Mycos phere 12(1): 430–518 (2021) www.mycosphere.org
ISSN 2077 7019
Article
Doi 10.5943/mycosphere/12/1/6
Microfungi associated with Camellia sinensis: A case study of leaf and
shoot necrosis on Tea in Fujian, China
Manawasinghe IS 1,2,4, Jayawardena RS 2, Li HL3, Zhou YY1, Zhang W1, Phillips
AJL5, Wanasinghe DN6, Dissanayake AJ 7, Li XH1, Li YH1, Hyde KD2,4 and
Yan JY1*
1
Institute of Plant and Environment Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097,
People’s Republic of China
2
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Tha iland
3
Tea Research Institute, Fujian Academy of Agricultural Sciences, Fu’an 355015, People’s Republic of China
4
Innovative Institute for Plant Health, Zhongkai University of Agriculture and Engineering, Guangzhou 510225,
People’s Republic of China
5
Universidade de Lisboa, Faculdade de Ciências, Biosystems and Integrative Sciences Institute (BioISI), Campo
Grande, 1749–016 Lisbon, Portugal
6
CAS, Key Laboratory for Plant Biodiversity and Biogeography of East Asia (KLPB), Kunming Institute of Botany,
Chinese Academy of Science, Kunming 650201, Yunnan , People’s Republic of China
7
School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu 6 1 1 7 3 1 ,
People’s Republic of China
Manawasinghe IS, Jayawardena RS, Li HL, Zhou YY, Zhang W, Phillips AJL, Wanasinghe DN,
Dissanayake AJ, Li XH, Li YH, Hyde KD, Yan JY 2021 – Microfungi associated with Camellia
sinensis: A case study of leaf and shoot necrosis on Tea in Fujian, China. Mycosphere 12(1), 430–
518, Doi 10.5943/mycosphere/12/1/6
Abstract
Camellia sinensis, commonly known as tea, is one of the most economically important crops
in China. Shoot and leaf necrosis in tea is of considerable concern as it directly affects the quality
and quantity of tea leaf harvest. In the present study, diseased leaves and shoots were collected
from Fujian Province to identify the fungal species associated with the disease. In total 110 strains
were isolated and they were identified by morphological characteristics and multi-locus
phylogenetic approaches. Thirty–two species belonging to 13 genera and 11 families associated
with shoot and leaf necrosis of tea were identified. Five new species; Chaetomium camelliae,
Diaporthe fujianensis, D. fusiformis, D. sinensis and Trichoderma camelliae are introduced. In
addition, nine novel host records are reported. These results indicate high species richness on tea
leaves and shoots. In addition, a checklist for fungi associated with C. sinensis worldwide is
provided. Information presented in this study provides new insights into fungi associated with leaf
necrosis and shoot blight of C. sinensis in China. However, further studies are necessary to
understand the pathogenic potential and biocontrol ability of the species identified in this study.
Keywords – Checklist – Five new species – Nine new host records – Tea pathogens
Table of contents
Ascomycota R.H. Whittaker
Submitted 16 January 2021, Accepted 29 March 2021, Published 4 May 2021
Corresponding Author: Jiye Yan – e-mail – jiyeyan@vip.163.com,
Huiling Li – e-mail – huilingli@163.co m
430
Dothideomycetes O.E. Erikss. & Winka
Dothideomycetidae P.M. Kirk, P.F. Cannon, J.C. David & Stalpers ex C.L. Schoch, Spatafora,
Crous & Shoemaker
Botryosphaeriales C.L. Schoch
Botryosphaeriaceae Theiss
1. Botryosphaeria dothidea (Moug.) Ces. & De Not., in Comm. Soc. crittog. Ital. 1(fasc. 4): 212
(1863)
Pleosporomycetidae C.L. Schoch, Spatafora, Crous & Shoemaker
Pleosporales Luttr. ex M.E. Barr
Didymellaceae Gruyter
2. Epicoccum layuense Qian Chen, Crous & L. Cai, in Chen et al., Stud. Mycol. 87: 145 (2017):
New host record
Phaeosphaeriaceae M.E. Barr
3. Setophoma yingyisheniae F. Liu & L. Cai, in Liu et al., Fungal Systematics and Evolution 4:
54 (2019)
Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan., Maharachch. & K.D. Hyde
Diaporthales Nannf
Diaporthaceae Höhn
4. Diaporthe biguttulata F. Huang, K.D. Hyde & Hong Y. Li, in Huang et al., Fungal Biology
(2015): New host record
5. Diaporthe eucalyptorum Crous & R.G. Shivas., in Crous et al. Persoonia 28: 153 (2012): New
host record
6. Diaporthe fujianensis Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
7. Diaporthe fusiformis Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
8. Diaporthe nobilis Sacc. & Speg., Michelia 1(no. 4): 386 (1878)
9. Diaporthe sackstonii R.G. Shivas, S.M. Thomps. & Y.P. Tan, in Thompson et al., Persoonia
35: 46 (2015)
10. Diaporthe sennae C.M. Tian & Qin Yang, in Yang et al., Phytotaxa 302(2): 149 (2017)
11. Diaporthe sinensis Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
12. Diaporthe unshiuensis F. Huang, K.D. Hyde & Hong Y. Li, in Huang et al., Fungal Biology
119(5): 344 (2015): New host record
13. Diaporthe viniferae Dissanayake, X.H. Li & K.D. Hyde, in Manawasinghe et al., Frontiers in
Microbiology 10: 21 (2019): New host record
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Glomerellales Chadef. ex Réblová, W. Gams & Seifert
Glomerellaceae Locq. ex Seifert & W. Gams
14. Colletotrichum camelliae Massee, in Bull. Misc. Inf., Kew: 91 (1899)
15. Colletotrichum fructicola Prihast., L. Cai & K.D. Hyde, in Prihastuti et al., Fungal Diversity
39: 96 (2009)
Hypocreales Lindau, Natürl. Pflanzenfam
Hypocreaceae De Not
16. Trichoderma atroviride P. Karst., in Bidr. Känn. Finl. Nat. Folk 51: 363 (1892): New host
record
17. Trichoderma camelliae Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
18. Trichoderma lixii (Pat.) P. Chaverri, in Chaverri et al., Mycologia 107(3): 578 (2015): New
host record
431
Nectriaceae Tul. & C. Tul
19. Fusarium asiaticum O’Donnell, T. Aoki, Kistler & Geiser in O’Donnell et al., Fungal Genetics
Biol. 41(6): 619 (2004): New host record
20. Fusarium concentricum Nirenberg & O’Donnell in Nirenberg & O’Donnell Mycologia 90(3):
442 (1998): New host record
21. Fusarium fujikuroi Nirenberg, in Nirenberg Mitt. biol. BundAnst. Ld–u. Forstw. 169: 32
(1976): New host record
22. Fusarium proliferatum (Matsush.) Nirenberg ex Gerlach & Nirenberg, in Nirenberg Mitt. biol.
BundAnst. Ld– u. Forstw. 169: 38 (1982): New host record
Subclass Sordariomycetidae O.E. Erikss., & Winka
Sordariales Chad., ex D. Hawksw. & O.E. Erikss
Chaetomiaceae G. Winter [as ‘Chaetomieae’], Rabenh
23. Chaetomium camelliae Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
Xylariomycetidae O.E. Erikss. & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor & M.E. Barr
24. Arthrinium jiangxiense M. Wang & L. Cai, in Wang, Tan, Liu & Cai, MycoKeys 34(1): 14
(2018)
25. Nigrospora camelliae–sinensis M. Wang & L. Cai, in Wang, Liu, Crous & Cai, Persoonia 39:
127 (2017)
Sporocadaceae Corda
26. Pestalotiopsis camelliae Yan M. Zhang, Maharachch. & K.D. Hyde, in in Zhang et al.,
Sydowia 64(2): 337 (2012)
27. Pestalotiopsis kenyana Maharachch., K.D. Hyde & Crous, in Maharachchikumbura et al.,
Studies in Mycology 79: 166 (2014)
28. Pestalotiopsis lushanensis F. Liu & L. Cai, in Liu et al., Scientific Reports (2017)
29. Pestalotiopsis rhodomyrtus Yu Song, K. Geng, K.D. Hyde & Yong Wang bis, in Song et al.,
Phytotaxa 126(1): 27 (2013)
30. Pseudopestalotiopsis camelliae–sinensis F. Liu & L. Cai, in Liu et al., Scientific Reports 7
(No. 866): 12 (2017)
31. Pseudopestalotiopsis chinensis F. Liu & L. Cai, in Liu et al., Scientific Reports 7 (No. 866): 12
(2017)
Xylariales Nannf
Xylariaceae Tul. & C. Tul
32. Nemania diffusa (Sowerby) Gray: New host record
Introduction
Tea is one of the oldest beverages in the world. The leaves and buds of Camellia sinensis (L.)
