ORE Open Research Exeter
TITLE
Generic names in Magnaporthales
AUTHORS
Zhang, N; Luo, J; Rossman, AY; et al.
JOURNAL
IMA Fungus
DEPOSITED IN ORE
13 February 2017
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http://hdl.handle.net/10871/25758
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IMA FUNGUS · 7(1): 155–159 (2016)
doi:10.5598/imafungus.2016.07.01.09
Ning Zhang1, Jing Luo1, Amy Y. Rossman2, Takayuki Aoki3, Izumi Chuma4, Pedro W. Crous5, Ralph Dean6, Ronald P. de Vries5,7,
Nicole Donofrio8, Kevin D. Hyde9, Marc-Henri Lebrun10, Nicholas J. Talbot11, Didier Tharreau12, Yukio Tosa4, Barbara Valent13,
Zonghua Wang14, and Jin-Rong Xu15
ART I CLE
Generic names in Magnaporthales
1
Department of Plant Biology and Pathology, Rutgers University, New Brunswick, NJ 08901, USA; corresponding author e-mail: zhang@aesop.
rutgers.edu
2
Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 97331, USA
3
Genetic Resources Center, National Agriculture and Food Research Organization, 2-1-2 Kannondai, Tsukuba, Ibaraki 305-8602, Japan
4
Kobe University, 1-1 Rokkodai cho, Nada-ku, Kobe 657-8501, Japan
5
CBS-KNAW Fungal Biodiversity Institute, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
6
Department of Plant Pathology, 2510 Thomas Hall, Raleigh, NC 27695, North Carolina State University, USA
7
Fungal Molecular Physiology, Utrecht University, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
8
Department of Plant and Soil Sciences, University of Delaware, 531 S. College Ave, 152 Townsend Hall, Newark, DE 19711, USA
9
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand
10
UMR BIOGER, INRA, AgroParisTech, Université Paris-Saclay, 78850 Thiverval-Grignon, France
11
University of Exeter, Northcote House, Exeter EX4 4QJ, UK
UMR BGPI, CIRAD, TA A 54 K, 34398 Montpellier, France
13
Department of Plant Pathology, Kansas State University, Manhattan, KS 66506-5502, USA
12
14
15
Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
Department of Botany and Plant Pathology, Purdue University, West Lafayette, IN 47907, USA
Abstract: The order Magnaporthales comprises about 200 species and includes the economically and
scientifically important rice blast fungus and the take-all pathogen of cereals, as well as saprotrophs and
endophytes. Recent advances in phylogenetic analyses of these fungi resulted in taxonomic revisions. In this
paper we list the 28 currently accepted genera in Magnaporthales with their type species and available gene and
genome resources. The polyphyletic Magnaporthe 1972 is proposed for suppression, and Pyricularia 1880 and
Nakataea 1939 are recommended for protection as the generic names for the rice blast fungus and the rice stem
rot fungus, respectively. The rationale for the recommended names is also provided. These recommendations
are made by the Pyricularia/Magnaporthe Working Group established under the auspices of the International
Commission on the Taxonomy of Fungi (ICTF).
Key words:
Ascomycota
Magnaporthe
Nakataea
one fungus-one name
pleomorphic fungi
Pyricularia
rice blast
take-all
Article info: Submitted: 2 March 2016; Accepted: 15 May 2016; Published: 8 June 2016.
INTRODUCTION
Magnaporthales (Sordariomycetes, Ascomycota) contains
important pathogens of cereals and grasses, including the
rice blast fungus Pyricularia oryzae (Magnaporthe oryzae),
the take-all pathogen of cereals Gaeumannomyces graminis,
the rice stem rot pathogen Nakataea oryzae (Magnaporthe
salvinii) and the summer patch pathogen of turf grass
Magnaporthiopsis poae (Cannon 1994, Thongkantha et al.
2009). To date, about 200 species have been described in
Magnaporthales, of which approximately 50 % are pathogens
of domesticated and wild monocotyledons.
The rice blast fungus has conidial (asexual) and ascosporic
(sexual) morphs and the rice stem rot fungus produces
sclerotial (asexual), conidial (asexual) and ascosporic
(sexual) morphs. Historically, such pleomorphy added to the
difficulty in resolving taxonomic and nomenclatural problems
associated with these species.
