ungal Systematics and Àolution JUNE 2021 PAGES 1–19
19
Fungal Systemacs and Evoluon is licensed under a Creave Commons Aribuon-NonCommercial-ShareAlike 4.0 Internaonal License © 2021 Westerdijk Fungal Biodiversity Instute 1 Fungal Systemacs and Evoluon doi.org/10.3114/fuse.2021.07.01 VOLUME 7 JUNE 2021 PAGES 1–19 INTRODUCTION Bananas (Musa spp.) originated from Indochina and South East Asia (Simmonds 1962), from where they spread across tropical and sub-tropical regions of the world, presently represenng the fourth most economically important food crop aſter rice, wheat and maize (www.fao.org). Bananas and plantains are, however, suscepble to several diseases that have been of serious concern to the industry, namely Fusarium wilt (Panama disease) caused by Fusarium odorassimum (formerly F. oxysporum f. sp. cubense tropical race 4) (Maryani et al. 2019), Banana bunchy top virus (Stainton et al. 2015), and the Sigatoka leaf spot complex, which include the most serious leaf spot diseases of banana (Churchill 2011). Species of the Sigatoka leaf spot complex have a confused taxonomic history, and were formerly treated in the genus Mycosphaerella, which was shown to be polyphylec, represenng numerous genera in the Mycosphaerellaceae (Crous et al. 2007, 2009, Videira et al. 2017), and in other families (Quaedvlieg et al. 2014). Species of the Sigatoka leaf spot complex are members of the genus Pseudocercospora (Arzanlou et al. 2008, Crous et al. 2013, Nakashima et al. 2016). Taxonomically, Pseudocercospora was formerly placed in the order Capnodiales in the class Dothideomycetes, which is the largest and most diverse class of ascomycetous fungi (Haridas et al. 2020). However, the Capnodiales represent sooty moulds that grow superficially on plant surfaces, and are associated with honeydew produced by insects, whereas the genera of plant pathogenic fungi formerly placed in Capnodiales, were shown to be members of the order Mycosphaerellales (Abdollahzadeh et al. 2020). More than 30 mycosphaerella-like species have in the past been associated with leaf spot diseases of banana (see list below). The primary agents of the Sigatoka leaf spot complex found on banana include P. musae (previously: Mycosphaerella musicola) causal agent of Sigatoka leaf spot or yellow Sigatoka; P. eumusae (previously: M. eumusae) causal agent of the eumusae leaf spot disease, and P. fijiensis (previously: M. fijiensis) causal agent of black Sigatoka or black leaf streak disease, which is the most aggressive and predominant member of the Sigatoka lead spot complex worldwide. Phylogenec reconstrucon based on a set of 46 conserved single-copy genes strongly supported an earlier evoluonary radiaon of P. fijiensis from P. musae and P. eumusae (between 15 to 40 MYA, Chang et al. 2016). Although Pseudocercospora and allied genera associated with leaf spots of banana (Musa spp.) P.W. Crous 1,2,3* , J. Carlier 4 , V. Roussel 4 , J.Z. Groenewald 1 1 Westerdijk Fungal Biodiversity Instute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands 2 Department of Biochemistry, Genecs and Microbiology, Forestry and Agricultural Biotechnology Instute (FABI), University of Pretoria, Pretoria, 0002, South Africa 3 Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands 4 Centre de Coopéraon Internaonal en Recherche Agronomique pour le Développement (CIRAD), TA 40/02, avenue Agropolis, 34 398 Montpellier, France *Corresponding author: [email protected] Abstract: The Sigatoka leaf spot complex on Musa spp. includes three major pathogens: Pseudocercospora, namely P. musae (Sigatoka leaf spot or yellow Sigatoka), P. eumusae (eumusae leaf spot disease), and P. fijiensis (black leaf streak disease or black Sigatoka). However, more than 30 species of Mycosphaerellaceae have been associated with Sigatoka leaf spots of banana, and previous reports of P. musae and P. eumusae need to be re-evaluated in light of recently described species. The aim of the present study was thus to invesgate a global set of 228 isolates of P. musae, P. eumusae and close relaves on banana using mulgene DNA sequence data [internal transcribed spacer regions with intervening 5.8S nrRNA gene (ITS), RNA polymerase II second largest subunit gene (rpb2), translaon elongaon factor 1-alpha gene (tef1), beta-tubulin gene (tub2), and the acn gene (act)] to confirm if these isolates represent P. musae, or a closely allied species. Based on these data one new species is described, namely P. pseudomusae, which is associated with leaf spot symptoms resembling those of P. musae on Musa in Indonesia. Furthermore, P. eumusae, P. musae and P. fijiensis are shown to be well defined taxa, with some isolates also represenng P. longispora. Other genera encountered in the dataset are species of Zasmidium (Taiwan leaf speckle), Metulocladosporiella (Cladosporium leaf speckle) and Scolecobasidium leaf speckle. Key words: mul-gene phylogeny Mycosphaerella new taxa Sigatoka leaf spots systemacs Citaon: Crous P, Carlier J, Roussel V, Groenewald JZ (2020). Pseudocercospora and allied genera associated with leaf spots of banana (Musa spp.). Fungal Systemacs and Evoluon 7: 1–19. doi: 10.3114/fuse.2021.07.01 Received: 1 September 2020; Accepted: 23 October 2020; Effectively published online: 30 October 2020 Corresponding editor: U. Braun
Transcript of ungal Systematics and Àolution JUNE 2021 PAGES 1–19
Fungal Systematics and Evolution is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License
© 2021 Westerdijk Fungal Biodiversity Institute 1
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
Fungal Systematics and Evolution
doi.org/10.3114/fuse.2021.07.01
VOLUME 7JUNE 2021PAGES 1–19
INTRODUCTION
Bananas (Musa spp.) originated from Indochina and South East Asia (Simmonds 1962), from where they spread across tropical and sub-tropical regions of the world, presently representing the fourth most economically important food crop after rice, wheat and maize (www.fao.org).
Bananas and plantains are, however, susceptible to several diseases that have been of serious concern to the industry, namely Fusarium wilt (Panama disease) caused by Fusarium odoratissimum (formerly F. oxysporum f. sp. cubense tropical race 4) (Maryani et al. 2019), Banana bunchy top virus (Stainton et al. 2015), and the Sigatoka leaf spot complex, which include the most serious leaf spot diseases of banana (Churchill 2011).
Species of the Sigatoka leaf spot complex have a confused taxonomic history, and were formerly treated in the genus Mycosphaerella, which was shown to be polyphyletic, representing numerous genera in the Mycosphaerellaceae (Crous et al. 2007, 2009, Videira et al. 2017), and in other families (Quaedvlieg et al. 2014). Species of the Sigatoka leaf spot complex are members of the genus Pseudocercospora (Arzanlou et al. 2008, Crous et al. 2013, Nakashima et al. 2016).
Taxonomically, Pseudocercospora was formerly placed in the order Capnodiales in the class Dothideomycetes, which is the largest and most diverse class of ascomycetous fungi (Haridas et al. 2020). However, the Capnodiales represent sooty moulds that grow superficially on plant surfaces, and are associated with honeydew produced by insects, whereas the genera of plant pathogenic fungi formerly placed in Capnodiales, were shown to be members of the order Mycosphaerellales (Abdollahzadeh et al. 2020).
More than 30 mycosphaerella-like species have in the past been associated with leaf spot diseases of banana (see list below). The primary agents of the Sigatoka leaf spot complex found on banana include P. musae (previously: Mycosphaerella musicola) causal agent of Sigatoka leaf spot or yellow Sigatoka; P. eumusae (previously: M. eumusae) causal agent of the eumusae leaf spot disease, and P. fijiensis (previously: M. fijiensis) causal agent of black Sigatoka or black leaf streak disease, which is the most aggressive and predominant member of the Sigatoka lead spot complex worldwide. Phylogenetic reconstruction based on a set of 46 conserved single-copy genes strongly supported an earlier evolutionary radiation of P. fijiensis from P. musae and P. eumusae (between 15 to 40 MYA, Chang et al. 2016). Although
Pseudocercospora and allied genera associated with leaf spots of banana (Musa spp.)
P.W. Crous1,2,3*, J. Carlier4, V. Roussel4, J.Z. Groenewald1
1Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands2Department of Biochemistry, Genetics and Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa3Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands4Centre de Coopération International en Recherche Agronomique pour le Développement (CIRAD), TA 40/02, avenue Agropolis, 34 398 Montpellier, France
*Corresponding author: [email protected]
Abstract: The Sigatoka leaf spot complex on Musa spp. includes three major pathogens: Pseudocercospora, namely P. musae (Sigatoka leaf spot or yellow Sigatoka), P. eumusae (eumusae leaf spot disease), and P. fijiensis (black leaf streak disease or black Sigatoka). However, more than 30 species of Mycosphaerellaceae have been associated with Sigatoka leaf spots of banana, and previous reports of P. musae and P. eumusae need to be re-evaluated in light of recently described species. The aim of the present study was thus to investigate a global set of 228 isolates of P. musae, P. eumusae and close relatives on banana using multigene DNA sequence data [internal transcribed spacer regions with intervening 5.8S nrRNA gene (ITS), RNA polymerase II second largest subunit gene (rpb2), translation elongation factor 1-alpha gene (tef1), beta-tubulin gene (tub2), and the actin gene (act)] to confirm if these isolates represent P. musae, or a closely allied species. Based on these data one new species is described, namely P. pseudomusae, which is associated with leaf spot symptoms resembling those of P. musae on Musa in Indonesia. Furthermore, P. eumusae, P. musae and P. fijiensis are shown to be well defined taxa, with some isolates also representing P. longispora. Other genera encountered in the dataset are species of Zasmidium (Taiwan leaf speckle), Metulocladosporiella (Cladosporium leaf speckle) and Scolecobasidium leaf speckle.
Key words: multi-gene phylogenyMycosphaerellanew taxaSigatoka leaf spotssystematics
Citation: Crous P, Carlier J, Roussel V, Groenewald JZ (2020). Pseudocercospora and allied genera associated with leaf spots of banana (Musa spp.). Fungal Systematics and Evolution 7: 1–19. doi: 10.3114/fuse.2021.07.01Received: 1 September 2020; Accepted: 23 October 2020; Effectively published online: 30 October 2020Corresponding editor: U. Braun
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
2
these species can be distinguished based on differences in morphology and symptomatology, considerable overlap exists among them, making definite identification only possible by means of additional markers such as DNA sequence data (Arzanlou et al. 2008). Furthermore, several previous records of P. musae might in fact belong to the recently described P. longispora, while the variation within P. eumusae (Crous & Mourichon 2002, Arzanlou et al. 2008) also requires further analysis. Because the distribution and relative importance of the latter taxa remain insufficiently known, the aim of the present study was to investigate a global set of isolates to better elucidate the distribution of Pseudocercospora spp. associated with leaf spot diseases of banana.
MATERIALS AND METHODS
Isolates
The set of isolates studied is presented in Table 1. Isolates were sourced from the CBS and CIRAD collections and were collected from various countries. The isolates from the CIRAD collection were derived either from single conidia or ascospores as described in Zapater et al. (2008). They were assumed to belong to P. musae, P. eumusae or other species based on morphological observations in culture media or sequencing of the internal transcribed spacer regions with intervening 5.8S nrRNA gene (ITS) of the nrDNA operon. The isolates from CBS collection were also derived either from single conidia or ascospore as described by Crous et al. (1991) and Crous (1998). In the present study, colonies were sub-cultured on 2 % potato-dextrose agar (PDA), oatmeal agar (OA), MEA (Crous et al. 2019), autoclaved banana leaf on 2 % tap water agar (BLA), and incubated at 25 °C under continuous near-ultraviolet light to promote sporulation. Reference strains and specimens of the studied fungi are all maintained from now in the CBS culture collection (CBS) of the Westerdijk Fungal Biodiversity Institute (WI), Utrecht, the Netherlands.
