. This disease is known as
annellations (collarettes). Conidia formed singly, globose to bur oak blight and is most common in Iowa and Minnesota, but it
broad ellipsoid-obovoid, sometimes subcylindrical or some- has been noted in western and southern Wisconsin, northern
what irregular, wall thin to somewhat thickened, smooth to Illinois, northeast Kansas, eastern Nebraska, and eastern South
faintly rough, hyaline to pigmented, apex rounded, base Dakota, with isolated groups of affected trees in counties of Illi-
rounded to attenuated, sometimes with distinct frill or peg-like nois and Missouri that border Iowa (Harrington & McNew 2016).
basal hilum (adapted from Braun et al. 2018).
References: Harrington et al. 2012 (morphology and pathoge-
Culture characteristics: Colonies on MEA flat, with dense or fluffy, nicity), Harrington & McNew 2016 (pathogenicity, bur oak blight),
sparse to moderate aerial mycelium, margins regular or scal- Harrington & McNew 2018 (morphology and phylogeny), Braun
loped; surface ivory white, dingy white to pale yellow, straw, et al. 2018 (morphology, pathogenicity and phylogeny).
cream to light grey, smoke grey, white with center green olive and
Authors: Y. Marin-Felix & P.W. Crous
brown hyphal stripes, creamy white with or without concentric
rings of olivaceous mycelium, or white to grey with wet conidial Zasmidium Fr., Summa veg. Scand., Sectio Post. (Stockholm):
masses olive green to black; reverse pale grey, greyish white 407. 1849. Fig. 83.
with olivaceous margins, smoke grey with olivaceous grey Synonyms: Periconiella Sacc., Atti Ist. Veneto Sci. Lett. Arti 3:
margins, golden yellow to slightly darker, yellowish grey with 727. 1885.
concentric rings, yellow with dark grey concentric rings, straw Biharia Thirum. & Mishra, Sydowia 7: 79. 1953.
with dark brown concentric rings, or middle dark grey and yellow Stenellopsis B. Huguenin, Bull. Trimestriel Soc. Mycol. France
to medium brown towards the rim. 81: 695. 1966.
Verrucispora D.E. Shaw & Alcorn, Proc. Linn. Soc. New South
Optimal media and cultivation conditions: MEA, OA, PDA and Wales 92: 171. 1967. (nom. illegit., Art. 53.1).
PNA at 25 °C under near-ultraviolet light. Verrucisporota D.E. Shaw & Alcorn, Austral. Syst. Bot. 6: 273.
Distribution: North America, Asia, Australia and Europe. 1993.
Hosts: Castanea spp., Chrysolepis chrysophylla, Lithocarpus Classification: Dothideomycetes, Dothideomycetidae, Capno-
densiflorus and Quercus spp (Fagaceae), Liquidambar styraci- diales, Mycosphaerellaceae.
flua (Hamamelidaceae), Lindera glauca (Lauraceae), and Pinus Type species: Zasmidium cellare (Pers.) Fr., basionym: Raco-
tabuliformis (Pinaceae). Reported from other hosts not verified dium cellare Pers. Neotype designated by Videira et al. (2017):
such as Acer spp. (Aceraceae), Carya spp. (Juglandaceae) and CBS 146.36 (duplicate cultures ATCC 36951 = IFO 4862 = IMI
Fraxinus spp. (Oleaceae). 044943 = LCP 52.402 = LSHB BB274 = MUCL 10089).
Disease symptoms: Leaf spots, necrosis and death. Tubakia DNA barcode (genus): LSU.
iowensis also causes petiole necrosis and death of whole leaves
on bur oak (bur oak blight). DNA barcodes (species): ITS, rpb2, act, tef1, tub2. Table 22.
Fig. 84.
Notes: Tubakia was recently revised by Braun et al. (2018),
resulting in the introduction of five new genera to accommodate Ascomata pseudothecial, amphigenous or epiphyllous, dark
species previously placed in Tubakia, i.e. Apiognomonioides, brown or black, globose, single to aggregated; necks rarely
Involutiscutellula, Oblongisporothyrium, Paratubakia and Sapro- perceptible, usually a paler coloured circular area, composed of
thyrium. All these genera together with Racheliella and convergent yellow hyphae; ascomatal wall composed of 2–3
Sphaerosporithyrium, both genera also described during the layers of cells of textura angularis. Asci bitunicate, fasciculate,