Recent advancement in gene, transcriptome and genome
sequencing of Magnaporthales fungi resulted in robust
phylogenies, which correspond well with the pathogenicity,
ecology and biology of these species. However, the
phylogenies conflict with certain traditional generic concepts
based on morphology. Magnaporthe and Gaeumannomyces,
for example, were shown to be polyphyletic. Taxonomic
revisions have been carried out for some of these taxa in
recent publications (Luo & Zhang 2013, Klaubauf et al.
2014, Luo et al. 2015a). In this paper, we list 28 accepted
genera in Magnaporthales and provide the rationale for the
recommended genera if there is competition.
© 2016 International Mycological Association
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VOLUME 7 · NO. 1
155
ART I CLE
Zhang et al.
A list of accepted generic names in Magnaporthales, with
the type species, is compiled in Table 1 including references
that serve as the basis for recognizing these genera. This
follows approval of their usage by the Pyricularia/Magnaporthe
Working Group, without prejudice. Cases that require action
to be approved by the Nomenclature Committee for Fungi
(NCF) are indicated by an asterisk in that Table.
RECOMMENDATIONS
(A) = a name typified by an asexual morph, (S) = a name
typified by a sexual morph.
Use Nakataea Hara 1939 (A) rather than
Magnaporthe R.A. Krause & R.K. Webster 1972
(S)
Cattaneo (1876) first named the rice stem rot pathogen as
Sclerotium oryzae based on the sclerotial morph. In the same
paper he also described Leptosphaeria salvinii, which was
later recognized as the sexual morph of the same fungus
(Tullis 1933). Hara (1939) named the conidial morph of the
species Nakataea sigmoidea. Krause & Webster (1972) then
established the new generic name Magnaporthe, typified by the
ascosporic morph, to accommodate the rice stem rot pathogen
as Magnaporthe salvinii, as necessitated by the nomenclatural
rules then in force. Sclerotium and Leptosphaeria currently
belong in Basidiomycota and Dothideomycetes respectively
(Xu et al. 2010), and are therefore not applicable to this
sordariomycetous species. Nakataea and Magnaporthe are
congeneric and their type species, Nakataea sigmoidea and
Magnaporthe salvinii, refer to the same species (Krause &
Webster 1972). Subsequent to the ending of the separate
naming of morphs of the same fungus species in 2011, under
Art. 59.1 of the International Code of Nomenclature for algae,
fungi, and plants (ICN; McNeill et al. 2012), Luo et al. (2013)
made a new combination for the rice stem rot fungus as
Nakataea oryzae, using the oldest legitimate generic name
and species epithet. Those authors did not, however, formally
propose the rejection or suppression of the later sexually
typified names as currently required by Art. 57.2. We note that
it is proposed that this requirement be eliminated from the ICN
in 2017 (Hawksworth 2015), but it is currently in force.
Use Pyricularia oryzae Cavara 1892 (A) rather
than Magnaporthe oryzae (Catt.) B.C. Couch
2002 (S)
In 1880, Saccardo established the generic name Pyricularia
based on the asexually typified P. grisea on crabgrass. The
rice isolates were designated as P. oryzae in 1892 by Cavara,
which now is known as the rice blast fungus. Since then, over
50 species have been listed as Pyricularia that cause blast
diseases of monocotyledonous plants.
The sexual morph of Pyricularia was first observed in
1970 from laboratory crossing experiments and believed to
belong to Magnaporthe because of the similarity in ascospore
morphology (Hebert 1970, Barr 1977, Couch & Kohn 2002).
However, recent phylogenetic and phylogenomic analyses
demonstrated that the sexually typified genus Magnaporthe
was polyphyletic. The rice blast fungus is not congeneric
156
with the type species of Magnaporthe, M. salvinii, and the
placement of the rice blast fungus in Magnaporthe was
based on an incorrect morphological identification (Zhang et
al. 2011, Luo & Zhang 2013, Luo et al. 2014, Murata et al.
2014, Luo et al. 2015a). This is not a nomenclatural issue
because the generic names Magnaporthe and Pyricularia are
not congeneric and so do not compete for priority.
Pyricularia and Magnaporthe are currently both widely used
generic names, and the rice blast fungus is an economically
and scientifically important species that deserves much
caution. The Pyricularia/Magnaporthe Working Group has
considered the possibility of conserving the name Magnaporthe
over Pyricularia. However, such conservation would require a
change in the type species of the genus Magnaporthe, and
would cause numerous name changes for those species
currently placed in Pyricularia.