DNA extraction, amplification (PCR) and phylogeny
Fungal mycelium (Table 1) was scraped from the agar surface of cultures with a sterile scalpel and genomic DNA was isolated using the Wizard® Genomic DNA Purification Kit (Promega Corporation, WI, USA) following the manufacturers’ protocols. Eight loci were amplified following previously published protocols. First, the partial ITS region was sequenced for all isolates included in this study (for amplification conditions, see Fan et al. 2018). Amplification of the partial DNA-directed RNA polymerase II second largest subunit gene (rpb2), the partial translation elongation factor 1-alpha gene (tef1) and the partial beta-tubulin gene (tub2) followed Braun et al. (2018), while the amplification of the partial actin gene (act) followed Videira et al. (2016). The resulting fragments were sequenced in both directions using the respective PCR primers and the BigDye Terminator Cycle Sequencing Kit v. 3.1 (Applied Biosystems Life Technologies, Carlsbad, CA, USA); DNA sequencing amplicons were purified through Sephadex G-50 Superfine columns (Sigma-Aldrich, St. Louis, MO) in MultiScreen HV plates (Millipore, Billerica, MA). Purified sequence reactions were analysed on an Applied Biosystems 3730xl DNA Analyzer (Life Technologies, Carlsbad, CA, USA). The DNA sequences were analysed and
consensus sequences were computed using SeqMan Pro v. 13 (DNASTAR, Madison, WI, USA).
The sequences for each gene region were subjected to megablast searches (Zhang et al. 2000) to identify closely related sequences in the NCBI’s GenBank nucleotide database. Sequences of the individual loci were aligned using MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server/index.html) (Katoh & Standley 2013), and the alignments were then manually edited in MEGA v. 7.0.21. Sequence Matrix v. 1.8 (http://www.ggvaidya.com/taxondna/) was used to concatenate the individual loci in various combinations. Phylogenetic trees on the combined datasets were generated using Bayesian analyses performed with MrBayes v. 3.2.7 (Ronquist et al. 2012) as explained in Braun et al. (2018) and Pseudocercospora individual gene trees were evaluated using distance and parsimony analyses with PAUP v. 4.0b10 (Swofford 2003). All resulting trees were printed with Geneious v. 11.1.5 3 (http://www.geneious.com, Kearse et al. 2012) and the layout of the trees was done in Adobe Illustrator v. CC 2017.
Morphology
Slide preparations were mounted in clear lactic acid or Shear’s mounting fluid. Observations were made with a Nikon SMZ25 dissection microscope, and with a Zeiss Axio Imager 2 light microscope using differential interference contrast (DIC) illumination and images recorded on a Nikon DS-Ri2 camera with associated software. Colony characters and pigment production were noted after 2–4 wk of growth on MEA, PDA and OA (Crous et al. 2019) incubated at 25 °C. Colony colours (surface and reverse) were scored using the colour charts of Rayner (1970). Sequences derived in this study were deposited in GenBank (Table 1), the alignment in TreeBASE (www.treebase.org; study number 27043), and taxonomic novelties in MycoBank (www.MycoBank.org; Crous et al. 2004).
RESULTS
Phylogeny
Based on the blast results, the majority of strains belonged to Pseudocercospora musae (104 strains) and Pseudocercospora eumusae (48 strains), while other Pseudocercospora species included Pseudocercospora fijiensis (two strains), Pseudocercospora longispora (two strains) and five strains representing a novel Pseudocercospora species related to Pseudocercospora musae (Fig. 1). The remainder of the strains tentatively identified as Mycosphaerella musae based on symptomatology (seven strains), Zasmidium musae (five strains), Scolecobasidium musicola (= Ochroconis musicola) (four strains, Fig. 2), Parapallidocercospora thailandica (three strains), while one strain was identical to Pantospora guazumae and the last strain represented a Penicillium infection and was discarded from further analyses (Table 1).
Two Bayesian analyses were performed (Table 2); the first on a concatenated ITS/act/tef1/rpb2 alignment of Pseudocercospora sequences (Fig. 1) and the second on a concatenated ITS/act/tef1/tub2 alignment of Scolecobasidium sequences (Fig. 2). The Pseudocercospora phylogeny (Fig. 1) delimits four known species clades, namely Pseudocercospora eumusae, Pseudocercospora fijiensis, Pseudocercospora longispora and Pseudocercospora musae, with five strains from Musa (Indonesia) clustering in a
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
3
Tabl
e 1.
Col
lecti
on in
form
ation
and
Gen
Bank
acc
essio
n nu
mbe
rs o
f str
ains
incl
uded
in th
is st
udy.
Tax
onom
ic n
ovel
ties a
nd se
quen
ces g
ener
ated
in th
is st
udy
are
show
n in
bol
d. n
/a: n
ot a
pplic
able
.
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
Pseu
doce
rcos
pora
eum
usae
CBS
1148
24 =
STE
-U 4
579
= CI
RAD
1156
, ex-
type
Mus
a; F
ranc
e: R
éuni
on; 2
001
EU51
4238
.1LF
ZN01
0000
53.1
LFZN
0100
0037
.1; L
FZN
0100
0034
.1; ‒
CBS
1213
78M
usa;
Mal
aysia
; ‒EU
5142
42.1
EU51
4305
.1‒;
‒; ‒
CBS
1213
80M
usa;
Sri
Lank
a; ‒
EU51
4243
.1EU
5143
06.1
‒; ‒
; ‒
CIRA
D 53
5M
usa;
Indi
a; ‒
AY92
3759
.1‒
‒; ‒
; ‒
CPC
3731
5 =
CIRA
D-VN
M 0
03M
usa;
Vie
tnam
; 199
5M
W06
3266
.1M
W07
0615
.1M
W07
0937
.1; M
W07
0796
.1; ‒
CPC
3731
6 =
CIRA
D-VN
M 0
04M
usa;
Vie
tnam
; 199
5M
W06
3267
.1M
W07
0616
.1M
W07
0938
.1; M
W07
0797
.1; ‒
CPC
3731
8 =
CIRA
D-M
YS 0
02M
usa;
Mal
aysia
; 200
1M
W06
3268
.1M
W07
0617
.1M
W07
0939
.1; M
W07
0798
.1; ‒
CPC
3731
9 =
CIRA
D-M
YS 0
03M
usa;
Mal
aysia
; 200
1M
W06
3269
.1M
W07
0618
.1M
W07
0940
.1; M
W07
0799
.1; ‒
CPC
3732
1 =
CIRA
D-M
YS 0
57M
usa;
Mal
aysia
; 199
3M
W06
3270
.1M
W07
0619
.1M
W07
0941
.1; M
W07
0800
.1; ‒
CPC
3732
2 =
CIRA
D-M
YS 0
58M
usa;
Mal
aysia
; 199
3M
W06
3271
.1M
W07
0620
.1M
W07
0942
.1; M
W07
0801
.1; ‒
CPC
3732
3 =
CIRA
D-N
GA 0
52M
usa;
Nig
eria
; 199
9M
W06
3272
.1M
W07
0621
.1M
W07
0943
.1; M
W07
0802
.1; ‒
CPC
3732
4 =
CIRA
D-N
GA 0
53M
usa;
Nig
eria
; 199
9M
W06
3273
.1M
W07
0622
.1M
W07
0944
.1; M
W07
0803
.1; ‒
CPC
3732
5 =
CIRA
D-N
GA 0
57M
usa;
Nig
eria
; 198
9M
W06
3274
.1M
W07
0623
.1M
W07
0945
.1; M
W07
0804
.1; ‒
CPC
3732
6 =
CIRA
D-N
GA 0
58M
usa;
Nig
eria
; 198
9M
W06
3275
.1M
W07
0624
.1M
W07
0946
.1; M
W07
0805
.1; ‒
CPC
3732
7 =
CIRA
D-N
GA 0
66M
usa;
Nig
eria
; 199
0M
W06
3276
.1M
W07
0625
.1M
W07
0947
.1; M
W07
0806
.1; ‒
CPC
3733
0 =
CIRA
D-RE
U 0
02M
usa;
Fra
nce:
Réu
nion
; 199
9M
W06
3277
.1M
W07
0626
.1‒;
MW
0708
07.1
; ‒
CPC
3733
1 =
CIRA
D-RE
U 0
03M
usa;
Fra
nce:
Réu
nion
; 199
9M
W06
3278
.1M
W07
0627
.1M
W07
0948
.1; M
W07
0808
.1; ‒
CPC
3733
2 =
CIRA
D-RE
U 0
10M
usa;
Fra
nce:
Réu
nion
; 200
1M
W06
3279
.1M
W07
0628
.1M
W07
0949
.1; M
W07
0809
.1; ‒
CPC
3733
3 =
CIRA
D-RE
U 0
11M
usa;
Fra
nce:
Réu
nion
; 200
1M
W06
3280
.1M
W07
0629
.1M
W07
0950
.1; M
W07
0810
.1; ‒
CPC
3733
4 =
CIRA
D-RE
U 0
12M
usa;
Fra
nce:
Réu
nion
; 200
1M
W06
3281
.1M
W07
0630
.1M
W07
0951
.1; M
W07
0811
.1; ‒
CPC
3733
5 =
CIRA
D-RE
U 0
13M
usa;
Fra
nce:
Réu
nion
; 200
1M
W06
3282
.1M
W07
0631
.1M
W07
0952
.1; M
W07
0812
.1; ‒
CPC
3733
6 =
CIRA
D-RE
U 0
35M
usa;
Fra
nce:
Réu
nion
; 200
2M
W06
3283
.1M
W07
0632
.1M
W07