subsessile, obpyriform, obovoid to ellipsoidal, obclavate to
118
fusoid-ellipsoidal, saccate or clavate to cylindrical, apar- Cyathea delgadii (Cyatheaceae), Dasypogon sp. (Dasypogona-
aphysate, 8-spored. Ascospores 2–3-seriate to multiseriate, ceae), Daviesia latifolia (Fabaceae), Elaeocarpus kirtonii
hyaline, smooth-walled, guttulate, without sheath, fusoid to (Elaeocarpaceae), Eucalyptus spp. (Myrtaceae), Gahnia sie-
ellipsoidal with obtuse ends, straight to slightly curved, uni- beriana (Cyperaceae), Geniostoma rupestre (Loganiaceae), Itea
septate, constricted or not at the septum, widest in the middle of parvifolia (Escalloniaceae), Lonicera japonica (Caprifoliaceae),
apical cell. In plant pathogenic species, mycelium mostly Maclura cochinchinensis (Moraceae), Malus spp. (Rosaceae),
immersed as well as superficial, rarely only immersed; hyphae Pittosporum tenuifolium (Pittosporaceae), Podocarpus sp.
branched, septate, hyaline or almost so to pigmented, pale (Podocarpaceae), Pseudotsuga menziesii and Tsuga hetero-
olivaceous to brown, wall thin to somewhat thickened, phylla (Pinaceae), Restio subverticillatus (Restionaceae), Roth-
immersed hyphae smooth-walled or almost so to faintly rough, mannia engleriana (Rubiaceae), Sasa sp. (Poaceae), Scaevola
external hyphae distinctly verruculose to verrucose (in culture taccada (Goodeniaceae), Schinus terebinthifolius (Anacardia-
immersed hyphae usually smooth-walled or almost so, aerial ceae), Strelitzia sp. (Strelitziaceae), and Syzygium cordatum
hyphae verruculose). Stromata lacking to well-developed, pig- (Myrtaceae).
mented. Conidiophores solitary, arising from superficial hyphae,
Disease symptoms: Causing various lesions, ranging from
lateral, occasionally terminal, in vivo (in plant pathogenic taxa)
yellowish discolorations to distinct leaf spots. Also associated
sometimes also fasciculate, arising from internal hyphae or
with sooty blotch and flyspeck diseases. Also isolated from wall
stromata, semimacronematous to macronematous, in culture
in wine cellar.
occasionally micronematous, cylindrical, filiform, subuliform,
straight to strongly geniculate-sinuous, mostly unbranched, Notes: The genus Zasmidium is morphologically similar to
aseptate, i.e. reduced to conidiogenous cells, to pluriseptate, Stenella, producing thickened and darkened conidiogenous loci
subhyaline to pigmented, pale olivaceous to medium dark and hila (Braun et al. 2013). However, these genera differ in the
brown, wall thin to somewhat thickened, smooth to verruculose. conidial hila and scars, being flat in Stenella and planate and
Conidiogenous cells integrated, terminal, occasionally interca- somewhat thickened, darkened in Zasmidium. Moreover, Sten-
lary, rarely pleurogenous, or conidiophores reduced to con- ella belongs to Teratosphaeriaceae while Zasmidium is located
idiogenous cells, mostly polyblastic, sympodial, with within Mycosphaerellaceae (Arzanlou et al. 2007, Quaedvlieg
conspicuous, somewhat thickened and darkened-refractive, et al. 2013, Videira et al. 2017).
planate loci. Conidia solitary or catenate, in simple or A recent phylogenetic analysis based on LSU, ITS and rpb2
branched acropetal chains, shape and size variable, ranging showed that species belonging to other genera, i.e. Mycos-
from amero- to scolecosporous, aseptate to transversely pluri- phaerella, Parastenella, Periconiella, Ramichloridium, Rasutoria,
euseptate, subhyaline to pigmented, pale olivaceous to brown, Stenella and Verrucisporota, were located in the monophyletic
wall thin to somewhat thickened, smooth or almost so to usually clade representing the genus Zasmidium (Videira et al. 2017).
distinctly verruculose (in plant pathogenic species without su- Therefore, 12 new combinations were introduced, and the genera