The asexually typified generic name Pyricularia is the
correct name for the rice blast fungus, which corresponds well
with pathogenicity and ecological and evolutionary features.
The name Pyricularia oryzae should therefore be used for
the rice blast fungus. The synonym Magnaporthe oryzae,
can nevertheless continue to be mentioned in publications
as “Pyricularia oryzae (syn. Magnaporthe oryzae)”. This
practice will help to bridge a potential gap in the literature and
knowledge for this important species.
Use Clasterosporium Schwein. 1832 (A) rather
than Clasterosphaeria Sivan. 1984 (S)
The generic name Clasterosphaeria, typified by C. cyperi,
was established for the sexual morph of Clasterosporium
cyperi and includes only two names. The generic name
Clasterosporium based on C. caricinum includes 158
names, many of which have been placed in other genera.
Whether or not Clasterosphaeria cyperi is congeneric with
Clasterosporium caricinum is not known, although this seems
likely given that both occur on Cyperaceae. If this is the
case, use of the older, more commonly used generic name
Clasterosporium is recommended.
Use Gaeumannomyces Arx & D.L. Olivier 1952
(S) rather than Harpophora Gams 2000 (A)
The generic name Gaeumannomyces, typified by G.
graminis, has long been used for the cause of take-all of
wheat disease (Walker 1972, 1980). Harpophora was
established for phialophora-like species that were known
to be asexual morphs related to Gaeumannomyces
and Magnaporthe but did not produce a sexual morph
(Gams 2000). With the change to one name, Harpophora
based on H. radicicola is to be considered a synonym of
Gaeumannomyces, based on phylogeny (Luo et al. 2015b).
Given the greater number of species, priority, and numerous
reports, we see no reason not to use the first published
name, Gaeumannomyces.
ACKNOWLEDGMENTS
This work was partially supported by the National Science Foundation
of the United States (grant number DEB 1145174 and DEB 1452971)
to Ning Zhang.
IMA FUNGUS
Generic names in Magnaporthales
Taxa
Type species
Number of
species
GenBank accession numbers for gene and
genome sequence data**
Buergenerula Syd. in Annls
mycol. 34: 392. 1936.
Buergenerula biseptata
(Rostr.) Syd. 1936.
(Metasphaeria biseptata
Rostr. 1904).
4
Buergenerula spartinae: transcriptome (SRX798618)
(Luo et al. 2015a).
Bussabanomyces Klaubauf et
al. in Stud. Mycol. 79: 100. 2014.
Bussabanomyces
longisporus (Bussaban)
Klaubauf et al. 2014.
(Pyricularia longispora
Bussaban 2003).
1
Bussabanomyces longisporus: transcriptome
(SRX798619) (Luo et al. 2015a).
Ceratosphaerella Huhndorf et al.
in Mycologia 100: 941. 2008.
Ceratosphaerella
castillensis (C.L. Sm.)
Huhndorf et al. 2008.
2
Ceratosphaerella castillensis: ITS (EU527997), LSU
(EU528003) (Huhndorf et al. 2008).
Ceratosphaeria Niessl in Verh.
nat. Ver. Brünn 14: 203. 1876.
Ceratosphaeria
lampadophora (Berk. &
Broome) Niessl 1876.
34
Ceratosphaeria lampadophora: ITS (AY761088),
LSU (AY346270) (Huhndorf et al. 2008).
* Clasterosporium Schwein. in
Trans. Am. phil. Soc., New Series
4: 300. 1832.
Clasterosporium caricinum
Schwein. 1832.
158
Clavatisporella K.D. Hyde in
Mycotaxon 55: 276. 1995.
Clavatisporella musicola
K.D. Hyde 1995.
1
Falciphora J. Luo & N. Zhang in
Mycologia 107: 643. 2015.
Falciphora oryzae (Z.L.
Yuan et al.) J. Luo & N.
Zhang 2015.
1
Falciphora oryzae genome (JNVV01000000) (Xu et
al. 2014).
Gaeumannomyces Arx & D.L.
Olivier in Trans. Brit. mycol. Soc.
35: 32. 1952.
Gaeumannomyces
graminis var. graminis
(Sacc.) Arx & D.L. Olivier
1952.