0953
.1; M
W07
0813
.1; ‒
CPC
3733
7 =
CIRA
D-RE
U 0
36M
usa;
Fra
nce:
Réu
nion
; 200
2M
W06
3284
.1M
W07
0633
.1M
W07
0954
.1; M
W07
0814
.1; ‒
CPC
3733
8 =
CIRA
D-M
US
001
Mus
a; Îl
e M
auric
e (M
auriti
us);
2003
MW
0632
85.1
MW
0706
34.1
MW
0709
55.1
; MW
0708
15.1
; ‒
CPC
3733
9 =
CIRA
D-M
US
002
Mus
a; Îl
e M
auric
e (M
auriti
us);
2003
MW
0632
86.1
MW
0706
35.1
MW
0709
56.1
; MW
0708
16.1
; ‒
CPC
3734
0 =
CIRA
D-M
US
009
Mus
a; Îl
e M
auric
e (M
auriti
us);
2003
MW
0632
87.1
MW
0706
36.1
MW
0709
57.1
; MW
0708
17.1
; ‒
CPC
3734
1 =
CIRA
D-M
US
010
Mus
a; Îl
e M
auric
e (M
auriti
us);
2003
MW
0632
88.1
MW
0706
37.1
MW
0709
58.1
; MW
0708
18.1
; ‒
CPC
3734
2 =
CIRA
D-M
US
011
Mus
a; Îl
e M
auric
e (M
auriti
us);
1996
MW
0632
89.1
MW
0706
38.1
MW
0709
59.1
; MW
0708
19.1
; ‒
CPC
3734
3 =
CIRA
D-M
US
012
Mus
a; Îl
e M
auric
e (M
auriti
us);
1996
MW
0632
90.1
MW
0706
39.1
MW
0709
60.1
; MW
0708
20.1
; ‒
CPC
3734
4 =
CIRA
D-M
US
015
Mus
a; Îl
e M
auric
e (M
auriti
us);
1996
MW
0632
91.1
MW
0706
40.1
MW
0709
61.1
; MW
0708
21.1
; ‒
CPC
3734
5 =
CIRA
D-M
US
016
Mus
a; Îl
e M
auric
e (M
auriti
us);
1996
MW
0632
92.1
MW
0706
41.1
MW
0709
62.1
; MW
0708
22.1
; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
4
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
CPC
3734
6 =
CIRA
D-IN
D 00
1M
usa;
Indi
a; 1
999
MW
0632
93.1
MW
0706
42.1
MW
0709
63.1
; MW
0708
23.1
; ‒
CPC
3734
7 =
CIRA
D-IN
D 00
2M
usa;
Indi
a; 1
999
MW
0632
94.1
MW
0706
43.1
MW
0709
64.1
; MW
0708
24.1
; ‒
CPC
3734
9 =
CIRA
D-IN
D 00
9M
usa;
Indi
a; ‒
MW
0632
95.1
MW
0706
44.1
MW
0709
65.1
; MW
0708
25.1
; ‒
CPC
3735
0 =
CIRA
D-IN
D 01
0M
usa;
Indi
a; 1
998
MW
0632
96.1
MW
0706
45.1
MW
0709
66.1
; MW
0708
26.1
; ‒
CPC
3735
1 =
CIRA
D-IN
D 01
1M
usa;
Indi
a; 1
995
MW
0632
97.1
MW
0706
46.1
MW
0709
67.1
; MW
0708
27.1
; ‒
CPC
3735
2 =
CIRA
D-IN
D 03
1M
usa;
Indi
a; 1
992
MW
0632
98.1
MW
0706
47.1
MW
0709
68.1
; MW
0708
28.1
; ‒
CPC
3735
3 =
CIRA
D-IN
D 03
2M
usa;
Indi
a; 1
992
MW
0632
99.1
MW
0706
48.1
MW
0709
69.1
; MW
0708
29.1
; ‒
CPC
3735
4 =
CIRA
D-LK
A 00
1M
usa;
Sri
Lank
a; ‒
MW
0633
00.1
MW
0706
49.1
MW
0709
70.1
; MW
0708
30.1
; ‒
CPC
3735
5 =
CIRA
D-LK
A 00
2M
usa;
Sri
Lank
a; 1
999
MW
0633
01.1
MW
0706
50.1
MW
0709
71.1
; MW
0708
31.1
; ‒
CPC
3735
6 =
CIRA
D-LK
A 00
3M
usa;
Sri
Lank
a; 1
999
MW
0633
02.1
MW
0706
51.1
MW
0709
72.1
; ‒; ‒
CPC
3735
7 =
CIRA
D-LK
A 00
6M
usa;
Sri
Lank
a; 1
995
MW
0633
03.1
MW
0706
52.1
MW
0709
73.1
; MW
0708
32.1
; ‒
CPC
3735
8 =
CIRA
D-LK
A 00
7M
usa;
Sri
Lank
a; 1
995
MW
0633
04.1
MW
0706
53.1
MW
0709
74.1
; MW
0708
33.1
; ‒
CPC
3735
9 =
CIRA
D-LK
A 01
6M
usa;
Sri
Lank
a; 1
995
MW
0633
05.1
MW
0706
54.1
MW
0709
75.1
; MW
0708
34.1
; ‒
CPC
3736
0 =
CIRA
D-LK
A 01
7M
usa;
Sri
Lank
a; 1
995
MW
0633
06.1
MW
0706
55.1
MW
0709
76.1
; MW
0708
35.1
; ‒
CPC
3736
1 =
CIRA
D-TH
A 00
1M
usa;
Tha
iland
; 199
4M
W06
3307
.1M
W07
0656
.1M
W07
0977
.1; ‒
; ‒
CPC
3736
2 =
CIRA
D-TH
A 00
2M
usa;
Tha
iland
; 199
4M
W06
3308
.1M
W07
0657
.1M
W07
0978
.1; M
W07
0836
.1; ‒
CPC
3736
3 =
CIRA
D-TH
A 00
5M
usa;
Tha
iland
; 199
4M
W06
3309
.1M
W07
0658
.1‒;
MW
0708
37.1
; ‒
CPC
3736
4 =
CIRA
D-TH
A 00
6M
usa;
Tha
iland
; 199
4M
W06
3310
.1M
W07
0659
.1M
W07
0979
.1; M
W07
0838
.1; ‒
CPC
3736
5 =
CIRA
D-TH
A 00
7M
usa;
Tha
iland
; 199
4M
W06
3311
.1M
W07
0660
.1M
W07
0980
.1; M
W07
0839
.1; ‒
CPC
3736
6 =
CIRA
D-TH
A 00
8M
usa;
Tha
iland
; 199
4M
W06
3312
.1M
W07
0661
.1M
W07
0981
.1; M
W07
0840
.1; ‒
CPC
3736
7 =
CIRA
D-TH
A 00
9M
usa;
Tha
iland
; 199
4M
W06
3313
.1M
W07
0662
.1M
W07
0982
.1; M
W07
0841
.1; ‒
S103
0BM
usa;
Mau
ritius
; ‒EU
5142
33.1
EU51
4300
.1‒;
‒; ‒
S103
7BM
usa;
Mau
ritius
; ‒EU
5142
34.1
EU51
4301
.1‒;
‒; ‒
S17
Mus
a; In
dia;
‒M
N86
0088
.1‒
‒; ‒
; ‒
S37
Mus
a cv
. Nen
dran
; Ind
ia; ‒
MN
9019
97.1
‒‒;
‒; ‒
S48
Mus
a; In
dia;
‒M
N94
7254
.1‒
‒; ‒
; ‒
Pseu
doce
rcos
pora
fijie
nsis
CBS
1202
58 =
CIR
AD 8
6, e
x-ep
itype
Mus
a; C
amer
oon;
‒EU
5142
48.1
NW
_006
9215
33.1
NW
_006
9215
32.1
; NW
_006
9215
35.1
; ‒
CBS
1213
62 =
CIR
AD 3
64 =
X84
7M
usa;
Tai
wan
; ‒EU
5142
54.1
EU51
4316
.1‒;
‒; ‒
CIRA
D 11
= X
843
Mus
a; H
ondu
ras;
‒EU
5142
53.1
EU51
4315
.1‒;
‒; ‒
CIRA
D 35
5 =
X850
Mus
a; Iv
ory
Coas
t; ‒
EU51
4255
.1‒
‒; ‒
; ‒
CPC
1630
1M
usa;
Mex
ico;
‒KX
4625
81.1
KX46
2548
.1KX
4626
67.1
; KX4
6261
3.1;
‒
CPC
3721
6 =
CMR
1853
Mus
a; C
amer
oon;
200
7M
W06
3314
.1M
W07
0663
.1M
W07
0983
.1; M
W07
0842
.1; ‒
CPC
3721
7 =
CIRA
D-CM
R 18
61M
usa;
Cam
eroo
n; 1
996
MW
0633
15.1
MW
0706
64.1
MW
0709
84.1
; MW
0708
43.1
; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
5
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
X104
Mus
a; C
olom
bia;
‒EU
5142
51.1
‒‒;
‒; ‒
Pseu
doce
rcos
pora
long
ispor
aCB
S 12
2469
= X
474
Mus
a; M
alay
sia; 1
988
EU51
4284
.1EU
5143
41.1
‒; ‒
; ‒
CBS
1224
70 =
X47
5, e
x-ty
peM
usa
cv. P
isang
Byo
k AA
A/AA
B;
Mal
aysia
; 198
8N
R_15
6515
.1EU
5143
42.1
GU38
4447
.1; ‒
; ‒
CPC
3731
7 =
CIRA
D-M
YS 0
01M
usa;
Mal
aysia
; 198
8M
W06
3316
.1M
W07
0665
.1M
W07
0985
.1; ‒
; ‒
CPC
3732
0 =
CIRA
D-M
YS 0
17M
usa;
Mal
aysia
; 198
8M
W06
3317
.1M
W07
0666
.1‒;
‒; ‒
Pseu
doce
rcos
pora
mus
aeCB
S 11
6634
= IM
I 123
823
= X4
2, e
x-ep
itype
Mus
a; C
uba;
‒EU
5142
65.1
LFZO
0100
0469
.1LF
ZO01
0000
01.1
; LFZ
O01
0004
53.1
; ‒
CBS
143.
36 =
ATC
C 10
688
‒; S
urin
ame;
‒M
H855
742.
1‒
‒; ‒
; ‒
CPC
3718
6 =
CIRA
D-AU
S 00
1M
usa;
Aus
tral
ia: N
ew S
outh
Wal
es;
1999
MW
0633
18.1
MW
0706
67.1
MW
0709
86.1
; MW
0708
44.1
; ‒
CPC
3718
7 =
CIRA
D-AU
S 00
2M
usa;
Aus
tral
ia: N
ew S
outh
Wal
es;
1999
MW
0633
19.1
MW
0706
68.1
MW
0709
87.1
; MW
0708
45.1
; ‒
CPC
3718
8 =
CIRA
D-AU
S 00
7M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
9M
W06
3320
.1M
W07
0669
.1M
W07
0988
.1; M
W07
0846
.1; ‒
CPC
3718
9 =
CIRA
D-AU
S 00
8M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
9M
W06
3321
.1M
W07
0670
.1M
W07
0989
.1; M
W07
0847
.1; ‒
CPC
3719
0 =
CIRA
D-AU
S 01
2M
usa;
Aus
tral
ia; 1
999
MW
0633
22.1
MW
0706
71.1
MW
0709
90.1
; MW
0708
48.1
; ‒
CPC
3719
2 =
CIRA
D-AU
S 12
7M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
4M
W06
3323
.1M
W07
0672
.1M
W07
0991
.1; M
W07
0849
.1; ‒
CPC
3719
3 =
CIRA
D-AU
S 12
8M
usa;
Aus
tral
ia; 1
992
MW
0633
24.1
MW
0706
73.1
MW
0709
92.1
; MW
0708
50.1
; ‒
CPC
3719
4 =
CIRA
D-AU
S 12
9M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
2M
W06
3325
.1M
W07
0674
.1M
W07
0993
.1; M
W07
0851
.1; ‒
CPC
3719
6 =
CIRA
D-AU
S 13
1M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
3M
W06
3326
.1M
W07
0675
.1M
W07
0994
.1; M
W07
0852
.1; ‒
CPC
3719
7 =
CIRA
D-BR
A 00
3M
usa;
Bra
zil; 1
983
MW
0633
27.1
MW
0706
76.1
MW
0709
95.1
; MW
0708
53.1
; ‒
CPC
3719
8 =
CIRA
D-BR
A 00
4M
usa;
Bra
zil; 1
983
MW
0633
28.1
MW
0706
77.1
MW
0709
96.1
; ‒; ‒
CPC
3720
1 =
CIRA
D-CI
V 09
1M
usa;
Cot
e d’
ivoi
re (I
vory
Coa
st);
1985
MW
0633
29.1
MW
0706
78.1
MW
0709
97.1
; MW
0708
54.1
; ‒
CPC
3720
2 =
CIRA
D-CI
V 09
2M
usa;
Cot
e d’
ivoi
re (I
vory
Coa
st);
1986
MW
0633
30.1
MW
0706
79.1
MW
0709
98.1
; MW
0708
55.1
; ‒
CPC
3720
3 =
CIRA
D-CM
R 11
75M
usa;
Cam
eroo
n; 1
999
MW
0633
31.1
MW
0706
80.1
MW
0709
99.1
; MW
0708
56.1
; ‒
CPC
3720
4 =
CIRA
D-CM
R 11
76M
usa;
Cam
eroo
n; 1
999
MW
0633
32.1
MW
0706
81.1
MW
0710
00.1
; MW
0708
57.1
; ‒
CPC
3720
5 =
CIRA