Kuntze, either as black tea or green tea, play an important role in traditional cultures especially in
Asia and Europe (Lu et al. 2016). Tea is popular due to its medicinal properties and as a stimulant
(Namita et al. 2012). Camellia comprises over 200 species (Sealy 1958) but C. sinensis is the most
cultivated species of tea. Camellia sinensis is grown in tropical and subtropical climatic regions
(Jayasinghe & Kumar 2019). Camellia sinensis is a perennial plant, belonging to Theaceae
(Meegahakumbura et al. 2016). It requires specific agro–climatic conditions with temperature of
10ºC–30ºC, annual precipitation of >1250 mm, acidic soil, 0.50–10–degree slopes and elevations
up to 2000m (Jayasinghe & Kumar 2019). These factors limit the world’s tea production to certain
countries and regions such as Far East Asia, Africa, Latin America and the Caribbean islands
(Meegahakumbura et al. 2016). Tea is grown as mostly a monocrop in over 52 countries in the
432
world (Lu et al. 2016). China is the largest tea exporting country in the world and has been for
centuries (FAOSTAT 2019). According to the FAOSTAT data, annual tea production in the world
is nearly 2.9 million tons. The world’s black tea production increased by 2.2% annually and green
tea production increased by 7.5% annually during the last decade. China is the world’s largest tea
producer followed by India. In 2016, China accounted for 42.6% of world tea production, with an
output of 2.44 million tonnes (FAOSTAT 2019). The main tea growing regions in China are
Shandong, Jiangsu, Zhejiang, Fujian, Guangdong, Guangxi, Yunnan, Guizhou, Sichuan,
Chongqiong, Shaanxi, Taiwan, Hainan, Tibet, Hubei, Hunan, Henan, Jiangxi, Anhui and Gansu
(Boehm et al. 2016).
Camellia sinensis is affected by a number of diseases caused by bacteria, fungi, insects,
nematodes and viruses. To increase the productivity and quality of tea, it is important to identify the
pathogens associated with different parts of the plant. The most devastating diseases of tea are
caused by fungi (Sarmah et al. 2016, Liu et al. 2019). Microfungi widely and commonly associated
with tea diseases are Colletotrichum spp., Exobasidium vexans (blister blight), Macrophoma
theicola (stem canker and twig dieback), Pellicularia koleroga (black blight, thread blight),
Pestalotiopsis (brown blight), Pseudopestalotiopsis theae (grey blight) and Tunstallia aculeate
(thorny stem blight) (Liu et al. 2017, Yang et al. 2018a, b). In China, over 100 fungal species have
been identified as causal organisms of diseases on buds, leaves and shoots, which are the most
economically important parts of the plant (Jayawardena et al. 2016b, Gao et al. 2016, Liu et al.
2016a, b, Li et al. 2019).
During the last few years there has been a significant improvement in the identification of
new diseases and fungal species from tea plantations in China (Jayawardena et al. 2016b, Li et al.
2019). To develop effective control measures, early detection and correct species identification are
essential. In the present study, we isolated fungi associated diseased leaves and shoots of
Camellia sinensis. The objectives of this study were to (i) identify and characterise the isolates, (ii)
provide detailed descriptions of fungi and (iii) provide a worldwide checklist of fungi associated
with Camellia species. These results will provide new insights into knowledge on microfungi
associated with tea in China.
Materials & Methods
Sampling and isolation
Field surveys were conducted during June 2015 in ten tea plantations in Zhangzhou County,
Fujian Province, China. Samples were collected from diseased leaves and shoots of Purple Rose
cultivar (Fig. 1). Symptomatic tissue samples were taken to the laboratory in zip–lock plastic bags
containing wet sterilised tissues. Samples were photographed and relevant data were documented.
Fungi were isolated by a tissue isolation method. Infected leaves or shoots were cut into small
pieces comprising both disease and healthy tissues. Tissue samples were surface sterilised by
dipping into 70% ethanol for 30 sec and then transferring to 10% NaOCl for 30 sec followed by
three washes with sterilised distilled water. Once the samples were dried on sterilised filter paper,
they were placed on potato dextrose agar (PDA) plates supplemented with ampicillin (100 μg/mL)
and incubated at 25°C. Pure cultures were obtained following 3–4 times of hypal tip isolation.
Cultures were deposited in the Culture Collection of Institute of Plant and Environmental
Protection, Beijing Academy of Agriculture and Forestry Sciences (JZB).
DNA extraction, PCR amplification and sequence assembly
Approximately 10 mg of aerial mycelium was scraped from five to seven days old cultures
grown on PDA. Total genomic DNA was extracted using the DNeasy Plant Mini Kit (QIAGEN
GmbH, QIAGEN Strasse 1, 40742 Hilden, Germany). The PCR mixtures for all gene regions were
as follows: 25 µl total volumes consisted of 0.3 µl of TaKaRa Ex–Taq DNA polymerase, 2.5 µl of
10 × Ex–Taq DNA polymerase buffer, 3.0 µl of dNTPs, 2 µl of genomic DNA, 1 µl of each primer
and 15.2 ddH2 O. Polymerase Chain Reactions (PCR) were conducted in a Bio–Rad C1000 thermal
433
cycler (Germany). The thermal cycler conditions for each locus are given in Table 1. The PCR
products were visualised on a 1% agarose gel stained with ethidium bromide under UV light using
a Gel DocT M XR Molecular Imager (Bio–Rad, USA). Positive amplicons were sequenced by
Beijing Biomed Gene Technology Co LTD. Resulting sequence chromatograms were checked with
BioEdit v.5 (Hall 1999) to confirm sequence quality. At first, the internal transcribed spacer (ITS)
region was sequenced and the resulting sequences were compared with those in GenBank using the
MegaBLAST tool (https://blast.ncbi.nlm.nih.gov/Blast.cgi). Depending on BLAST identification
and morphological characteristics for each isolate, other relevant gene regions were sequenced
(Table 2). Consensus sequences were obtained using DNAStar v. 5.1 (DNASTAR, Inc.).
Figure 1 – Camellia sinensis plantation and field symptoms on Purple Rose cultivar bushes at
different ages. a, b Collection site of the study Zhangzhou, Fujian Province, China. c, d Healthy
young tea buds. e Healthy shrub. f–h Diseased shrubs (sample collected).