7
Gaeumannomyces graminis var. avenae:
transcriptome (SRX798620) (Luo et al. 2015a);
Gaeumannomyces graminis var. graminis:
transcriptome (SRX798621) (Luo et al. 2015a);
Gaeumannomyces graminis var. tritici: genome
(ADBI00000000) (Okagaki et al. 2015).
Herbampulla Scheuer &
Nograsek in Mycotaxon 47: 419.
1993.
Herbampulla crassirostris
Scheuer & Nograsek 1993
1
Kohlmeyeriopsis Klaubauf et al.
in Stud. Mycol. 79: 101. 2014.
Kohlmeyeriopsis
medullaris (Kohlm., Volkm.Kohlm. & O.E. Erikss.)
Klaubauf et al. 2014.
1
Magnaporthiopsis J. Luo & N.
Zhang in Mycologia 105: 1021.
2013.
Magnaporthiopsis poae
(Landsch. & N. Jacks.) J.
Luo & N. Zhang 2013.
5
MAGNAPORTHACEAE
ART I CLE
Table 1. Accepted generic names in Magnaporthales with type species, number of species in each genus, and gene or genomic resources.
Names needing approval are indicated by an asterisk (*).
= Clasterosphaeria Sivan. in
Trans. Brit. mycol. Soc. 83: 710.
1984.
= Harpophora W. Gams in Stud.
Mycol. 45: 192. 2000.
Kohlmeyeriopsis medullaris: SSU(FJ176801),
ITS(KM484852), LSU(FJ176801), RPB1(KM485069)
(Klaubauf et al. 2014).
Magnaporthiopsis incrustans: genome (SRX795321),
transcriptome (SRX798625) (Luo et al. 2015a);
Magnaporthiopsis panicorum: transcriptome
(SRX798626) (Luo et al. 2015a);
Magnaporthiopsis poae: genome (ADBL01000000)
(Okagaki et al. 2015);
Magnaporthiopsis rhizophila: genome (SRX798599),
transcriptome (SRX798627) (Luo et al. 2015a).
Muraeriata Huhndorf et al. in
Mycologia 100: 948. 2008.
VOLUME 7 · NO. 1
Muraeriata collapsa
Huhndorf, Greif, Mugambi &
A.N. Mill. 2008.
2
Muraeriata collapsa: LSU (EU527996) (Huhndorf et
al. 2008).
157
Zhang et al.
ART I CLE
Table 1. (Continued).
Taxa
Type species
Number of
species
GenBank accession numbers for gene and
genome sequence data**
* Nakataea Hara, Diseases Rice
Plant, 2nd : 185. 1939.
Nakataea oryzae (Catt.) J.
Luo & N. Zhang 2013.
7
Nakataea oryzae: genome (SRX798605),
transcriptome (SRX798628) (Luo et al. 2015a).
Omnidemptus P.F. Cannon &
Alcorn in Mycotaxon 51: 483.
1994.
Omnidemptus affinis P.F.
Cannon & Alcorn 1994.
1
Omnidemptus affinis: transcriptome (SRX798629)
(Luo et al. 2015a).
Pseudophialophora J. Luo &
N. Zhang in Mycologia 106: 581.
2014.
Pseudophialophora
eragrostis J. Luo & N.
Zhang 2014.
8
Pseudophialophora eragrostis: transcriptome
(SRX798634) (Luo et al. 2015a);
= Magnaporthe R.A. Krause &
R.K. Webster in Mycologia 64:
110. 1972.
Pseudophialophora panicorum: transcriptome
(SRX798635) (Luo et al. 2015a);
Pseudophialophora schizachyrii: transcriptome
(SRX798637) (Luo et al. 2015a).
Pyriculariopsis M.B. Ellis,
Demat. Hyphom.: 206. 1971.
Pyriculariopsis parasitica
(Sacc. & Berl.) M.B. Ellis
1971.
1
Pyriculariopsis parasitica: LSU(DQ341514) (Klaubauf
et al. 2014).
Slopeiomyces Klaubauf et al. in
Stud. Mycol. 79: 102. 2014.
Slopeiomyces
cylindrosporus (D. Hornby
et al.) Klaubauf et al. 2014.
1
Slopeiomyces cylindrosporus: transcriptome
(SRX798639) (Luo et al. 2015a).
Ophioceras
dolichostomum (Berk. &
M.A. Curtis) Sacc. 1883.