D-CM
R 12
92M
usa;
Cam
eroo
n; 1
999
MW
0633
33.1
MW
0706
82.1
MW
0710
01.1
; MW
0708
58.1
; ‒
CPC
3720
6 =
CIRA
D-CM
R 12
93M
usa;
Cam
eroo
n; 1
999
MW
0633
34.1
MW
0706
83.1
MW
0710
02.1
; MW
0708
59.1
; ‒
CPC
3720
7 =
CIRA
D-CM
R 13
28M
usa;
Cam
eroo
n; 1
999
MW
0633
35.1
MW
0706
84.1
MW
0710
03.1
; ‒; ‒
CPC
3720
8 =
CIRA
D-CM
R 13
29M
usa;
Cam
eroo
n; 1
999
MW
0633
36.1
MW
0706
85.1
MW
0710
04.1
; ‒; ‒
CPC
3720
9 =
CIRA
D-CM
R 28
57M
usa;
Cam
eroo
n; 1
988
MW
0633
37.1
MW
0706
86.1
MW
0710
05.1
; MW
0708
60.1
; ‒
CPC
3721
0 =
CIRA
D-CM
R 28
56M
usa;
Cam
eroo
n; 1
988
MW
0633
38.1
MW
0706
87.1
MW
0710
06.1
; MW
0708
61.1
; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
6
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
CPC
3721
1 =
CIRA
D-CM
R 28
59M
usa;
Cam
eroo
n; 1
987
MW
0633
39.1
MW
0706
88.1
MW
0710
07.1
; MW
0708
62.1
; ‒
CPC
3721
2 =
CIRA
D-CM
R 28
61M
usa;
Cam
eroo
n; 1
987
MW
0633
40.1
MW
0706
89.1
MW
0710
08.1
; MW
0708
63.1
; ‒
CPC
3721
9 =
CIRA
D-CO
L 05
6M
usa;
Col
ombi
a; 1
987
MW
0633
41.1
MW
0706
90.1
MW
0710
09.1
; MW
0708
64.1
; ‒
CPC
3722
0 =
CIRA
D-CO
L 05
7M
usa;
Col
ombi
a; 1
988
MW
0633
42.1
MW
0706
91.1
MW
0710
10.1
; ‒; ‒
CPC
3722
1 =
CIRA
D-CO
L 05
9M
usa;
Col
ombi
a; 1
989
MW
0633
43.1
MW
0706
92.1
MW
0710
11.1
; ‒; ‒
CPC
3722
2 =
CIRA
D-CO
L 06
7M
usa;
Col
ombi
a; 1
990
MW
0633
44.1
MW
0706
93.1
MW
0710
12.1
; ‒; ‒
CPC
3722
3 =
CIRA
D-CR
I 001
Mus
a; C
osta
Ric
a; 1
991
MW
0633
45.1
MW
0706
94.1
MW
0710
13.1
; ‒; ‒
CPC
3722
4 =
CIRA
D-CU
B 04
59M
usa;
Cub
a; 1
986
MW
0633
46.1
MW
0706
95.1
MW
0710
14.1
; MW
0708
65.1
; ‒
CPC
3722
5 =
CIRA
D-GI
N 0
02M
usa;
Gui
nea;
199
4M
W06
3347
.1M
W07
0696
.1M
W07
1015
.1; ‒
; ‒
CPC
3722
6 =
CIRA
D-GI
N 0
03M
usa;
Gui
nea;
199
4M
W06
3348
.1M
W07
0697
.1M
W07
1016
.1; ‒
; ‒
CPC
3722
7 =
CIRA
D-GI
N 0
04M
usa;
Gui
nea;
199
4M
W06
3349
.1M
W07
0698
.1M
W07
1017
.1; ‒
; ‒
CPC
3722
8 =
CIRA
D-GI
N 0
05M
usa;
Gui
nea;
199
4M
W06
3350
.1M
W07
0699
.1M
W07
1018
.1; ‒
; ‒
CPC
3722
9 =
CIRA
D-GI
N 0
07M
usa;
Gui
nea;
199
4M
W06
3351
.1M
W07
0700
.1M
W07
1019
.1; ‒
; ‒
CPC
3723
0 =
CIRA
D-GI
N 0
08M
usa;
Gui
nea;
199
4M
W06
3352
.1M
W07
0701
.1M
W07
1020
.1; ‒
; ‒
CPC
3723
1 =
CIRA
D-GL
P 01
01M
usa;
Gua
delo
upe;
200
3M
W06
3353
.1M
W07
0702
.1M
W07
1021
.1; ‒
; ‒
CPC
3723
2 =
CIRA
D-GL
P 01
02M
usa;
Gua
delo
upe;
200
3M
W06
3354
.1M
W07
0703
.1M
W07
1022
.1; ‒
; ‒
CPC
3723
3 =
CIRA
D-GL
P 01
30M
usa;
Gua
delo
upe;
200
3M
W06
3355
.1M
W07
0704
.1M
W07
1023
.1; M
W07
0866
.1; ‒
CPC
3723
4 =
CIRA
D-GL
P 01
31M
usa;
Gua
delo
upe;
200
3M
W06
3356
.1M
W07
0705
.1M
W07
1024
.1; M
W07
0867
.1; ‒
CPC
3723
5 =
CIRA
D-GL
P 01
60M
usa;
Gua
delo
upe;
200
3M
W06
3357
.1M
W07
0706
.1M
W07
1025
.1; M
W07
0868
.1; ‒
CPC
3723
6 =
CIRA
D-GL
P 07
75M
usa;
Gua
delo
upe;
200
3M
W06
3358
.1M
W07
0707
.1M
W07
1026
.1; M
W07
0869
.1; ‒
CPC
3723
7 =
CIRA
D-GL
P 01
62M
usa;
Gua
delo
upe;
200
3M
W06
3359
.1M
W07
0708
.1M
W07
1027
.1; M
W07
0870
.1; ‒
CPC
3723
8 =
CIRA
D-GL
P 01
63M
usa;
Gua
delo
upe;
200
3M
W06
3360
.1M
W07
0709
.1M
W07
1028
.1; M
W07
0871
.1; ‒
CPC
3723
9 =
CIRA
D-GL
P 01
92M
usa;
Gua
delo
upe;
200
3M
W06
3361
.1M
W07
0710
.1M
W07
1029
.1; M
W07
0872
.1; ‒
CPC
3724
0 =
CIRA
D-GL
P 01
93M
usa;
Gua
delo
upe;
200
3M
W06
3362
.1M
W07
0711
.1M
W07
1030
.1; ‒
; ‒
CPC
3724
1 =
CIRA
D-GL
P 02
22M
usa;
Gua
delo
upe;
200
3M
W06
3363
.1M
W07
0712
.1M
W07
1031
.1; ‒
; ‒
CPC
3724
2 =
CIRA
D-GL
P 02
23M
usa;
Gua
delo
upe;
200
3M
W06
3364
.1M
W07
0713
.1M
W07
1032
.1; M
W07
0873
.1; ‒
CPC
3724
3 =
CIRA
D-GL
P 02
54M
usa;
Gua
delo
upe;
199
2M
W06
3365
.1M
W07
0714
.1M
W07
1033
.1; M
W07
0874
.1; ‒
CPC
3724
4 =
CIRA
D-GL
P 02
55M
usa;
Gua
delo
upe;
199
2M
W06
3366
.1M
W07
0715
.1M
W07
1034
.1; M
W07
0875
.1; ‒
CPC
3724
5 =
CIRA
D-GL
P 02
61M
usa;
Gua
delo
upe;
199
2M
W06
3367
.1M
W07
0716
.1M
W07
1035
.1; M
W07
0876
.1; ‒
CPC
3724
6 =
CIRA
D-GL
P 02
56M
usa;
Gua
delo
upe;
199
2M
W06
3368
.1M
W07
0717
.1M
W07
1036
.1; M
W07
0877
.1; ‒
CPC
3724
7 =
CIRA
D-GL
P 02
57M
usa;
Gua
delo
upe;
199
2M
W06
3369
.1M
W07
0718
.1M
W07
1037
.1; M
W07
0878
.1; ‒
CPC
3724
8 =
CIRA
D-GL
P 02
58M
usa;
Gua
delo
upe;
199
2M
W06
3370
.1M
W07
0719
.1M
W07
1038
.1; M
W07
0879
.1; ‒
CPC
3724
9 =
CIRA
D-GL
P 02
62M
usa;
Gua
delo
upe;
199
2M
W06
3371
.1M
W07
0720
.1M
W07
1039
.1; M
W07
0880
.1; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
7
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
CPC
3725
0 =
CIRA
D-GL
P 02
63M
usa;
Gua
delo
upe;
199
2M
W06
3372
.1M
W07
0721
.1M
W07
1040
.1; M
W07
0881
.1; ‒
CPC
3725
1 =
CIRA
D-GL
P 02
65M
usa;
Gua
delo
upe;
199
2M
W06
3373
.1M
W07
0722
.1M
W07
1041
.1; M
W07
0882
.1; ‒
CPC
3725
2 =
CIRA
D-GL
P 02
66M
usa;
Gua
delo
upe;
199
2M
W06
3374
.1M
W07
0723
.1M
W07
1042
.1; M
W07
0883
.1; ‒
CPC
3725
3 =
CIRA
D-GL
P 02
67M
usa;
Gua
delo
upe;
199
2M
W06
3375
.1M
W07
0724
.1M
W07
1043
.1; M
W07
0884
.1; ‒
CPC
3725
4 =
CIRA
D-GL
P 02
68M
usa;
Gua
delo
upe;
199
2M
W06
3376
.1M
W07
0725
.1M
W07
1044
.1; M
W07
0885
.1; ‒
CPC
3725
5 =
CIRA
D-GL
P 02
69M
usa;
Gua
delo
upe;
199
2M
W06
3377
.1M
W07
0726
.1M
W07
1045
.1; M
W07
0886
.1; ‒
CPC
3725
6 =
CIRA
D-GL
P 02
70M
usa;
Gua
delo
upe;
199
2M
W06
3378
.1M
W07
0727
.1M
W07
1046
.1; M
W07
0887
.1; ‒
CPC
3725
7 =
CIRA
D-GL
P 02
71M
usa;
Gua
delo
upe;
199
2M
W06
3379
.1M
W07
0728
.1M
W07
1047
.1; M
W07
0888
.1; ‒
CPC
3725
8 =
CIRA
D-GL
P 02
73M
usa;
Gua
delo
upe;
199
2M
W06
3380
.1M
W07
0729
.1M
W07
1048
.1; ‒
; ‒
CPC
3725
9 =
CIRA
D-GL
P 02
74M
usa;
Gua
delo
upe;
199
2M
W06
3381
.1M
W07
0730
.1M
W07
1049
.1; M
W07
0889
.1; ‒
CPC
3726
0 =
CIRA
D-GL
P 02
75M
usa;
Gua
delo
upe;
199
2M
W06
3382
.1M
W07
0731
.1M
W07
1050
.1; M
W07
0890
.1; ‒
CPC
3726
1 =
CIRA
D-GL
P 02
76M
usa;
Gua
delo
upe;
199
2M
W06
3383
.1M
W07
0732
.1M
W07
1051
.1; M
W07
0891
.1; ‒
CPC
3726
2 =
CIRA
D-GL
P 02
77M
usa;
Gua
delo
upe;
199
2M
W06
3384
.1M
W07
0733
.1M
W07
1052
.1; ‒
; ‒
CPC
3726
3 =
CIRA
D-GL
P 02
78M
usa;
Gua
delo
upe;
199
2M
W06
3385
.1M
W07
0734
.1M
W07
1053
.1; M
W07
0892
.1; ‒
CPC
3726
4 =
CIRA
D-GL
P 02
79M
usa;
Gua
delo
upe;
199
2M
W06
3386
.1M
W07
0735
.1M
W07
1054
.1; M
W07
0893
.1; ‒
CPC
3726
5 =
CIRA
D-GL
P 02
81M
usa;
Gua
delo
upe;
199
2M
W06
3387
.1M
W07
0736
.1M
W07
1055
.1; M
W07
0894
.1; ‒
CPC
3726
6 =
CIRA
D-GL
P 02
82M
usa;
Gua
delo
upe;
199
2M
W06
3388
.1M
W07
0737
.1M
W07
1056
.1; M
W07
0895
.1; ‒
CPC
3726
7 =
CIRA
D-GL
P 07
73M
usa;
Gua
delo
upe;
199
8M
W06
3389
.1M
W07
0738
.1M
W07
1057
.1; M
W07
0896
.1; ‒
CPC
3726
8 =
CIRA
D-GL
P 07
74M
usa;
Gua
delo
upe;
199
8M
W06
3390
.1M
W07
0739
.1M
W07
1058
.1; M
W07
0897
.1; ‒
CPC
3727
1 =
CIRA
D-ID
N 0
6M
usa;
Indo
nesia
; 198
8M
W06
3391
.1M
W07
0740
.1M
W07
1059
.1; ‒
; ‒
CPC
3727
2 =
CIRA
D-ID
N 0
7M
usa;
Indo
nesia
; 198
8M
W06
3392
.1M
W07
0741
.1M
W07
1060
.1; ‒
; ‒
CPC
3727
4 =
CIRA
D-ID
N 3
0M
usa;
Indo
nesia
; 201
0M
W06
3393
.1M
W07
0742
.1M
W07
1061
.1; M
W07
0898
.1; ‒
CPC
3727
5 =
CIRA
D-ID
N 3
1M
usa;
Indo
nesia
; 201
0M
W06
3394
.1M
W07
0743
.1M
W07
1062
.1; ‒
; ‒
CPC
3727
6 =
CIRA
D-ID
N 5
2M
usa;
Indo
nesia
; 201
0M
W06
3395
.1M
W07
0744
.1M
W07
1063
.1; M
W07
0899
.1; ‒
CPC
3727
7 =
CIRA
D-ID
N 5
3M
usa;
Indo
nesia
; 201
0M
W06
3396
.1M
W07
0745
.1M
W07
1064
.1; M
W07
0900
.1; ‒
CPC
3728
0 =
CIRA
D-JA
M 0
02M
usa;
Jam
aica
; 199
5M
W06