434
Table 1 Gene regions and primer pairs used in the present study
GAPDH
Primers
(Forward and
Reverse)
ACT–512F &
ACT–783R
GDF & GDR
ITS
ITS4 & ITS5
LSU
LR0R & LR5
rpb2
fRPB2–5F &
fRPB2–7cR
NS1 & NS4
Locus
ACT
SSU
tef1
tub2
EF1728F &
EF1986R,
Bt2a & Bt2b
BT2Fw &
BT4Rd
PCR amplification
Reference
95°C: 5 min, (95°C: 30 s, 55°C: 50 s,72°C: 1
min) ×39 cycles 72°C: 10 min
95°C 3 min (95°C 1 min 60°C 30 s 72°C 45 s)
×34 cycles 72°C: 10 min
94°C: 3 min, (94°C: 30 s, 58°C: 30 s, 72°C: 1
min) × 32 cycles; 72°C: 10 min
94°C: 5 min, (94°C: 1 min, 53°C: 50 s, 72°C: 1
min) × 37 cycles, 72°C: 10 min
95°C: 5 min, (95°C: 15 s, 56°C: 50 s, 72°C: 2
min) × 37 cycles, 72°C: 10 min
94°C: 4 min, (94°C: 50 s, 56°C: 1 min, 72°C: 1
min,72°C: 10 min) × 37 cycles
(95°C: 5 min, 95°C: 30 s, 58°C: 50 s, 72°C: 1
min) × 40 cycles, 72°C: 10 min
95°C: 5 min, (95°C: 30 s, 58°C: 50 s, 72°C: 1
min) × 40 cycles; 72°C: 10 min
95°C: 5 min, (94°C: 30 s, 55°C: 50 s, 72°C: 1
min) × 40 cycles; 72°C: 7 min
Carbone & Kohn
(1999)
Templeton et al.
(1992)
White et al. (1990)
Vilgalys & Hester
(1990)
Liu et al. (1999)
White et al. (1990)
Carbone & Kohn
(1999)
Glass & Donaldson
(1995)
Woudenberg et al.
(2009)
Phylogenetic analyses
Reference sequences were obtained from GenBank for each genus. The sequences obtained in
this study were aligned with sequences downloaded from GenBank using MAFFT (Katoh & Toh
2010) and manually adjusted using BioEdit v.5 (Hall 1999) wherever necessary. Ambiguous
regions in the alignment were excluded from further analyses, and gaps were treated as missing
data. Phylogenetic relationships were inferred using maximum parsimony (MP) implemented in
PAUP (v4.0) (Swofford & Sullivan 2003), maximum likelihood (ML) in RAxML (Silvestro &
Michalak 2016) and Bayesian posterior probability analysis (BYPP) in MrBayes (v3.0b4)
(Ronquist & Huelsenbeck 2003).
In PAUP, the stability of the trees was evaluated by 1000 bootstrap replications. Branches of
zero length were collapsed and all multiple most parsimonious trees were saved. Parameters,
including tree–length (TL), consistency index (CI), retention index (RI), relative consistency index
(RC) and homoplasy index (HI) were calculated. Differences between the trees inferred under
different optimality criteria were evaluated using Kishino–Hasegawa tests (KHT) (Kishino &
Hasegawa 1989).
The evolutionary models for Bayesian and ML analyses were selected using MrModeltest v.
2.3 (Nylander 2004). The GTR + I + G model of evolution with 1000 non–parametric
bootstrapping iterations was used for the ML analyses. For the BYPP, different evolutionary
models were used in response to the gene regions and gene combinations. The ML analyses were
done with RAxML–HPC2 on XSEDE (8.2.8) (Stamatakis et al. 2008, Stamatakis 2014) in the
CIPRES Science Gateway platform (Miller et al. 2010). For each phytogenetic tree, 1000
nonparametric bootstrapping iterations were used.
In Bayesian posterior probability analysis, posterior probabilities (PPs) were determined by
Markov chain Monte Carlo sampling (BMCMC). Six simultaneous Markov chains were run for 10 6
generations, sampling the trees at every 100th generation. From the 10,000 trees obtained, the first
2,000 representing the burn–in phase were discarded. The remaining 8000 trees were used to
calculate PPs in a majority rule consensus tree (Ronquist & Huelsenbeck 2003).
Taxonomic novelties were submitted to the Faces of Fungi database (Jayasiri et al. 2015) and
Index Fungorum (2020). Sequences generated in this study were deposited in GenBank (Table 2).
Species descriptions, phylogenetic results and notes for these identified taxa are presented under the
435
relevant family and genus. Classes, orders, families and genera were treated according to Wijayawardene et al. (2020).
Table 2 Genbank accession numbers of taxa isolated in the present study
No.
1
Species
B. dothidea
2
3
E. layuense
S. yingyisheniae
4
5
6
D. biguttulata
D. eucalyptorum
D. fuijianensis
7
D. fusiformis
8
9
10
11
D. sackstonii
D. sennae
D. sinensis
12
D. unshiuensis
13
D. viniferae
ID
JZB310190
JZB310191
JZB310192
JZB310193
JZB310194
JZB380035
JZB3270001
JZB3270002
JZB3270003
JZB3270004
JZB320166
JZB320153
JZB320149
JZB320150
JZB320151
JZB320152
JZB320154
JZB320155
JZB320156
JZB320157
JZB320158
JZB320159
JZB320165
JZB320147
JZB320167
JZB320168
JZB320169
JZB320160
JZB320161
JZB320162
JZB320163
JZB320164
JZB320148
ITS
MT497875
MT497876
MT497877
MT497878
MT497879
MT497880
MT523022
MT523023
MT523024
MT523025
MW010210
MW010211
MW010212
MW010213
MW010214
MW010215
MW010216
MW010217
MW010218
MW010219
MW010220
MW010221
MW010222
MW010223
MW010224
MW010225
MW010226
MW010227
MW010228
MW010229
MW010230
MW010231
MW010232
LSU
–
–
–
–
–
MT497881
MT523028
MT523029
MT523030
MT523031
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
tub2
MT513138
MT513139
MT513140
MT513141
MT513142
–
–
–
–
–
MW055998
MW055999
MW056008
MW056009
MW056010
MW056011
MW056012
MW056013
MW056014
MW056015
–
MW056000
–
MW056001
MW056016
MW056017
MW056018
MW056002
MW056003
MW056004
MW056005
MW056006
MW056007
tef1
MT513143
MT513144
MT513145
MT513146
MT513147
–
–
–
–
–
–
MW205223
MW205231
MW205232
–
MW205233
–
–
MW205234
–
–
MW205224
–
MW205225
MW205235
MW205236
MW205237
MW205226
–
MW205227
MW205228
MW205229
MW205230
rpb2
–
–
–
–
–
MT513137
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
SSU
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
ACT
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
CAL
–
–
–
–
–
–
–
–
–
–
MW205204
MW205205
MW205212
MW205213
MW205214
MW205215
MW205216
MW205217
MW205218
MW205219
–
–
–
MW205206
MW205220
MW205221
MW205222
–
MW205207
MW205208
MW205209
MW205210
MW205211
CHS
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
GAPDH
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
436
Table 2 Continued.
No.