perficial mycelium always verruculose); hila somewhat thick- Periconiella and Verrucisporota reduced to synonymy with Zas-
ened and darkened-refractive, planate, conidial secession midium. Based on our phylogenetic analysis, 48 species are
schizolytic (asexual morph description adapted from Braun accepted in the genus together with one new species described
et al. 2013). here, and a new combination based on Ramichloridium ducassei.
The type species of the genus, Z. cellare, has been isolated
Culture characteristics: Colonies slow growing, with sparse to
from wine cellars in Europe and America, while the other species
moderate aerial mycelium, rarely with aerial mycelium absent,
of the genus are associated to plants as saprobic or mostly
sometimes with mucoid exudate, margins smooth and regular,
biotrophic, usually foliicolous, symptomless or causing various
lobate or feathery. On MEA olivaceous, olivaceous green with
lesions, ranging from yellowish discolorations to distinct leaf
margins whitish, brown olivaceous, olivaceous grey, iron mouse
spots. Zasmidium spp. are pathogens of a wide range of hosts
grey, grey, cream, yellowish brown, vinaceous buff to olivaceous
such as Z. biverticillatum and Z. musigenum, which cause
buff, dark brown, or dark brown with margins grey; reverse pale
tropical speckle disease on members of Musaceae (Stahel 1937,
or dark grey olivaceous, olivaceous black, mouse grey, iron-grey,
Jones 2000), and Z. fructicola and Z. fructigenum, both patho-
greenish black, pale orange, isabelline, buff, dark brown, or
gens of Citrus causing a disease known as citrus greasy spot
brown vinaceous. On PDA pale white with margins pale oliva-
(Huang et al. 2015).
ceous grey, pale mouse grey, olivaceous grey, iron-grey, iron-
grey with patches orange, or brown olivaceous; reverse oliva- References: Arzanlou et al. 2007, Videira et al. 2017
ceous grey, iron-grey, iron-grey with patches orange, or isabel- (morphology and phylogeny), Braun et al. 2013 (morphology).
line. On OA mouse grey, olivaceous grey, smoke grey with
Zasmidium ducassei (R.G. Shivas et al.) Y. Marín & Crous,
margins olivaceous grey, iron-grey, or iron-grey with broad
margins of orange; reverse olivaceous, olivaceous grey, dark
Basionym: Ramichloridium ducassei R.G. Shivas et al., Aus-
mouse grey, or iron-grey.
tralas. Pl. Path. 40: 63. 2010.
Optimal media and cultivation conditions: MEA, OA, PDA and
Description and illustration: Shivas et al. (2011).
SNA at 25 °C under near-ultraviolet light.
Typus: Australia, Queensland, Daintree, on leaves of Musa
acuminata × balbisiana (Musaceae), 14 Apr. 2010, M. Berridge &
Hosts: Wide range of hosts belonging to 25 different families, K.R.E. Grice (holotype BRIP 53367, culture ex-type BRIP 53367).
including Alocasia odora and Anthurium sp. (Araceae), Aporosa Additional material examined: Malaysia, on leaves of Musa sp., 2016, P.W.
villosa (Euphorbiaceae), Brabejum stellatifolium, Grevillea spp. Crous, CPC 32929.
and Hakea undulata (Proteaceae), Citrus spp. (Rutaceae),
MARIN-FELIX ET AL.
Notes: This species was initially introduced as R. ducassei to Krueger-Hadfield. We are thankful to Dr. Paul Kirk for helpful advice regarding the
accommodate some isolates associated with a severe leaf treatment of the invalid name Seiridium cupressi (Guba) Boesew. The study of
the genus Alternaria was supported by the Spanish Ministerio de Economía y
speckle disease of Ducasse banana (Musa Competitividad, Grant CGL2017-88094-P.
acuminata × babisiana cv. Pisang awak) in northern Queensland
(Shivas et al. 2011). The authors noticed that this species was
similar to Zasmidium in having pigmented conidiophores with
integrated conidiogenous cells that sympodially proliferate near
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Genera of Phytopathogenic Fungi GOPHY3