3397
.1M
W07
0746
.1M
W07
1065
.1; ‒
; ‒
CPC
3728
1 =
CIRA
D-JA
M 0
03M
usa;
Jam
aica
; 199
5M
W06
3398
.1M
W07
0747
.1M
W07
1066
.1; M
W07
0901
.1; ‒
CPC
3728
2 =
CIRA
D-LC
A 00
1M
usa;
Sai
nt L
ucia
; 198
6M
W06
3399
.1M
W07
0748
.1M
W07
1067
.1; M
W07
0902
.1; ‒
CPC
3728
3 =
CIRA
D-M
TQ 1
033
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3400
.1M
W07
0749
.1M
W07
1068
.1; M
W07
0903
.1; ‒
CPC
3728
4 =
CIRA
D-M
TQ 1
034
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3401
.1M
W07
0750
.1M
W07
1069
.1; ‒
; ‒
CPC
3728
5 =
CIRA
D-M
TQ 1
063
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3402
.1M
W07
0751
.1M
W07
1070
.1; ‒
; ‒
CPC
3728
6 =
CIRA
D-M
TQ 1
064
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3403
.1M
W07
0752
.1M
W07
1071
.1; M
W07
0904
.1; ‒
CPC
3728
7 =
CIRA
D-M
TQ 1
093
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3404
.1M
W07
0753
.1M
W07
1072
.1; ‒
; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
8
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
CPC
3728
8 =
CIRA
D-M
TQ 1
094
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3405
.1M
W07
0754
.1M
W07
1073
.1; M
W07
0905
.1; ‒
CPC
3728
9 =
CIRA
D-M
TQ 1
123
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3406
.1M
W07
0755
.1M
W07
1074
.1; M
W07
0906
.1; ‒
CPC
3729
0 =
CIRA
D-M
TQ 1
124
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3407
.1M
W07
0756
.1M
W07
1075
.1; ‒
; ‒
CPC
3729
1 =
CIRA
D-M
TQ 1
153
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3408
.1M
W07
0757
.1M
W07
1076
.1; M
W07
0907
.1; ‒
CPC
3729
2 =
CIRA
D-M
TQ 1
154
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3409
.1M
W07
0758
.1M
W07
1077
.1; M
W07
0908
.1; ‒
CPC
3729
3 =
CIRA
D-M
TQ 1
183
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3410
.1M
W07
0759
.1M
W07
1078
.1; M
W07
0909
.1; ‒
CPC
3729
4 =
CIRA
D-M
TQ 1
184
Mus
a; F
ranc
e: M
artin
ique
; 200
3M
W06
3411
.1M
W07
0760
.1M
W07
1079
.1; M
W07
0910
.1; ‒
CPC
3729
5 =
CIRA
D-M
TQ 1
213
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3412
.1M
W07
0761
.1M
W07
1080
.1; ‒
; ‒
CPC
3729
6 =
CIRA
D-M
TQ 1
214
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3413
.1M
W07
0762
.1M
W07
1081
.1; ‒
; ‒
CPC
3729
8 =
CIRA
D-M
TQ 1
221
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3414
.1M
W07
0763
.1M
W07
1082
.1; M
W07
0911
.1; ‒
CPC
3729
9 =
CIRA
D-M
TQ 1
222
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3415
.1M
W07
0764
.1M
W07
1083
.1; M
W07
0912
.1; ‒
CPC
3730
0 =
CIRA
D-M
TQ 1
223
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3416
.1M
W07
0765
.1M
W07
1084
.1; M
W07
0913
.1; ‒
CPC
3730
1 =
CIRA
D-M
TQ 1
225
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3417
.1M
W07
0766
.1M
W07
1085
.1; M
W07
0914
.1; ‒
CPC
3730
2 =
CIRA
D-M
TQ 1
226
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3418
.1M
W07
0767
.1M
W07
1086
.1; M
W07
0915
.1; ‒
CPC
3730
3 =
CIRA
D-M
TQ 1
227
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3419
.1M
W07
0768
.1M
W07
1087
.1; ‒
; ‒
CPC
3731
0 =
CIRA
D-VE
N 0
01M
usa;
Ven
ezue
la; 1
986
MW
0634
20.1
MW
0707
69.1
MW
0710
88.1
; MW
0709
16.1
; ‒
CPC
3731
1 =
CIRA
D-VE
N 0
02M
usa;
Ven
ezue
la; 1
989
MW
0634
21.1
MW
0707
70.1
MW
0710
89.1
; MW
0709
17.1
; ‒
PM11
= A
TCC
3614
3M
usa
AAA;
Hon
dura
s; ‒
AY26
6148
.1‒
‒; ‒
; ‒
X588
Mus
a cv
. Will
iam
s; A
ustr
alia
; ‒EU
5142
68.1
EU51
4326
.1‒;
‒; ‒
X596
Mus
a cv
. SH-
3362
AA;
Aus
tral
ia; ‒
EU51
4270
.1EU
5143
28.1
‒; ‒
; ‒
X602
Mus
a cv
. Lak
atan
; Aus
tral
ia; ‒
EU51
4271
.1EU
5143
29.1
‒; ‒
; ‒
Pseu
doce
rcos
pora
pse
udom
usae
CBS
1471
47 =
CPC
372
70 =
CIR
AD-
IDN
02,
ex-
type
Mus
a; In
done
sia; 1
988
MW
0634
23.1
MW
0707
72.1
MW
0710
91.1
; MW
0709
19.1
; ‒
CBS
1471
48 =
CPC
372
69 =
CIR
AD-
IDN
01
Mus
a; In
done
sia; 1
988
MW
0634
22.1
MW
0707
71.1
MW
0710
90.1
; MW
0709
18.1
; ‒
CBS
1471
49 =
CPC
372
73 =
CIR
AD-
IDN
29
Mus
a; In
done
sia; 1
988
MW
0634
24.1
MW
0707
73.1
MW
0710
92.1
; ‒; ‒
CBS
1471
50 =
CPC
372
78 =
CIR
AD-
IDN
57
Mus
a; In
done
sia; 1
989
MW
0634
25.1
MW
0707
74.1
MW
0710
93.1
; ‒; ‒
CBS
1471
51 =
CPC
372
79 =
CIR
AD-
IDN
58
Mus
a; In
done
sia; 1
989
MW
0634
26.1
MW
0707
75.1
MW
0710
94.1
; MW
0709
20.1
; ‒
Pseu
dosig
moi
dea
exce
ntric
a (O
utgr
oup)
CBS
469.
95 =
INIF
AT C
94/2
02 =
M
UCL
392
27, e
x-ty
peLa
urac
eae
leaf
litte
r; Cu
ba; 1
994
NR_
1565
45.1
KF15
5934
.1KF
1559
75.1
; ‒; K
F156
196.
1
Scol
ecob
asid
ium
aila
nthi
MFL
U 1
8-21
10Ai
lant
hus s
p.; T
haila
nd; 2
017
MK3
4773
1.1
MK4
1289
2.1
‒; ‒
; MK4
1288
1.1
MFL
UCC
17-
0923
, ex-
type
Aila
nthu
s sp.
; Tha
iland
; 201
7N
R_16
3326
.1M
K412
893.
1‒;
‒; M
K412
883.
1
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
9
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
Scol
ecob
asid
ium
bac
illifo
rme
CBS
1004
42Bi
ofilm
on
stai
nles
s ste
el in
drin
king
w
ater
; Ger
man
y; ‒
NR_
1555
66.1
KT27
2051
.1KT
2720
70.1
; ‒; K
T272
059.
1
Scol
ecob
asid
ium
con
stric
tum
CBS
106.
65So
il un
der B
rass
ica
napu
s; G
erm
any;
‒
HQ66
7518
.1HQ
9169
60.1
JF44
0551
.1; ‒
; ‒
Scol
ecob
asid
ium
cor
dana
eCB
S 41
2.51
= M
UCL
947
2 =
QM
37
3b‒;
USA
; ‒HQ
6675
40.1
KF15
5907
.1KF
1559
80.1
; ‒; K
F156
200.
1
Scol
ecob
asid
ium
cor
dana
eCB
S 47
5.80
, ex-
type
Mau
ritia
min
or; C
olom
bia;
197
9N
R_13
2051
.1HQ
9169
76.1
KF15
5981
.1; ‒
; KF1
5619
7.1
Scol
ecob
asid
ium
hum
icol
aCB
S 11
6655
= IM
I 110
131
= UA
MH
1024
1, e
x-ty
pePe
at so
il; C
anad
a; ‒
NR_
1114
37.1
HQ91
6963
.1JF
4405
53.1
; ‒; H
Q87
7631
.1
Scol
ecob
asid
ium
icar
usCB
S 11
6645
Sand
y so
il; C
anad
a; ‒
HQ66
7525
.1LM
6445
99.1
‒; ‒
; LM
6446
04.1
CBS
423.
64 =
MU
CL 1
0160
Rhizo
sphe
re o
f Sol
anum
tube
rosu
m,
in sa
ndy
soil;
Net
herla
nds;
‒HQ
6675
23.1
HQ91
6965
.1KF
1560
08.1
; ‒; ‒
CBS
536.
69 =
MU
CL 1
5054
= O
AC
1021
2, e
x-ty
peFo
rest
soil;
Can
ada;
‒N
R_14
5367
.1KF
1559
44.1
‒; ‒
; KF1
5617
4.1
Scol
ecob
asid
ium
long
ipho
rum
CBS
435.
76 =
UAM
H 39
72So
il un
der a
spha
lt pa
ving
of c
ar
park
; Can
ada;
‒KF
1560
38.1
KF15
5908
.1KF
1559
78.1
; ‒; K
F156
182.
1
Scol
ecob
asid
ium
min
imum
CBS
510.
71 =
ATC
C 22
631
= IM
I 08
2933
, ex-
type
Rhizo
sphe
re o
f Gos
sypi
um
arbo
reum
; Nig
eria
; ‒N
R_14
5366
.1KF
1559
45.1
KF15
6007
.1; ‒
; KF1
5617
2.1
Scol
ecob
asid
ium
mus
aeCB
S 11
9790
= IM
I 138
059
Soil;
Egy
pt; ‒
KT27
2077
.1KT
2720
53.1
KT27
2072
.1; ‒
; KT2
7206
1.1
Scol
ecob
asid
ium
mus
aeCB
S 13
5928
Blac
k bi
ofilm
, was
hing
mac
hine
, de
terg
ent d
raw
er, p
rivat
e re
siden
ce;
Germ
any;
‒
KT27
2080
.1KT
2720
56.1
KT27
2074
.1; ‒
; KT2
7206
4.1
CBS
1359
30Bl
ack
biofi
lm, s
ink
drai
n, p
rivat
e re
siden
ce; G
erm
any;
‒KT
2720
82.1
KT27
2058
.1KT
2720
76.1
; ‒; K
T272
066.
1
CBS
1359
31Bl
ack
biofi
lm, b
atht
ub, w
ater
tap;
Ge
rman
y; ‒
KT27
2081
.1KT
2720
57.1
KT27
2075
.1; ‒
; KT2
7206
5.1
CBS
1450
61 =
CPC
339
47Pe
rsea
am
eric
ana;
Tha
iland
; 200
8M
K442
605.
1M
K442
639.
1M
K442
698.
1; ‒
; ‒
CBS
312.
96De
sert
soil;
Isra
el; 1
996
KT27
2078
.1KT
2720
55.1
KF15
6002
.1; ‒
; KT2
7206
3.1
CBS
729.
95, e
x-ty
pe o
f Och
roco
nis
mira
bilis
Regu
lato
r of d
iver
; ‒; ‒
KF15
6029
.1KF
1559
48.1
KF15
5999
.1; ‒
; KF1
5617
1.1
Scol
ecob
asid
ium
mus
icol
aCB
S 14
4441
= C
PC 3
2927
, ex-
type
Mus
a; M
alay
sia; 2
010
NR_
1603
60.1
‒M
H327
887.