14
15
16
17
Species
C. camelliae
C. fructicola
T. atroviride
T. camelliae
18
T. lixii
19
F. asiaticum
20
F. concentricum
21
F. fijikuroi
22
23
F. proliferatum
Ch. camelliae
24
25
26
A. jiangxiense
Ni. camelliae–
sinensis
P. camelliae
27
P. kenyana
28
29
P. lushanensis
P. rhodomyrtus
ID
JZB330153
JZB330154
JZB3360001
JZB3360002
JZB3360003
JZB3360004
JZB3360005
JZB3360006
JZB3360007
JZB3360008
JZB3110018
JZB3110019
JZB3110020
JZB3110021
JZB3110022
JZB3110010
JZB3110011
JZB3110012
JZB3110013
JZB3110014
JZB3110016
JZB3110017
JZB3110015
JZB3340001
JZB3340002
JZB3260001
JZB3230016
ITS
MW007830
MW007831
MW008450
MW008451
MW008452
MW008453
MW008454
MW008455
MW008456
MW008457
–
–
–
–
–
–
–
–
–
–
–
–
–
MT535751
MT535752
MT525316
MT525317
LSU
–
–
–
–
–
–
–
–
–
tub2
MW013330
MW013331
–
–
–
–
–
–
–
tef1
MW056027
MW056028
MW056029
MW056030
MW056031
MW056019
MW056020
MW056021
MW056022
MW056023
MW056025
MW056026
MW056024
MT535535
MT535536
MW026028
MW026029
rpb2
–
–
–
–
–
–
–
–
–
–
MW055992
MW055993
MW055994
MW055995
MW055996
MW055984
MW055985
MW055986
MW055987
MW055988
MW055990
MW055991
MW055989
MT535537
MT535538
–
–
SSU
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
ACT
MW013328
MW013329
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
CAL
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
CHS
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
GAPDH
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
MT535749
MT535750
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
MT535533
MT535534
MW034378
MW034379
JZB340064
JZB340063
JZB340062
JZB340061
JZB340059
JZBH340060
MT509821
MT509822
MT509823
MT509824
MT509825
MT509826
–
–
–
–
–
–
MT535513
MT535514
MT535515
MT535516
MT535517
MT535518
MW205238
MW205239
MW205240
MW205241
MW205242
MW205243
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
437
Table 2 Continued.
tub2
tef1
rpb2
ID
ITS
LSU
SSU
ACT
CAL
CHS GAPDH
JZB340040
MT509827
–
MT535519
MW034366
–
–
–
–
–
–
JZB340041
MT509828
–
MT535520
MW034367
–
–
–
–
–
–
JZB340042
MT509829
–
–
MW034368
–
–
–
–
–
–
JZB340043
MT509830
–
–
–
–
–
–
–
–
–
JZB340044
MT509831
–
MT535521
–
–
–
–
–
–
–
JZB340045
MT509832
–
MT535522
–
–
–
–
–
–
–
JZB340046
MT509833
–
MT535523
–
–
–
–
–
–
–
JZB340047
MT509834
–
MT535524
MW034369
–
–
–
–
–
–
JZB340048
MT509835
–
–
–
–
–
–
–
–
–
JZB340049
MT509836
–
MT535525
MW034370
–
–
–
–
–
–
JZB340050
MT509837
–
MT535526
MW034371
–
–
–
–
–
–
JZB340051
MT509838
–
MT535527
MW034372
–
–
–
–
–
–
JZB340052
MT509839
–
MT535528
–
–
–
–
–
–
–
JZB340053
MT509840
–
–
–
–
–
–
–
–
–
JZB340054
MT509841
–
MT535529
MW034373
–
–
–
–
–
–
31
Ps. chinensis
JZB340055
MT509842
–
MT535530
MW034374
–
–
–
–
–
–
JZB340056
MT509843
–
MT535531
MW034375
–
–
–
–
–
–
JZB340057
MT509844
–
–
MW034376
–
–
–
–
–
–
JZB340058
MT509845
–
MT535532
MW034377
–
–
–
–
–
–
32
Nemania diffusa JZB3370001
MT509575
–
–
–
MT512899
–
–
–
–
–
JZB3370002
MT509576
–
–
–
MT512900
–
–
–
–
–
JZB3370003
MT509577
–
–
–
MT512901
–
–
–
–
–
Ex–type cultures are bold. ITS: internal transcribed spacer regions 1 & 2 including 5.8S nrDNA gene; LSU: Large subunit nrDNA gene; tef1: Partial translation elongation
factor 1–α; tub2: partial sequences of beta–tubulin; rpb2: RNA polymerase II gene; SSU: small subunit nrDNA gene; ACT: Partial actine; CAL: calmodulin; CHS:
Chalcone synthase; GAPDH: Glyceraldehyde 3–phosphate dehydrogenase. (JZB: Culture Collection of Institute of Plant and Environmental Protection, Beijing Academy
of Agriculture and Forestry Sciences. Type sequnces of newly generated taxa are bold.
No.
30
Species
Ps. camelliae–
sinensis
Morphology and culture characteristics
Colony morphology and conidial characteristics were examined for each species isolated. Colony colours were recorded according to the colour
charts of Rayner (1970) after five to seven days of growth on PDA at 25°C. Digital images of morphological structures mounted in water were taken
using an Axio Imager Z2 photographic microscope (Carl Zeiss Microscopy, Oberkochen, Germany). Measurements were taken using ZEN PRO 2012
(Carl Zeiss Microscopy). For each isolate, conidial length and width were measured for 40 conidia and the mean values were calculated. In addition,
conidial shape, colour and guttulation were recorded.
438
Checklist
The checklist was based on articles in refereed journals, Index to Saccardo’s Sylloge
Fungorum, Petrak’s Lists, Index of Fungi, graduate student theses, books and web–based resources
such as annual reports on tea and the USDA fungal database Fungus–Host Distributions database
(https://nt.ars–grin.gov/fungaldatabases/fungushost/fungushost.cfm) (Accessed 10th June 2020).
The mode of life, i.e. pathogen, endophyte or saprotroph, is listed. The checklist includes species
names, family, life modes, disease name (if any), locality and references. The current name used is
according to Index Fungorum (2020) and the classification follows Wijayawardene et al. (2020).
Genera and species are listed in alphabetical order. In some cases, the host names given in the
original citation were changed to be consistent with current taxonomy. In a few cases, neither the
species cited nor a proper synonym was identified and the species name was used as originally
cited.
Results
In this study, we identified 32 species belong to 11 fungal families. Species descriptions,
phylogenetic results and notes for these identified taxa are presented under the relevant family and
genus. Classes, orders, families and genera were treated according to Wijayawardene et al. (2020).
Dothideomycetes P.M. Kirk, P.F. Cannon, J.C. David & Stalpers ex C.L. Schoch, Spatafora, Crous
& Shoemaker, Mycologia 98 (6): 1045 (2007)
For taxonomic treatments, we follow Hongsanan et al. (2020a, b).
Botryosphaeriales C.L. Schoch, Crous & Shoemaker, Mycologia 98 (6): 1050 (2007)
Notes –
Six families; Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae,
Phyllostictaceae, Planistromellaceae and Saccharataceae are accepted in Botryosphaeriales.
Taxonomic treatments follow Phillips et al. (2019) and Hongsanan et al. (2020b).
Botryosphaeriaceae Theiss. & Syd [as ‘Botryosphaeriacae’], Annls mycol. 16(1/2): 16 (1918)
Notes – Botryosphaeriaceae species are endophytes, pathogens and saprobes on a wide range
of hosts (Manawasinghe et al. 2016, Rashmi et al. 2019). They are well–known opportunists on
many economically important crops (Chethana et al. 2016). Currently, more than 279 species and
22 genera are included with this family (Dissanayake et al. 2016, Phillips et al. 2013, 2019,
Hongsanan et al. 2020b).
Botryosphaeria Ces. & De Not. Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 211 (1863)
Botryosphaeria comprises 13 species based on both morphology and molecular data
(Dissanayake et al. 2016, Jayawardena et al. 2019). In the present study, five isolates clustered in
the main clade with B. dothidea type sequence. (ML and BYPP) (Fig. 2). Depending on
morphology and sequence similarites we confirmed these five strains as B. dothidea.
Botryosphaeria dothidea (Moug.: Fr.) Ces. & De Not., Comm. Soc. crittog. Ital. 1 (fasc. 4): 212
(1863)
Fig. 3
Index Fungorum: IF 183247; Facesoffungi number: FoF03512
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: Not observed.
Asexual morph: Conidiomata stromatic, Conidiophores hyaline, cylindrical, smooth.
Conidiogenous cells 11.5–14 × 4–6.5 μm (x = 13 × 6 m, n = 20), hyaline, sub–cylindrical.
Conidia 18–40 × 5–10 μm (x̅ = 24 × 7 μm, n = 20), hyaline, unicellular, narrowly fusiform, with a
sub–truncate to bluntly rounded base, forming a septum before germination, smooth–walled with
granular contents.