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available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 94: 1–124 (2019).

Genera of phytopathogenic fungi: GOPHY 3

Y. Marin-Felix1,2*, M. Hernandez-Restrepo1, I. Iturrieta-Gonzalez2, D. García2, J. Gene2, J.Z. Groenewald1, L. Cai3, Q. Chen3,

*Correspondence: Y. Marin-Felix, y.marin@wi.knaw.nl; P.W. Crous, p.crous@wi.knaw.nl

Key words: DNA barcodes, Fungal systematics, New taxa.

Available online 13 June 2019; https://doi.org/10.1016/j.simyco.2019.05.001.

Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.

INTRODUCTION (VPRI). For fresh collections, we followed the procedures pre-

Table 1. DNA barcodes of accepted Allophoma spp.

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

Species Section Isolates1 GenBank accession numbers2 References

CBS 119398T Alternaria daucicaulis

IRAN 16888FT Alternaria kordkuyana

86 X1272T Alternaria parvicaespitosa

80 CBS 121331T Alternaria breviramosa

CBS 109381T Alternaria carotiincultae

100 CBS 245.67T Alternaria radicina

CBS 481.90 Alternaria scirpicola

CBS 480.90T Alternaria caricis

Table 3. DNA barcodes of accepted Brunneosphaerella spp.

DNA barcodes (genus): LSU, ITS. Hosts: Protea spp. (Proteaceae).

Table 4. DNA barcodes of accepted Elsinoe spp.

Fig. 22. (Continued).

Table 5. DNA barcodes of accepted Exserohilum spp.

Fig. 28. (Continued).

Table 6. DNA barcodes of accepted Neosetophoma spp.

Table 7. DNA barcodes of accepted Neostagonospora spp.

Table 8. DNA barcodes of accepted Nothophoma spp.

CBS 381.91ET Nothophoma anigozanthi

CBS 142457T Nothophoma variabilis

CBS 125.93 Nothophoma arachidis-hypogaeae

CBS 123395NT Nothophoma infossa

100 LC12187 Nothophoma quercina

CBS 377.67 Nothophoma gossypiicola

97 MCC 1082T Nothophoma raii

CBS 140674T Nothophoma macrospora

Phoma herbarum CBS 615.75

Table 9. DNA barcodes of accepted Parastagonospora spp.

Table 10. DNA barcodes of accepted Phaeosphaeriopsis spp.

Table 11. DNA barcodes of accepted Pleiocarpon spp.

Table 12. DNA barcodes of accepted Pyrenophora spp.

Table 12. (Continued).

Table 13. DNA barcodes of accepted Ramichloridium spp.

Table 14. DNA barcodes of accepted Seifertia spp.

Table 15. DNA barcodes of accepted Seiridium spp.

length width CBS 908.85 Cu uprreessus lusittanica South Africa

Table 16. DNA barcodes of accepted Septoriella spp.

Table 17. DNA barcodes of accepted Setophoma spp.

Table 18. DNA barcodes of the accepted Wingﬁeldomyces sp.

Table 19. DNA barcodes of accepted Stagonosporopsis spp.

Table 20. DNA barcodes of accepted Stemphylium spp.

Fig. 78. (Continued).

Table 21. DNA barcodes of accepted Tubakia spp.

Table 22. DNA barcodes of accepted Zasmidium spp.

Table 22. (Continued).

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