1; M
H327
876.
1; M
H327
898.
1
CPC
3721
8 =
CIRA
D-CM
R 20
64M
usa;
Cam
eroo
n; 1
999
MW
0634
27.1
MW
0707
76.1
MW
0710
95.1
; MW
0709
21.1
; MW
0711
15.1
CPC
3730
8 =
CIRA
D-M
TQ 0
605
Mus
a; F
ranc
e: M
artin
ique
; 199
2M
W06
3428
.1M
W07
0777
.1M
W07
1096
.1; M
W07
0922
.1; M
W07
1116
.1
CPC
3730
9 =
CIRA
D-M
TQ 0
606
Mus
a; F
ranc
e: M
artin
ique
; 199
2M
W06
3429
.1M
W07
0778
.1M
W07
1097
.1; M
W07
0923
.1; M
W07
1117
.1
CPC
3734
8 =
CIRA
D-IN
D 00
8M
usa;
Indi
a; ‒
MW
0634
30.1
MW
0707
79.1
MW
0710
98.1
; ‒; ‒
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
10
Tabl
e 1.
(Con
tinue
d).
Spec
ies
Colle
ction
num
ber(
s)1
Subs
trat
e in
clud
ing
host
, orig
in a
nd
colle
ction
yea
rG
enBa
nk a
cces
sion
num
ber2
ITS
act
tef1
; rpb
2; tu
b2
Scol
ecob
asid
ium
ram
osum
CBS
1371
71 =
FM
R 12
512
= U
THSC
03
-367
7Hu
man
skin
; USA
: Pen
nsyl
vani
a;
2003
LM64
4522
.1LM
6446
01.1
‒; ‒
; LM
6446
06.1
CBS
1371
73 =
FM
R 12
514
= U
THSC
12
-108
2, e
x-ty
peHu
man
nai
l; U
SA: C
alifo
rnia
; 201
2N
R_15
5606
.1LM
6446
03.1
‒; ‒
; LM
6446
08.1
Troc
hoph
ora
simpl
ex (O
utgr
oup)
CBS
1247
44 =
KAC
C 42
362
Daph
inip
hyllu
m m
acro
podu
m;
Kore
a; ‒
GU26
9872
.1GU
3205
68.1
GU38
4580
.1; K
X462
666.
1; ‒
Stra
ins i
denti
fied
base
d on
sequ
ence
sim
ilarit
y
Myc
osph
aere
lla m
usae
CPC
3719
1 =
CIRA
D-AU
S 12
6M
usa;
Aus
tral
ia: N
ew S
outh
Wal
es;
1993
MW
0634
31.1
MW
0707
80.1
MW
0710
99.1
; ‒; ‒
CPC
3719
9 =
CIRA
D-BR
A 00
1M
usa;
Bra
zil; 1
989
MW
0634
32.1
MW
0707
81.1
MW
0711
00.1
; MW
0709
24.1
; ‒
CPC
3721
3 =
CIRA
D-CM
R 28
62M
usa;
Cam
eroo
n; 1
987
MW
0634
33.1
MW
0707
82.1
MW
0711
01.1
; MW
0709
25.1
; ‒
CPC
3721
5 =
CIRA
D-CM
R 28
63M
usa;
Cam
eroo
n; 1
987
MW
0634
34.1
MW
0707
83.1
MW
0711
02.1
; MW
0709
26.1
; ‒
CPC
3730
7 =
CIRA
D-M
TQ 1
241
Mus
a; F
ranc
e: M
artin
ique
; 198
6M
W06
3435
.1M
W07
0784
.1M
W07
1103
.1; M
W07
0927
.1; ‒
CPC
3731
2 =
CIRA
D-CO
K 00
2M
usa;
Coo
k Is
land
s; 1
989
MW
0634
36.1
MW
0707
85.1
MW
0711
04.1
; MW
0709
28.1
; ‒
CPC
3731
3 =
CIRA
D-CO
K 00
3M
usa;
Coo
k Is
land
s; 1
989
MW
0634
37.1
MW
0707
86.1
MW
0711
05.1
; MW
0709
29.1
; ‒
Pant
ospo
ra g
uazu
mae
CPC
3719
5 =
CIRA
D-AU
S 13
0M
usa;
Aus
tral
ia: Q
ueen
sland
; 199
3M
W06
3438
.1M
W07
0787
.1M
W07
1106
.1; M
W07
0930
.1; ‒
Para
palli
doce
rcos
pora
thai
land
ica
CPC
3720
0 =
CIRA
D-BR
A 00
2M
usa;
Bra
zil; 1
988
MW
0634
39.1
MW
0707
88.1
MW
0711
07.1
; MW
0709
31.1
; ‒
CPC
3721
4 =
CIRA
D-CM
R 00
43M
usa;
Cam
eroo
n; ‒
MW
0634
40.1
MW
0707
89.1
MW
0711
08.1
; MW
0709
32.1
; ‒
CPC
3730
5 =
CIRA
D-M
TQ 1
235
Mus
a; F
ranc
e: M
artin
ique
; 198
8M
W06
3441
.1M
W07
0790
.1M
W07
1109
.1; M
W07
0933
.1; ‒
Zasm
idiu
m m
usae
CPC
3730
4 =
CIRA
D-M
TQ 1
228
Mus
a; F
ranc
e: M
artin
ique
; 198
7M
W06
3442
.1M
W07
0791
.1M
W07
1110
.1; ‒
; ‒
CPC
3730
6 =
CIRA
D-M
TQ 1
240
Mus
a; F
ranc
e: M
artin
ique
; 198
6M
W06
3443
.1M
W07
0792
.1M
W07
1111
.1; ‒
; ‒
CPC
3731
4 =
CIRA
D-TO
N 0
07M
usa;
Tong
a; 1
990
MW
0634
44.1
MW
0707
93.1
MW
0711
12.1
; MW
0709
34.1
; ‒
CPC
3732
8 =
CIRA
D-GA
B 03
0M
usa;
Gab
on; 1
998
MW
0634
45.1
MW
0707
94.1
MW
0711
13.1
; MW
0709
35.1
; ‒
CPC
3732
9 =
CIRA
D-GA
B 03
1M
usa;
Gab
on; 1
998
MW
0634
46.1
MW
0707
95.1
MW
0711
14.1
; MW
0709
36.1
; ‒1 A
TCC:
Am
eric
an T
ype
Cultu
re C
olle
ction
, Virg
inia
, USA
; CBS
: Wes
terd
ijk F
unga
l Bio
dive
rsity
Insti
tute
, Utr
echt
, The
Net
herla
nds;
CIR
AD: C
entr
e de
coo
péra
tion
inte
rnati
onal
e en
rech
erch
e ag
rono
miq
ue
pour
le d
ével
oppe
men
t, M
ontp
ellie
r, Fr
ance
; CPC
: Cul
ture
col
lecti
on o
f Ped
ro C
rous
, hou
sed
at C
BS; I
MI:
Inte
rnati
onal
Myc
olog
ical
Insti
tute
, CAB
I-Bio
scie
nce,
Egh
am, B
akeh
am L
ane,
UK;
INIF
AT: A
lexa
nder
Hu
mbo
ldt I
nstit
ute
for B
asic
Res
earc
h in
Tro
pica
l Agr
icul
ture
, Ciu
dad
de L
a Ha
bana
, Cub
a; K
ACC:
Kor
ean
Agric
ultu
ral C
ultu
re C
olle
ction
, Nati
onal
Insti
tute
of A
gric
ultu
ral B
iote
chno
logy
, Rur
al D
evel
opm
ent
Adm
inist
ratio
n, S
uwon
, Rep
ublic
of K
orea
; MFL
UCC
: Mae
Fah
Luan
g U
nive
rsity
Cul
ture
Col
lecti
on, C
hian
g Ra
i, Th
aila
nd; M
UCL
: Uni
vers
ité C
atho
lique
de
Louv
ain,
Louv
ain-
la-N
euve
, Bel
gium
; OAC
: Dep
artm
ent
of B
otan
y an
d Ge
netic
s, U
nive
rsity
of
Guel
ph, O
nt.,
Cana
da; S
TE-U
: Dep
artm
ent
of P
lant
Pat
holo
gy, U
nive
rsity
of
Stel
lenb
osch
, Sou
th A
fric
a; U
AMH:
Uni
vers
ity o
f Al
bert
a M
icro
fung
us C
olle
ction
and
He
rbar
ium
, Edm
onto
n, A
lber
ta, C
anad
a; U
THSC
: Fun
gus T
estin
g La
bora
tory
at t
he U
nive
rsity
of T
exas
Hea
lth S
cien
ce C
ente
r, Sa
n An
toni
o, T
X, U
SA.
2 ITS:
inte
rnal
tran
scrib
ed sp
acer
regi
ons a
nd in
terv
enin
g 5.
8S n
rRN
A ge
ne; a
ct: p
artia
l acti
n ge
ne; t
ef1:
par
tial t
rans
latio
n el
onga
tion
fact
or 1
-alp
ha g
ene;
rpb2
: par
tial D
NA-
dire
cted
RN
A po
lym
eras
e II
seco
nd
larg
est s
ubun
it; tu
b2: p
artia
l bet
a-tu
bulin
gen
e.
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
11
0.05
Trochophora simplex CBS 124744 CPC 37315 Musa Vietnam CPC 37316 Musa Vietnam CPC 37318 Musa Malaysia CPC 37319 Musa Malaysia CPC 37321 Musa Malaysia CPC 37322 Musa Malaysia CPC 37324 Musa Nigeria CPC 37325 Musa Nigeria CPC 37326 Musa Nigeria CPC 37327 Musa Nigeria CPC 37330 Musa Réunion CPC 37331 Musa Réunion CPC 37332 Musa Réunion CPC 37333 Musa Réunion CPC 37334 Musa Réunion CPC 37335 Musa Réunion CPC 37336 Musa Réunion CPC 37337 Musa Réunion CPC 37338 Musa Mauritius CPC 37339 Musa Mauritius CPC 37340 Musa Mauritius CPC 37341 Musa Mauritius CPC 37342 Musa Mauritius CPC 37343 Musa Mauritius CPC 37344 Musa Mauritius CPC 37345 Musa Mauritius CPC 37346 Musa India CPC 37347 Musa India CPC 37349 Musa India CPC 37351 Musa India CPC 37352 Musa India CPC 37353 Musa India CPC 37354 Musa Sri Lanka CPC 37355 Musa Sri Lanka CPC 37356 Musa Sri Lanka CPC 37357 Musa Sri Lanka CPC 37358 Musa Sri Lanka CPC 37359 Musa Sri Lanka CPC 37360 Musa Sri Lanka CPC 37361 Musa Thailand CPC 37362 Musa Thailand CPC 37363 Musa Thailand CPC 37364 Musa Thailand CPC 37365 Musa Thailand CPC 37366 Musa Thailand CPC 37367 Musa Thailand CBS 114824 Musa Réunion CBS 121378 Musa Malaysia CBS 121380 Musa Sri Lanka CIRAD 535 Musa India S1030B Musa Mauritius S1037B Musa Mauritius S17 Musa India S37 Musa cv. Nendran India S48 Musa India CPC 37323 Musa Nigeria CPC 37350 Musa India
Pseudocercospora eumusae
Fig. 1. Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the multigene Pseudocercospora sequence alignment. Bayesian posterior probabilities (PP) > 0.84 are shown at the nodes and thickened lines represent nodes with PP = 1.00. The scale bar represents the expected changes per site. Species are indicated with coloured blocks to the right of the tree. Culture collection numbers are followed by the host and origin, where known. The tree was rooted to Trochophora simplex (culture CBS 124744). The taxonomic novelty described in this study and cultures with type status are indicated in bold face.
lineage sister to Pseudocercospora musae and representing a novel species which is described below. All five species clades were fully supported in the Bayesian analyses (posterior probability value of 1.0). Based on the Scolecobasidium phylogeny (Fig. 2), two strains are very closely related to the ex-type strain of Scolecobasidium musicola while two other
strains form a closely related sister lineage. The two lineages have the following numbers of fixed nucleotide differences: ITS (19 substitutions and one indel), act (13 substitutions; no sequence available for the ex-type), tef1 (23 substitutions) and tub2 (8 substitutions and one indel; no sequence available for CPC 37348).