Culture characteristics – Colonies on PDA reaching 50 mm diam., after four days at 28°C.
Initially, white becoming grey, moderately dense, margin smooth, olivaceous.
439
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead leaves and shoots
of Camellia sinensis, June 2015, H.L. Li (dried cultures JZBH310190–JZBH310194), and living
cultures JZB310190–JZB31094.
Notes – The colony morphology of taxa isolated in this study are similar to typical strains of
B. dothidea (Phillips et al. 2013). In the multilocus phylogenetic analysis, the five isolates from the
present study clustered together with 56% ML bootstrap and less than 0.90 BYPP. Morphologically
these taxa are similar to the type description of B. dothidea (Phillips et al. 2013). Botryosphaeria
dothidea has a wide range of hosts and it is a well–known woody host–pathogen (Phillips et al.
2005, 2013, Dissanayake et al. 2016, Hyde et al. 2020a). Botryosphaeria dothidea had been
reported to cause diseases on many different hosts in China (Manawasinghe et al. 2018). This
species was first reported as shoot blight pathogen in Chinese tea plants in 2016 (Jayawardena et al.
2016b).
Figure 2 – The phylogenetic tree generated by ML analysis of combined ITS and translation
elongation factor 1–alpha (tef1) sequence data of Botryosphaeria species. The phylogenetic tree is
rooted with Neofusicoccum parvum (ATCC 58191). Tree topology of the ML analysis was similar
440
to the BYPP. The best scoring RAxML tree with a final likelihood value of –24349.980578 is
presented. The matrix had 1172 distinct alignment patterns, with 9.91% of undetermined characters
or gaps. Estimated base frequencies were as follows: A = 0.251668, C = 0.245757, G = 0.259668, T
= 0.242908; substitution rates AC = 1.353890, AG = 4.605576, AT = 1.059439, CG = 0.801610,
CT = 9.121730, GT = 1.000000; gamma distribution shape parameter α = 0.944898. RAxML
bootstrap support values ≥50% and Bayesian posterior probabilities ≥0.95 (BYPP) are given near
the nodes. The scale bar indicates 0.02 changes per site. Ex–type/ex–epitype strains are in bold and
isolates belong to this study are given in red.
Figure 3 – Botryosphaeria dothidea (JZB310193) a Material examined. b Upper view of the
colony on PDA after four days. c Reverse view of the colony on PDA after four days. d Mycelia.
e–i Conidia. Scale bars: e–i = 20 µm.
Pleosporales Luttr. ex M.E. Barr, Prodromus to class Loculoascomycetes: 67 (1987)
Notes – Pleosporales is the largest order of Dothideomycetes (Liu et al. 2017). It comprises
highly diverse taxa that are endophytes, epiphytes, parasites, lichenicolous, or saprobes in terrestrial
or aquatic environments or even occur on animal dung (Zhang et al. 2009). For the taxonomic
treatment of Pleosporales, we follow Kirk et al. (2008), Zhang et al. (2009) and Hongsanan et al.
(2020a).
Didymellaceae Gruyter, Aveskamp & Verkley, Mycol. Res. 113(4): 516 (2009)
Notes – Zhang et al. (2009) included Didymellaceae in Pleosporales within the suborder
Pleosporineae. The family Didymellaceae was established by de Gruyter et al. (2009).
Didymellaceae includes economically important plant pathogens (Salam et al. 2011, de Gruyter et
al. 2013) endophytes, as well as fungicolous and lichenicolous taxa (Aveskamp et al. 2010, Chen et
al. 2015, Valenzuela-Lopez et al. 2018). Recent taxonomic treatments are given in Wanasinghe et
al. (2018), Marin-Felix et al. (2019) and Hongsanan et al. (2020a).
Epicoccum Link., Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816) [1815]
Notes – Epicoccum is characterized by epicoccoid and sub–cylindrical conidia (Chen et al.
2015). Species belonging to this genus are ubiquitous (Chen et al. 2015). They have been reported
441
as common causal agents of leaf spot diseases in various hosts (Chen et al. 2015, Liu et al. 2019).
The taxon isolated in the present study formed a clade together with Epicoccum layuense with 64%
ML and 77% MP bootstrap values in the phylogenetic tree (Fig. 4).
Figure 4 – Phylogenetic tree generated by ML analysis of combined LSU, ITS, and rpb2 sequence
data of Epicoccum species. The tree is rooted with Allophoma cylindrispora (CBS 142453). Tree
topology of the ML analysis was similar to the MP. The best scoring RAxML tree with a final
likelihood value of –4626.221244 is presented. The matrix had 298 distinct alignment patterns,
with 1.09% of undetermined characters or gaps. Estimated base frequencies were as follows: A =
0.252747, C = 0.210997, G = 0.301466, T = 0.234790; substitution rates AC = 1.405315, AG =
3.684928, AT = 2.423691, CG = 1.931133, CT = 14.809240, GT = 1.000000; gamma distribution
shape parameter α = 0.269921. Maximum parsimony analysis of 1737 constant characters and 221
informative characters resulted in 738 equally most parsimonious tree of 462 steps (CI = 0.564, RI
= 0.672, RC = 0.379, HI = 0.436). RAxML bootstrap support values ≥50% and MP bootstrap
442
support values ≥50% are shown near the nodes. The scale bar indicates 20.0 changes per site. Ex–type/
ex–epitype strains are in bold and taxon isolated in this study is in red.
Epicoccum layuense Qian Chen, Crous & L. Cai, in Chen et al., Stud. Mycol. 87: 145 (2017)
Fig. 5
Index Fungorum: IF818963; Facesoffungi number: FoF09381
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Not observed. Asexual
morph: Hyphae about 2.5 μm, septate, branched, conidiomata on PDA, aggregated, superficial,
clavate, Conidiomatal wall pseudoparenchymatous, multi–layered, outer wall brown olivaceous.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells, light brown, simple. Conidia
12–13 × 20–18 μm (x̅ = 12 × 20 μm n = 30) μm diam, globose to subglobose, one basal cell,
terminal, solitary, smooth, dark brown.
Culture characteristics – Colonies on PDA, 60 mm diam., after seven days, margin irregular,
aerial mycelia floccose, bright yellow; reverse yellow to pale brown, with a brown concentric ring
near the centre.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead leaves of
Camellia sinensis, June 2015, H.L. Li (dried culture JZBH380035), and living cultures JZB380035.
Notes – The phylogenetic analysis of combined LSU, ITS and rpb2 DNA data set placed this
taxon with the Epicoccum layuense with 67% and 77% bootstrap support values. Epicoccum
layuense (JZB380035) shares similar colony morphology and spore characters with the type
description of Epicoccum layuense (Chen et al. 2015). A recent study conducted by Del Frari et al.
(2019) has shown the potential of Epicoccum layuense as a biocontrol agent against grapevine
trunk disease caused by Phaeomoniella chlamydospora, Fomitiporia mediterranea, and
Phaeoacremonium minimum. This is the first report of E. layuense on C. sinensis (Farr & Rossman
2020).
Figure 5 – Epicoccum layuense (JZB380035). a conidiomata on PDA. b Developing spore.
c Conidium. d Upper view of a colony on PDA after seven days. e Reverse view of the colony on
PDA after seven days. Scale bars: a = 100 µm, b–c = 10 μm.
Phaeosphaeriaceae M.E. Barr, Mycologia 71(5): 948 (1979)
Notes – Phaeosphaeriaceae consists of economically important plant pathogens
(Phookamsak et al. 2014), endophytes or saprobes on plants. Phaeosphaeriaceae has undergone
several revisions and species additions during the last years (Phookamsak et al. 2014). For the
taxonomic treatment of Phaeosphaeriaceae, we follow Hongsanan et al. (2020a).