33
Ophioceras dolichostomum: genome (SRX798611)
(Luo et al. 2015a);
OPHIOCERACEAE
Ophioceras Sacc., Syll. Fung. 2:
358. 1883.
Ophioceras commune: transcriptome (SRX798630)
(Luo et al. 2015a);
Ophioceras leptosporum: transcriptome
(SRX798632) (Luo et al. 2015a).
Pseudohalonectria Minoura &
T. Muroi in Trans. Mycol. Soc.
Japan 19: 132. 1978.
Pseudohalonectria
lignicola Minoura & T. Muroi
1978.
13
Pseudohalonectria lignicola: genome (SRX798616),
transcriptome (SRX798633) (Luo et al. 2015a).
Bambusicularia Klaubauf et al.
in Stud. Mycol. 79: 104. 2014.
Bambusicularia brunnea
Klaubauf et al. 2014.
1
Bambusicularia brunnea: ITS(KM484830),
LSU(KM484948), ACT(AB274449), CAL(AB274482),
RPB1(KM485043) (Klaubauf et al. 2014).
Barretomyces Klaubauf et al. in
Stud. Mycol. 79: 104. 2014.
Barretomyces calatheae
(D.J. Soares et al.) Klaubauf
et al. 2014.
1
Barretomyces calatheae: ITS(KM484831),
LSU(KM484950), ACT(KM485162),
CAL(KM485231), RPB1(KM485045) (Klaubauf et al.
2014).
Deightoniella S. Hughes in
Mycol. Pap. 48: 27. 1952.
Deightoniella africana S.
Hughes 1952.
20
Macgarvieomyces Klaubauf et
al. in Stud. Mycol. 79: 106. 2014.
Macgarvieomyces borealis
(de Hoog & Oorschot)
Klaubauf et al. 2014.
2
PYRICULARIACEAE
Macgarvieomyces borealis: SSU(DQ341511),
ITS(KM484854), LSU(DQ341511), ACT(KM485170),
CAL(KM485239), MCM7(KM009174),
RPB1(KM485070), TEF1(KM009198) (Klaubauf et
al. 2014, Luo et al. 2015a);
Macgarvieomyces juncicola: transcriptome
sequenced (SRX798624) (Luo et al. 2015a).
158
Neopyricularia Klaubauf et al. in
Stud. Mycol. 79: 108. 2014.
Neopyricularia
commelinicola (M.J. Park
& H.D. Shin) Klaubauf et al.
2014.
1
Neopyricularia commelinicola: SSU(KM009211),
ITS(FJ850122), LSU(KM484985), ACT(KM485175),
CAL(KM485243), MCM7(KM009175),
RPB1(KM485087), TEF1(KM009199) (Klaubauf et
al. 2014, Luo et al. 2015a).
Proxipyricularia Klaubauf et al.
in Stud. Mycol. 79: 109. 2014.
Proxipyricularia zingiberis
(Y. Nisik.) Klaubauf et al.
2014.
1
Proxipyricularia zingiberis: ITS(KM484869),
LSU(KM484986), ACT(AB274448), CAL(KM485244),
RPB1(KM485088) (Klaubauf et al. 2014).
IMA FUNGUS
Generic names in Magnaporthales
Table 1. (Continued).
Type species
Number of
species
GenBank accession numbers for gene and
genome sequence data**
Pseudopyricularia Klaubauf et
al. in Stud. Mycol. 79: 109. 2014.
Pseudopyricularia
kyllingae Klaubauf et al.
2014.
3
Pseudopyricularia kyllingae: ITS(KM484876),
LSU(KM484992), ACT(AB274451), CAL(AB274484),
RPB1(KM485096) (Klaubauf et al. 2014).
Pyricularia Sacc. in Michelia 2:
20. 1880.
Pyricularia grisea Sacc.
1880.
55
Pyricularia grisea: transcriptome (SRX798638) (Luo
et al. 2015a); genome (PRJEB7653 at http://genome.
jouy.inra.fr/gemo/)
ART I CLE
Taxa
Pyricularia oryzae:
Genome (Dean et al. 2005).
Xenopyricularia Klaubauf et al.
in Stud. Mycol. 79: 116. 2014.
Xenopyricularia zizaniicola
(Hashioka) Klaubauf et al.
2014.
1
Xenopyricularia zizaniicola: transcriptome
(SRX798640) (Luo et al. 2015a).
** Unpublished genome data are not listed.
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