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
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0.96
0.97
0.97
0.88
0.89
CIRAD 355 Musa Ivory Coast CBS 121362 Musa Taiwan CPC 16301 Musa Mexico
CPC 37216 Musa Cameroon CPC 37217 Musa Cameroon CBS 120258 Musa Cameroon CIRAD 11 Musa Honduras X104 Musa Colombia
CPC 37317 Musa Malaysia CPC 37320 Musa Malaysia CBS 122469 Musa Malaysia CBS 122470 Musa cv. Pisang Byok AAA/AAB Malaysia
CPC 37278 Musa Indonesia CPC 37279 Musa Indonesia CPC 37269 Musa Indonesia CPC 37270 Musa Indonesia CPC 37273 Musa Indonesia CPC 37196 Musa Australia CPC 37197 Musa Brazil CPC 37198 Musa Brazil CPC 37209 Musa Cameroon CPC 37210 Musa Cameroon CPC 37232 Musa Guadeloupe CPC 37233 Musa Guadeloupe CPC 37236 Musa Guadeloupe CPC 37237 Musa Guadeloupe CPC 37239 Musa Guadeloupe CPC 37240 Musa Guadeloupe CPC 37244 Musa Guadeloupe CPC 37245 Musa Guadeloupe CPC 37246 Musa Guadeloupe CPC 37250 Musa Guadeloupe CPC 37253 Musa Guadeloupe CPC 37254 Musa Guadeloupe CPC 37255 Musa Guadeloupe CPC 37257 Musa Guadeloupe CPC 37259 Musa Guadeloupe CPC 37260 Musa Guadeloupe CPC 37262 Musa Guadeloupe CPC 37263 Musa Guadeloupe CPC 37282 Musa Saint Lucia CPC 37283 Musa Martinique CPC 37284 Musa Martinique CPC 37286 Musa Martinique CPC 37288 Musa Martinique CPC 37293 Musa Martinique CBS 143.36 Unknown Suriname CPC 37188 Musa Australia CPC 37189 Musa Australia CPC 37220 Musa Colombia CPC 37221 Musa Colombia CPC 37272 Musa Indonesia CPC 37290 Musa Martinique CPC 37298 Musa Martinique CPC 37300 Musa Martinique CPC 37227 Musa Guinea CPC 37228 Musa Guinea CPC 37238 Musa Guadeloupe CPC 37285 Musa Martinique CPC 37226 Musa Guinea CPC 37229 Musa Guinea CPC 37230 Musa Guinea CPC 37294 Musa Martinique CPC 37301 Musa Martinique CPC 37302 Musa Martinique
0.05
Pseudocercospora fijiensis
Pseudocercospora longispora
Pseudocercospora pseudomusae
Pseudocercospora musae
Fig. 1. (Continued).
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
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0.98
0.96
0.94
CPC 37202 Musa Ivory Coast CPC 37203 Musa Cameroon CPC 37204 Musa Cameroon CPC 37206 Musa Cameroon CPC 37207 Musa Cameroon CPC 37208 Musa Cameroon CPC 37212 Musa Cameroon CPC 37247 Musa Guadeloupe CPC 37271 Musa Indonesia CPC 37274 Musa Indonesia CPC 37275 Musa Indonesia CPC 37190 Musa Australia CPC 37222 Musa Colombia CPC 37201 Musa Ivory Coast CPC 37205 Musa Cameroon CPC 37211 Musa Cameroon CPC 37219 Musa Colombia CPC 37223 Musa Costa Rica CPC 37224 Musa Cuba CPC 37234 Musa Guadeloupe CPC 37235 Musa Guadeloupe CPC 37241 Musa Guadeloupe CPC 37242 Musa Guadeloupe CPC 37243 Musa Guadeloupe CPC 37248 Musa Guadeloupe CPC 37249 Musa Guadeloupe CPC 37251 Musa Guadeloupe CPC 37252 Musa Guadeloupe CPC 37256 Musa Guadeloupe CPC 37258 Musa Guadeloupe CPC 37261 Musa Guadeloupe CPC 37264 Musa Guadeloupe CPC 37265 Musa Guadeloupe CPC 37266 Musa Guadeloupe CPC 37267 Musa Guadeloupe CPC 37268 Musa Guadeloupe CPC 37280 Musa Jamaica CPC 37281 Musa Jamaica CPC 37289 Musa Martinique CPC 37292 Musa Martinique CPC 37296 Musa Martinique CPC 37299 Musa Martinique CPC 37310 Musa Venezuela PM11 Musa AAA Honduras X596 Musa cv. SH-3362 AA Australia X602 Musa cv. Lakatan Australia CPC 37192 Musa Australia CPC 37311 Musa Venezuela CPC 37276 Musa Indonesia CPC 37277 Musa Indonesia CPC 37231 Musa Guadeloupe CPC 37287 Musa Martinique CPC 37303 Musa Martinique CPC 37225 Musa Guinea CPC 37291 Musa Martinique CPC 37295 Musa Martinique CBS 116634 Musa Cuba CPC 37193 Musa Australia X588 Musa cv. Williams Australia CPC 37186 Musa Australia CPC 37187 Musa Australia CPC 37194 Musa Australia
0.05
Pseudocercospora musae(continued)
Fig. 1. (Continued).
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
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0.78
0.99
0.97
0.73
0.75
0.71
Pseudosigmoidea excentrica CBS 469.95 CPC 37218
CPC 37348
CBS 144441 CPC 37308
CPC 37309
CBS 100442
CBS 435.76
CBS 116655 CBS 412.51
CBS 475.80 MFLU 18-2110
MFLUCC 17-0923
CBS 106.65
CBS 510.71 UTHSC 03-3677
UTHSC 12-1082 CBS 116645
CBS 536.69 CBS 423.64
CBS 119790
CBS 145061
CBS 312.96
CBS 729.95, ex-type of Ochroconis mirabilis
CBS 135931
CBS 135928
CBS 135930 0.05
Scolecobasidium musicola
Scolecobasidium bacilliforme
Scolecobasidium longiphorum
Scolecobasidium humicola
Scolecobasidium cordanae
Scolecobasidium ailanthi
Scolecobasidium constrictum
Scolecobasidium minimum
Scolecobasidium ramosum
Scolecobasidium icarus
Scolecobasidium musae
Fig. 2. Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the multigene Scolecobasidium sequence alignment. Bayesian posterior probabilities (PP) > 0.70 are shown at the nodes and thickened lines represent nodes with PP = 1.00. The scale bar represents the expected changes per site. Species are indicated with coloured blocks to the right of the tree. The tree was rooted to Pseudosigmoidea excentrica (= Scolecobasidium excentricum; culture CBS 469.95). The taxonomic novelty described in this study and the cultures with a type status are indicated in bold face.
Table 2. Substitution models and other statistical measures used for Bayesian analyses in this study.
Analysis Number of ingroup sequences
Number of generations
Number of trees used
1Substitution models used for Bayesian analyses; Number of unique site patterns
ITS act rpb2 tef1 tub2
Ochroconis 26 100 000 15 002 GTR+I+G; 462 HKY+G; 186 ‒ HKY+I+G; 332 GTR+I+G; 262
Pseudocercospora 184 6 680 000 100 202 HKY+I; 75 HKY+G; 69 GTR+I; 178 HKY+G; 183 ‒1 ITS: internal transcribed spacer regions and intervening 5.8S nrRNA gene; act: partial actin gene; tef1: partial translation elongation factor 1-alpha gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit; tub2: partial beta-tubulin gene.
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Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
15
Taxonomy
Pseudocercospora pseudomusae Crous & Carlier, sp. nov. MycoBank MB837605. Fig. 3.
Etymology: Name reflects the genetic similarity to Pseudocercospora musae.
Colonies sporulating on PDA. Conidiophores aggregated in dense fascicles forming sporodochia on agar surface; consisting of septate, medium brown, thin-walled, smooth, subcylindrical conidiophores frequently branched below, 20–60 × 3–5 µm. Conidiogenous cells integrated, terminal and intercalary, proliferating sympodially, subcylindrical, smooth, olivaceous, 13–20 × 3–4 µm. Conidia solitary, olivaceous, thin-walled, smooth, subcylindrical, straight to curved, apex obtuse, base truncate, (60–)70–80(–100) × (2.5–)3 µm, (2–)4–5(–6)-septate.
Culture characteristics: Colonies erumpent, spreading, surface folded, with sparse to moderate aerial mycelium and even, lobate margins, reaching 8 mm after 2 wk at 25 °C in the dark. On MEA and PDA surface olivaceous grey, reverse iron-grey. On OA surface olivaceous grey with diffuse yellow pigment in agar (more prominent in isolates CPC 37273, 37378 and 37279).
Typus: Indonesia, on leaves of Musa sp., 1988, J. Carlier (holotype CBS H-24557, culture ex-type CPC 37270 = IDN 02 = CBS 147147).
Additional materials examined: Indonesia, on leaves of Musa sp., 1988, J. Carlier CPC 37269 = IDN 01 = CBS 147148; on leaves of Musa sp., 1988, J. Carlier, 37273 = IDN 29 = CBS 147149; on leaves of Musa sp., 1989, J. Carlier, CPC 37278 = IDN 57 = CBS 147150, CPC 37279 = IDN 58 = CBS 147151.
Notes: Pseudocercospora pseudomusae is closely related to P. musae, which has more obclavate-cylindrical conidia, that are (10–)20–80(–110) × (2–)2.5–5(–6) µm, (0–)2–7(–9)-septate (Braun et al. 2014). Conidia of P. pseudomusae differ in being more subcylindrical (not obclavate), and on average being longer than those of P. pseudomusae.
The five isolates of P. pseudomusae clustered together with full support in the multi-gene phylogeny (Fig. 1). In the individual
gene phylogenies based on distance and parsimony analyses (data not shown), the species can be distinguished from P. musae based on ITS and rpb2 while the distinction is less well-defined for tef1 and the strains are intermingled on act. The ITS sequence of CPC 37270 is 455/468 (97 %, including one indel) to 460/468 (98 %, no indels) similar to the included P. musae ITS sequences. The act sequence of CPC 37270 is 545/550 (99 %, no indels) to 498/501 (99 %, no indels) similar to the included P. musae act sequences. The rpb2 sequence of CPC 37270 is 588/592 (99 %, no indels) to 647/650 (99 %, no indels) similar to the included P. musae rpb2 sequences. The tef1 sequence of CPC 37270 is 448/453 (99 %, no indels) to 449/453 (99 %, no indels) similar to the included P. musae tef1 sequences.
Several of the isolates associated leaf speckle turned out to be representative of the genus Scolecobasidium, which together with its generic synonym, Ochroconis, were recently treated by Shen et al. (2020). All species of Ochroconis for which DNA data are available have since been transferred to Scolecobasidium, except O. ailanthi, which is thus treated below.
Scolecobasidium ailanthi (Jayasiri et al.) Crous, comb. nov. MB837607.Basionym: Ochroconis ailanthi Jayasiri et al., Mycosphere 10: 171. 2019.
Description and illustration: Jayasiri et al. (2019).
List of Cercosporoid taxa associated with leaf spots of Musa
Cercospora apii Fresen. (= Cercospora hayi Calp.)Type: Musa paradisiaca var. sapientum, Cuba (ex-type culture of C. hayi, ATCC 12234); Musa cv. Cavendish, India (CBS H-20035, culture CBS 119395).Disease: Leaf spots.Reference: Groenewald et al. (2013).
Cercospora musae var. paradisiaca Bat. & R. Garnier Type: Musa paradisiaca, Brazil. Disease: Leaf spots.Reference: Braun et al. (2014).[Not known from culture.]
Fig. 3. Pseudocercospora pseudomusae (CPC 37270). A–C. Conidiogenous cells giving rise to conidia (arrows indicate loci). D. Conidia. Scale bars = 10 µm.
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
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Cercospora pingtungensis T.Y. Lin & J.M. YenType: Musa acuminata, M. cavendishii, China, Taiwan. Disease: Leaf spots.Reference: Braun et al. (2014), tentatively reduced to synonymy with Pseudocercospora fijiensis.[Not known from culture.]