Setophoma Gruyter, Aveskamp & Verkley, in de Gruyter, Woudenberg, Aveskamp, Verkley,
Groenewald & Crous, Mycologia 102(5): 1077 (2010)
Notes – Setophoma was introduced by de Gruyter et al. (2010) and is typified by S. terrestris
(= Phoma terrestris). Setophoma species are characterised by setose pycnidia, phialidic
conidiogenous cells and hyaline, ellipsoidal to subcylindrical, aseptate conidia (de Gruyter et al.
2010, Quaedvlieg et al. 2013). Species belonging to this genus are well–known pathogens on
443
economically important crops including tea (Liu et al. 2019). Four isolates belonging to Setophoma
were isolated and identified here (Fig. 6).
Figure 6 – Phylogenetic tree generated by ML analysis of combined LSU and ITS sequence data of
Setophoma species. The tree is rooted with Didymella pinodella (CBS 531.66). Tree topology of
the ML analysis was similar to the MP. The best scoring RAxML tree with a final likelihood value
of – 4077.859174 is presented. The matrix had 232 distinct alignment patterns, with 8.92% of
undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.242298, C =
0.217889, G = 0.274550, T = 0.265263; substitution rates AC = 1.307115, AG = 3.702431, AT =
3.436097, CG = 0.690224, CT = 8.860042, GT = 1.000000; gamma distribution shape parameter α
= 0 0.692111. Maximum parsimony analysis of 1111 constant characters and 173 informative
characters resulted in 62 equally most parsimonious tree of 462 steps (CI = 0.727, RI = 0.836, RC =
0.608, HI = 0.273). RAxML bootstrap support values ≥50% and MP bootstrap support values
≥50% are shown near the nodes. The scale bar indicates 0.03 changes per site. Ex–type/ex–epitype
strains are in bold. Isolates from this study are in red.
444
Setophoma yingyisheniae F. Liu & L. Cai, in Liu, Wang, Li, Wang & Cai, Fungal Systematics and
Evolution 4: 54 (2019)
Fig. 7
Index Fungorum: IF829903; Facesoffungi number: FoF09382
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: not observed. Asexual
morph: Conidiomata 100–200 μm, pycnidial, black, globose or subglobose. Pycnidial wall brown,
with 3–5 layers, walls pseudoparenchymatous. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells hyaline, smooth, ampulliform, aseptate. Conidia 3.5–4.5 × 2–3 μm (x̅ = 4 ×
2.5 μm, n = 40) hyaline, aseptate, ellipsoid.
Culture characteristics – Grows up to 50 mm diam., after five days on PDA, irregular,
filamentous margins, flat, superficial, grey and wrinkled with a white margin. Reverse black with a
white margin.
Material examined – CHINA, Fujian Province, Zhangzhou County, on deseased leaves of
Camellia sinensis, June 2015, H.L. Li (dried cultures JZBH3270001–4), and living cultures
JZB3270001–4.
Notes – In the BLAST results, four isolates obtained in the present study showed similarities
to the species in Phaeosphaeriaceae. A phylogenetic analysis was conducted using combined LSU
and ITS gene regions for Phaeosphaeriaceae. In the phylogenetic tree of Setophoma, isolates from
this study grouped with the ex–type strain of Setophoma yingyisheniae (CGMCC 3.195.27).
Morphologically, isolates in this study were similar to the original description of S. yunnanensis
(Liu et al. 2019) and all the sequences generated in this study were similar to the Setophoma
yingyisheniae (CGMCC 3.195.27). Setophoma yingyisheniae was introduced by Liu et al. (2019) as
a species associated with leaf spots of tea plants in Yunnan province. This is a new geographical
report for S. yunnanensis.
Figure 7 – Setophoma yunnanensis (JZB3270002). a Diseased leaf. b Upper view of a colony on
PDA after five days. c Reverse view of the colony on PDA after five days. d, e Conidia. Scale bar:
d, e = 10 µm.
Sordariomycetes
Notes – For the taxonomic treatments of Sordariomycetes we follow Hyde et al. (2020b).
Subclass Diaporthomycetidae Senan., Maharachch. & K.D. Hyde, in Maharachchikumbura et al.,
Fungal Diversity 72: 208 (2015)
Notes – Maharachchikumbura et al. (2016) introduced Diaporthomycetidae based on
combined analysis of LSU, small subunit ribosomal RNA gene (SSU), tef1 and rpb2 sequence data.
For the taxonomic treatment of Diaporthomycetidae we follow Hyde et al. (2020b).
Diaporthales Nannf., Nova Acta Regiae Societatis Scientiarum Upsaliensis 8 (2): 53 (1932)
Notes – Based on morphology and molecular data, currently 27 families and 138 genera are
accepted within Diaporthales (Senanayake et al. 2017, Hyde et al. 2020b).
445
Diaporthe Nitschke Pyrenomyc. Germ. 2: 240 (1870)
Notes – Species in this genus are well known pathogens on many hosts including
economically important plants (Hyde et al. 2014, Udayanga et al. 2014a, b, 2015, Dissanayake et
al. 2017a). Diaporthe species are cryptic species, therefore the modern taxonomic classification and
identification are based on molecular phylogeny. Hence in this study, latest classification as
proposed by Marin-Felix et al. (2019), Manawasinghe et al. (2019), Hyde et al. (2019) was
followed.
In the present study, 45 isolates were obtained from tea leaves and shoots. However, only 23
isolates were used in the phylogenetic analysis due to sequencing errors and to obtain better
resolved phylogenies. A preliminary analysis was conducted using ITS, tef1, β–tubulin (tub2),
calmodulin (cal) and partial histone (his) gene regions with 250 Diaporthe species (including ex–
type strains) and tree was rooted with Diaportherella corylina. Once the placements of the species
were confirmed, the final phylogenetic tree was arranged including only the taxa from respective
species complex. (Fig. 8).
In the phylogenetic analysis, nine Diaporthe isolates clustered together with the Diaporthe
eucalyptoum (CBS132525), D. lithocarpus (CGMCC 3.15175) and D. hongkongensis
(CBS115448). In this clade, branch lengths and divergence times were indistinguishable. Therefore,
we conducted a recombination test for delimitation of species. In this analysis, we included four
strains comprising ex–type strains of D. lithocarpus (CGMCC3.15175), D. eucalyptorum
(CBS132525), D. fujianensis (JZBH3340150) and D. fusiformis (JZBH3340154). The pairwise
homoplasy index (PHI) test results using both LogDet transformation and splits decomposition
revealed that the PHI test did not find statistically significant evidence for recombination (p = 1.0)
(Fig. 9). Therefore, the two species identified in this study were treated as novel taxa.
Diaporthe biguttulata F. Huang, K.D. Hyde & Hong Y. Li, in Huang et al., Fungal Biology (2015)
Fig. 10
Index Fungorum: IF810579; Facesoffungi number: FoF09383
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Not observed. Asexual
morph: Pycnidia in culture, black, erumpent; walls 3–6 layers, light brown textura angularis.
Paraphyses not observed. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 10–
30 × 2–3 µm, phialidic, cylindrical, terminal and lateral. Alpha conidia 5.5–8 × 2–3.5 µm (x̅ = 6 ×
2.5 µm, n = 30), aseptate, hyaline, smooth, guttulate, fusoid, tapering towards both ends, apex
subobtuse, base subtruncate. Beta conidia and gamma conidia not seen.
Culture characteristics – Colonies on PDA covers entire petri dish after10 days at 25ºC.
Abundant tufted white aerial mycelia, buff, numerous black pycnidia 0.5 mm in diam., typically in
the direction of the edge of the colony. Reverse buff with concentric lines.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH320166), and living culture JZB320166
Notes – A toxon isolated in the present study formed a well–supported cluster with the
Diaporthe biguttulata (ZJUD47) with 100% ML, 99%, MP and 1.00 BYPP values. The
morphological characteristics of the isolated taxa are similar to the ex–type description (Huang et
al. 2015). Diaporthe biguttulata was introduced from Citrus limon in China (Huang et al. 2015).
This species has also been reported on Juglans regia in China. This is the first report of D.
biguttulata on Camellia sinensis (Farr & Rossman 2020).
Diaporthe eucalyptorum Crous & R.G. Shivas., in Crous et al. Persoonia 28: 153 (2012)
Fig. 11
Index Fungorum: IF 800374; Facesoffungi number: FoF09077
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Not observed. Asexual
morph: Pycnidia black, erumpent, cream conidial droplets exuding from central ostioles. Pycnidial
wall consisting of 3–6 layers of hyaline outer layers and light brown inner layers, textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, cylindrical, terminal
and lateral, with a slight taper towards the apex. Paraphyses hyaline, smooth, cylindrical, 1–3
446
septa. Alpha conidia 5.5–7 × 2–3 µm (x̅ = 6 × 2.5 µm, n = 30), aseptate, hyaline, smooth, guttulate,
fusoid, tapering towards both ends, straight, apex subobtuse, base subtruncate, Beta and gamma
conidia not seen.
Culture characteristics – Colonies on PDA reach 90 mm diam., after 10 days at 25ºC (covers
the total surface), abundant tufted white aerial mycelia, buff, numerous black pycnidia 0.5 mm in
diam. occur in the mycelium, typically in the direction of the edge of the colony; reverse buff with
concentric lines.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH320153), and living culture JZB320153.
Figure 8 – Phylogenetic tree generated by ML analysis of combined ITS, tef1, tub2, Cal and HIS
sequence data of Diaporthe species. The analyses included 166 strains and the tree was rooted with
Diaporthella corylina (CBS 121124). Tree topology of the ML analysis was similar to the MP and
447
BYPP. The best scoring RAxML tree with a final likelihood value of – 24349.980578 is presented.
The matrix had 1172 distinct alignment patterns, with 9.91% of undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.251668, C = 0.245757, G = 0.259668, T =
0.242908; substitution rates AC = 1.353890, AG = 4.605576, AT = 1.059439, CG = 0.801610, CT
= 9.121730, GT = 1.000000; gamma distribution shape parameter α = 0.944898. Maximum
parsimony analysis of 1145 constant characters and 1168 informative characters resulted in 100
equally most parsimonious tree of 1000 steps (CI = 0.325, RI = 0.757, RC = 0.246, HI = 0.675).
RAxML bootstrap support values ≥50% and maximum parsimony bootstrap support values ≥50%
are shown near the nodes. Nodes with ≥0.95 (BYPP) Bayesian posterior probabilities are indicated
with thickened lines. The scale bar indicates 0.1 changes per site. Ex–type/ ex–epitype strains are in
bold. New isolates recovered in this are in red.
Figure 8 – Continued.
448
Figure 8 – Continued.
449
Figure 9 – Results of the pairwise homoplasy index (PHI) test of closely related species using both
LogDet transformation and splits decomposition. The phi test did not find statistically significant
evidence for recombination (p = 1.0).
Figure 10 – Diaporthe biguttulata (JZBH3340160). a Diseased leaf. b Upper view of colony on
PDA after 10 days. c Reverse view of colony on PDA after 10 days. d, e Pycnidia on PDA.
f Conidium. Scale bars: d, e = 100, f = 10 µm. µm.
Figure 11 – Diaporthe eucalyptorum (JZBH3340153). a Diseased leaf. b Upper view on of colony
PDA after 10 days. c Reverse view of colony on PDA after 10 days. d, e Pycnidia on PDA.
f Pycnidial wall. g conidiogenous cell. f hypal end. i–k alpha conidia. Scale bars: d, e = 100 µm,
f–h = 20 µm, h–k = 10 µm.
450
Notes – The taxon isolated in the present study formed a well–supported cluster with
Diaporthe eucalyptorum (CBS 132525) with 88% ML, 54% MP and 0.98 BYPP values. The
morphological characteristics of the isolated taxon are similar to the ex–type isolate of this species
(Crous et al. 2012). Diaporthe eucalyptorum was introduced by Crous et al. (2012) as a leaf spot
causing fungus on Eucalyptus L. This is the first report of D. eucalyptorum on Camellia sinensis
(Farr & Rossman 2020).
Diaporthe fujianensis Jayaward., Manawas., X.H. Li, J.Y. Yan, & K. D. Hyde, sp. nov.
Fig. 12
Index Fungorum: IF557997; Facesoffungi number: FoF09384
Etymology – Epithet refers to the Fujian province from where the type was collected.
Holotype – JZBH3340150
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: not observed; Asexual
morph: Pycnidia on PDA superficial, scattered, black, globose, solitary in most. Conidiophores not
observed. Conidiogenous cells terminal, hyaline and smooth. Alpha conidia 4–6 × 2–3 μm (x̅ = 5 ×
2.5 μm n = 40), biguttulate, hyaline, oval and or ellipsoidal, both ends obtuse. Beta conidia and
gamma conidia were not observed.
Culture characteristics – Colonies on PDA reach 90 mm diam. after five days at 25°C,
producing abundant white aerial mycelia and reverse fuscous white.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried cultures JZBH320150 holotype; JZBH320149,
JZBH320151 and JZBH320152 paratype), and living cultures JZBH320150 ex–holotype;
JZBH320149, JZBH320151 and JZBH320152 ex–Paratype.
Notes – In the phylogenetic analysis four isolates obtained in this study clustered in a well–
supported clade with 100% ML and 84% MP bootstrap values and 0.98 BYPP. In the
recombination analysis, PHI test indicated that the current isolates belong to a species separated
from all other Diaporthe species included in the phylogenetic tree. Diaporthe fujianensis resides in
a sister clade to Diaporthe eucalyptorum. Morphologically the alpha conidia produced by this
species are smaller than those in Diaporthe eucalyptorum (6 × 2.5 µm). A pairwise nucleotide
comparison between Diaporthe eucalyptorum ex type strain (CBS 132525) and Diaporthe
fujianensis ex type strain (JZBH320150) in ITS region showed 1.75% base pair differences along
519 bp. Based on the molecular evidences we consider that these isolates belong to a novel species.
Figure 12 – Diaporthe fujianensis (JZBH3340150 holotype) a Diseased shoot. b Upper view on of
colony PDA after five days. c Reverse view of colony on PDA after five days. d–f alpha conidia.
Scale bars: d–f = 10 µm.
Diaporthe fusiformis Jayaward., Manawas., X.H. Li, J.Y. Yan, & K. D. Hyde, sp. nov.
Fig. 13
Index fungorum: IF557998; Facesoffungi number: FoF09385
Etymology – refers to the fusiform conidia
Holotype – JZBH3340154
451
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: not observed; Asexual
morph: Pycnidia on PDA superficial, scattered, black, globose, solitary and clustered.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth–walled
clustered. Alpha conidia 8–5 × 2–3 μm (x̅ = 7 × 2 μm, n = 40), eguttulate, hyaline, fusiform, both
ends angular. Beta conidia 23–32 ×1.2–1.6 μm (x̅ = 27× 1.5 μm, n = 40), aseptate, hyaline, hamate,
filiform, tapering towards both ends. Gamma conidia not observed.
Culture characteristics – Colonies on PDA reach 90 mm diam. after five days at 25°C,
producing abundant white aerial mycelia and reverse fuscous white becoming gray.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH320154–JZBH320157), and living cultures
JZB320154–7, dried cultures JZBH320154 holotype; JZBH320155–JZBH320157 paratype), and
living cultures JZBH320154 ex–holotype; JZBH320155–7 ex–paratype.