Cladocillium musae Chun-Hao Chen & R. KirschnerType: Musa itinerans, Taiwan (TNM, culture BCRC FU30634).Disease: Not associated with distinct leaf spots; secondary coloniser.Reference: Chen et al. (2020).
Metulocladosporiella chiangmaiensis Y. Marín, Cheew. & CrousType: Thailand, Musa sp. (holotype CBS H-23393, culture ex-type CBS 143918 = CPC 18646).Disease: Metulocladosporiella leaf speckle.Reference: Marin-Felix et al. (2019).
Metulocladosporiella malaysiana Y. Marín & CrousType: Malaysia, Musa sp. (holotype CBS H-23394, culture ex-type CBS 143919 = CPC 18131).Disease: Metulocladosporiella leaf speckle.Reference: Marin-Felix et al. (2019).
Metulocladosporiella musae (E.W. Mason) Crous et al. (= Cladosporium musae E.W. Mason)Type: Honduras, Musa sp. (epitype CBS H-14788, CBS 161.74 = ATCC 36973).Disease: Metulocladosporiella leaf speckle.Reference: Marin-Felix et al. (2019).
Metulocladosporiella musicola Crous, Schroers & J.Z. Groenew.Type: Africa, Musa acuminata subgr. Cavendish ‘Grand Nain’ (holotype CBS H-14787, culture ex-type CBS 110960 = CPC 4629).Disease: Metulocladosporiella leaf speckle.Reference: Crous et al. (2006).
Metulocladosporiella musigena Y. Marín, Cheew. & CrousType: Thailand, Musa sp. (holotype CBS H-23395, culture ex-type CBS 143920 = CPC 31490).Disease: Metulocladosporiella leaf speckle.Reference: Marin-Felix et al. (2019).
Metulocladosporiella samutensis Y. Marín, Luangsa-ard & CrousType: Thailand, Musa sp. (holotype CBS H-23396, culture ex-type CBS 143921 = CPC 33939).Disease: Metulocladosporiella leaf speckle. Reference: Marin-Felix et al. (2019).
Mycosphaerella formosana T.Y. Lin & J.M. YenType: Taiwan, Musa sp.Disease: Leaf spots.Reference: Aptroot (2006).[Not known from culture.]
Mycosphaerella henriquesiana G. WinterType: Africa, Musa sp.Disease: Leaf spots.Reference: Aptroot (2006).[Not known from culture.]
Mycosphaerella liukiuensis SawadaType: Taiwan, Musa formosana.Disease: Leaf spots.Reference: Aptroot (2006).[Not known from culture.]
Mycosphaerella musae (Speg.) Syd. & P. Syd. Type: Argentina, Musa sapientum (holotype LPS, slide ex-type IMI 91165).Disease: Leaf speckle.Reference: Aptroot (2006), Arzanlou et al. (2008).[Not known from culture.]
Pseudocercospora assamensis Arzanlou & Crous Type: India, Musa cv. Nanderan (Plantain) (holotype CBS H-20044, culture ex-type X988 = CBS 122467).Disease: Leaf spots.Reference: Arzanlou et al. (2008).
Pseudocercospora eumusae Crous & Mour. (= Mycosphaerella eumusae Crous & Mour.)Type: Réunion (France), Musa sp. (holotype PREM 57315, cultures ex-type (CIRAD 1156, 1157 = CPC 4579, 4580 = CBS 114824, CBS 114825).Disease: Eumusae leaf spot.Reference: Crous & Mourichon (2002).
Pseudocercospora fengshanensis (T.Y. Lin & J.M. Yen) J.M. Yen & S.K. SunType: China, Taiwan, Musa acuminata.Disease: Leaf spots.Reference: Braun et al. (2014).[Not known from culture.]
Pseudocercospora fijiensis (M. Morelet) Deighton (= Myco-sphaerella fijiensis M. Morelet)Type: Cameroon, Musa sp. (epitype CBS H-20037, culture ex-epitype CIRAD 86 = CBS 120258).Disease: Black Sigatoka or black leaf streak disease.Reference: Arzanlou et al. (2008).
Pseudocercospora indonesiana Arzanlou & CrousType: Indonesia, Musa cv. Buai (holotype CBS H-20045, culture ex-type X992 = CBS 122473).Disease: Leaf spots.Reference: Arzanlou et al. (2008).
Pseudocercospora longispora Arzanlou & CrousType: Malaysia, Musa cv. Pisang Byok AAA/AAB (holotype CBS H-20043, culture ex-type X475 = CBS 122470).Disease: Leaf spots.Reference: Arzanlou et al. (2008).
Pseudocercospora musae (Zimm.) Deighton (= Mycosphaerella musicola R. Leach ex J.L. Mulder)Type: Cuba, Musa sp. (epitype CBS H-20038, culture ex-epitype IMI 123823 = CBS 116634).Disease: Yellow Sigatoka disease.Reference: Arzanlou et al. (2008)
© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
17
Pseudocercospora musae-sapienti (A.K. Kar & M. Mandal) U. Braun & Mouch.Type: India, Wallis, Musa paradisiaca.Disease: Leaf spots.Reference: Braun et al. (2014).[Not known from culture.]
Pseudocercospora musicola U. Braun Type: Taiwan, Musa acuminata.Disease: Leaf spots.Reference: Braun et al. (2014).[Not known from culture.]
Pseudocercospora pseudomusae Crous & CarlierType: Indonesia, Musa sp. (holotype CBS H-24557, culture ex-type CPC 37270 = IDN 02 = CBS 147147).Disease: Leaf spots.Reference: Present study.
Rhachisphaerella mozambica (Arzanlou & Crous) Videira & Crous (= Mycosphaerella mozambica Arzanlou & Crous)Type: Mozambique, Musa sp. (holotype CBS H-20039, culture ex-type X34 = CBS 122464).Disease: Leaf spots.Reference: Arzanlou et al. (2008), Videira et al. (2017).
Scolecobasidium musae G.Y. Sun & Lu Hao [= Ochroconis musae (G.Y. Sun & Lu Hao) Samerp. & de Hoog]Type: China, Musa basjoo (holotype HMAS 243664, culture ex-type CGMCC 3.14990 = 0HLHKBJ-22).Disease: Leaf speckle.Reference: Samerpitak et al. (2015).
Scolecobasidium musicola (Crous) Crous, M. Shen & Y. Zhang ter (= Ochroconis musicola Crous)Type: Malaysia, Musa sp. (holotype CBS H-23562, culture ex-type CBS 144441).Disease: Leaf speckle.Reference: Shen et al. (2020).
Uwebraunia musae (Arzanlou & Crous) Crous (= Dissoconium musae Arzanlou & Crous)Type: India, Musa cv. Nendran (Plantain) AAB (holotype CBS H-20036, culture ex-type X1021 = CBS 122453).Disease: Leaf spots.Reference: Arzanlou et al. (2008).
Zasmidium biverticillatum (Arzanlou & Crous) Videira & Crous (= Ramichloridium biverticillatum Arzanlou & Crous)Type: Surinam, Musa sapientum (reference strain CBS 335.36).Disease: Zasmidium leaf speckle.References: Arzanlou et al. (2007), Videira et al. (2017).
Zasmidium ducassei (R.G. Shivas et al.) Y. Marín & Crous (= Ramichloridium ducassei R.G. Shivas et al.)Type: Australia, Musa acuminata × balbisiana (holotype and ex-type culture BRIP 53367).Disease: Zasmidium leaf speckle.References: Shivas et al. (2011), Marin-Felix et al. (2019).
Zasmidium musae (Arzanlou & Crous) Crous & U. Braun (= Stenella musae Arzanlou & Crous)Type: Tonga, Wind Ward Isles, Musa cv. TU8 AAAA, Musa cv. (holotype CBS H-20047, culture ex-type X745 = CBS 122477).Disease: Zasmidium leaf speckle.Reference: Arzanlou et al. (2008).
Zasmidium musae-banksii Videira & Crous (= Ramichloridium australiense Arzanlou & Crous)Type: Australia, Musa banksii (holotype CBS H-19928, culture ex-type CBS 121710).Disease: Zasmidium leaf speckle.Referencse: Arzanlou et al. (2007), Videira et al. (2017).
Zasmidium musicola (Arzanlou & Crous) Crous & U. Braun (= Stenella musicola Arzanlou & Crous)Type: India, Musa cv. Grand Nain AAA (holotype CBS H-20046, culture ex-type X1019 = CBS 122479).Disease: Zasmidium leaf speckle.Reference: Arzanlou et al. (2008).
Zasmidium musigenum Videira & Crous [= Ramichloridium musae (M.B. Ellis) de Hoog]Type: Surinam, Musa sapientum (ex-type culture CBS 365.36 = JCM 6973 = MUCL 9556).Disease: Zasmidium leaf speckle.References: Arzanlou et al. (2007), Videira et al. (2017).
Zasmidium queenslandicum (Arzanlou & Crous) Crous & U. Braun (= Stenella queenslandica Arzanlou & Crous)Type: Australia, Musa banksii (holotype CBS H-20050, culture ex-type CBS 122475).Disease: Zasmidium leaf speckle.Reference: Arzanlou et al. (2008)
DISCUSSION
The Sigatoka leaf spot complex is the most important complex of leaf diseases of banana (Mourichon & Fullerton 1990, Jones 2019). A revision of the taxonomy of this complex by Arzanlou et al. (2008) saw the introduction of eight new species, although their importance as foliar pathogens remains largely unknown. Many isolates that were in the past identified as P. musae based on general symptomatology, are now ascribed to one of these new taxa. The aim of the present study, therefore, was to revisit a global set of 228 isolates identified as P. musae, P. eumusae or other close relatives based on preliminary morphological analysis and ITS sequencing, and resolve their identity by employing multigene DNA sequence analysis and deep morphological observations. Based on these results (Fig. 1), 48 isolates were confirmed as P. eumusae, from India, Malaysia, Mauritius, Nigeria, Réunion (France), Sri Lanka, Thailand and Vietnam, while 104 isolates originally identified as P. musae were confirmed as this taxon from Australia, Brazil, Cameroon, Colombia, Costa Rica, Cuba, Guadeloupe, Guinea, Indonesia, Ivory Coast, Jamaica, Martinique (France), Saint Lucia and Venezuela, and two isolates as P. fijiensis (Cameroon), two isolates as P. longispora (Malaysia) and five isolates as a new species, P. pseudomusae, which is thus far only known from Indonesia. The remainder of the strains studied belonged to
© 2021 Westerdijk Fungal Biodiversity Institute
Crous et al.
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
18
Zasmidium musae (five strains), Scolecobasidium musicola (four strains, Fig. 2), Parapallidocercospora thailandica (three strains), while one strain was identical to Pantospora guazumae and the last strain represented a Penicillium infection and was discarded from further analyses (Table 1). Seven strains were tentatively identified as Mycosphaerella musae (Australia, Brazil, Cameroon, Martinique and the Cook Islands) (Table 1). Although leaf speckle of Musa spp. has traditionally been ascribed to M. musae, this species was originally described from a sexual morph collected on Musa sapientum in Argentina, and its exact identity remains to be determined (Arzanlou et al. 2008).
Other than the Sigatoka disease complex, several other disease complexes have also recently been revised. “Cladosporium leaf speckle” was shown to be caused by several species of Metulocladosporiella (Marin-Felix et al. 2019) and should better be renamed as “Metulocladosporiella leaf speckle”, and “Taiwan leaf speckle” ascribed to a complex of various Zasmidium spp. (Videira et al. 2017), which would in future be better referred to as “Zasmidium leaf speckle”, as these taxa have a wider distribution than Taiwan only. Several other species also causing leaf speckle symptoms have been assigned to Scolecobasidium (Shen et al. 2020), but the majority of foliar pathogens on banana have to date simply been recorded as causing “leaf spots”. A complicating factor is that many of the taxa listed here have not been recorded since they were initially described, and more needs to be done to resolve their phylogeny and ecology.
Conflict of interest: The authors declare that there is no conflict of interest.
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© 2021 Westerdijk Fungal Biodiversity Institute
Pseudocercospora spp. on Musa
Editor-in-ChiefProf. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.E-mail:[email protected]
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