Notes – In the phylogenetic analysis four isolates obtained in this study formed a well–
supported clade with 100% ML and 0.98 PP values. In the recombination analysis, PHI test
indicated that these isolates belong to a species separate from all other species in Diaporthe
(Fig. 9). Diaporthe fusiformis is phylogenetically close to Diaporthe eucalyptorum (CBS132525)
and Diaporthe fujianensis (This study). Diaporthe eucalyptorum (CBS132525) has larger conidia
and Diaporthe fujianensis has smaller conidia (4–6 × 2–3 μm) than Diaporthe fusiformis (8–5 × 2–
3 μm). In addition, the conidia of Diaporthe fusiformis are fusiform whereas Diaporthe
eucalyptorum has biguttulate fusoid conidia and Diaporthe fujianensis has oval to ellipsoidal
conidia. In comparison with Diaporthe lithocarpus; Diaporthe fujianensis has smaller conidia (4–6
× 2–3 μm) than Diaporthe lithocarpus (6–8 × 2–3 μm). Diaporthe lithocarpus develop both alpha
conidia and beta conidia whereas Diaporthe fujianensis is prominent with alpha conidia. Based on
morphological and phylogenetic characters we identified this taxon as a novel species.
Figure 13 – Diaporthe fusiformis (JZBH3340154 Holotype). a–c Pycnidia on PDA.
d Conidiogenus cells. e–g Alpha conidia. h Beta conidia. i Upper view of colony on PDA after five
days. j Reverse view of colony on PDA after five days. Scale bars: b–c = 100 µm, d = 20 µm,
e–h = 10 µm.
Diaporthe nobilis Sacc. & Speg., Michelia 1(no. 4): 386 (1878)
Fig. 14
Index fungorum: IF 153616; Facesoffungi number: FoF02717
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: Not observed.
Asexual morph: Conidiomata 200–350 μm in widest diam, globose, ostiolate, embedded in the
PDA, scattered. Conidiophores 15–22 × 1.5–2 μm, cylindrical, hyaline, rough, branched, septate,
452
straight or slightly curved. Alpha conidia 5.5–8 × 2–3 μm (x̅ = 6 × 2.5 μm, n = 30), unicellular,
hyaline, aseptate, oval, rounded at both ends. Beta and gamma conidia not seen.
Culture characteristics – Cultures incubated on PDA at 25°C in darkness, reach 70 mm diam.,
after seven days. Upper view white, cottony, regular margin. Reverse becoming brownish with age.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li (dried cultures JZBH320158 and JZBH320159), and
living cultures JZB320158 and JZB320159.
Notes – In combined multigene phylogenetic analysis of ITS, tef1, tub2, Cal and HIS, two
strains clustered together with the Diaporthe nobilis (CBS 124030) with 65% ML 61%, MP and
0.99 BYPP values. Colony morphology, spore shape and dimensions are similar to those of
Diaporthe nobilis (Li et al. 2017). So far, this species has been reported on several woody hosts
including tea (Farr & Rossman 2020). In China, Diaporthe nobilis associated with tea was first
reported by Li et al. (2017). However, the pathogenicity of this species has not yet been confirmed.
Figure 14 – Diaporthe nobilis (JZBH3340158). a Diseased shoot. b Upper view of the colony on
PDA after seven days. c Reverse view of the colony on PDA after seven days. d Conidiogenous
cell. e An alpha conidium. Scale bars: d–e = 10 µm.
Diaporthe sackstonii R.G. Shivas, S.M. Thomps. & Y.P. Tan, Persoonia 35: 46 (2015)
Fig. 15
Index Fungorum: IF 808674; Facesoffungi number: FoF09386
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: Not observed. Asexual
morph: Pycnidia on PDA solitary, scattered, ostiolate, cream conidial droplets exuded from some
ostioles. Conidiophores reduced to conidiogenous cells. Conidiogenous cells filiform, hyaline to
pale yellowish–brown. Alpha conidia 6–8 × 2–2.5 μm (x̅ = 6.5 × 2 μm, n = 30), abundant, fusiform,
rounded at the apex, obconically truncate at base, hyaline. Beta conidia not observed.
Culture characteristics – Colonies on PDA covering entire plate after 10 days. White areal
myclilum, entire margine, with age a few scattered dark stromata up to 1 mm diam., buff. Reverse
white and become black with age.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried culture JZBH320165), and living culture JZB320165.
Notes – The single isolate obtained from the present study formed a well–supported clade
with the ex–type strain of Diaporthe sackstonii (BRIP54669b) with 83% ML, 83% MP and 0.98
BYPP values. Morphologically these two isolates are similar and they share sequences difference
of less than 1% at each gene region (at three genes ITS, tub2 and tef1). Diaporthe sackstonii was
introduced by Thompson et al. (2015) on Helianthus annuus in Australia. This is the first report of
Diaporthe sackstonii on Camellia sinensis (Farr & Rossman 2020).
Diaporthe sennae C.M. Tian & Qin Yang, in Yang et al., Phytotaxa 302(2): 149 (2017)
Fig. 16
Index Fungorum: IF820452; Facesoffungi number: FoF08696
453
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: Not observed. Asexual
morph: Conidiomata pycnidial, circular to ovoid, immersed, scattered on PDA, Conidiophores
reduced to conidiogenous cells. Conidiogenous cells hyaline, phialidic, straight or slightly curved.
Alpha conidia 5–7 × 1.5–2 μm (x̅ = 6.0 × 2 μm, n = 30), hyaline, aseptate, smooth, ellipsoidal to
oval, usually one guttulate at each end, rarely 3 small guttulate. Beta conidia not observed.
Figure 15 – Diaporthe sackstonii (JZBH3340165) a Diseased shoot. b Upper view of mycelium on
PDA after 10 days. c Reverse view of mycelium on PDA after 10 days. d Pycnidia on PDA.
e Alpha conidia. Scale bars: d = 100 µm, e = 10 µm.
Figure 16 – Diaporthe sennae (JZBH3340147). a Diseased shoot. b Upper view of mycelium on
PDA after 10 days. c Reverse view of mycelium on PDA after 10 days. d–e Alpha conidia. Scale
bars: d–e = 10 μm.
Culture characteristics – Colonies on PDA covering the entire plate after 10 days. Colony flat
with white flat aerial mycelium, becoming pale brown mycelium due to pigment formation,
conidiomata absent
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried culture JZBH320147), and living culture JZB320147.
Notes – One isolate from the present study clustered together with the ex–type of Diaporthe
sennae (CFCC 51636) with 76% ML, 50% MP and 1.00 BYPP. Morphologically the strain isolated
in this study shares similar characters with the type description (Yang et al. 2017). Diaporthe
454
sennae was introduced from infected branches/twigs of Senna bicapsularis in China (Yang et al.
2017). However, pathogenicity of this species has not been confirmed. To our knowledge, this is
the first report of Diaporthe sennae on Camellia sinensis (Farr & Rossman 2020).
Diaporthe sinensis Jayaward., Manawas., X.H. Li, J.Y. Yan, & K. D. Hyde, sp. nov.
Fig. 17
Index Fungorum: IF557999; Facesoffungi number: FoF09387
Etymology – Name derived from the epithet of the host
Holotype – JZBH320167
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: not observed: Asexual
morph: Pycnidia on PDA 360–900 μm (x̄ = 500 µm, n = 20) in diam., superficial, scattered, dark
brown to black, globose, solitary in most. Conidiophores reduced to Conidiogenous cells.
Conidiogenous cells hyaline, simple, smooth terminal. Alpha conidia 7–4 ×2–3 μm (x̅ = 5 × 3 μm,
n = 40) hyaline, oval, both ends obtuse. Beta conidia and gamma conidia not observed.
Culture characteristics – Colonies on PDA reach 90 mm diam., after five days at 25°C,
producing abundant white aerial mycelia and reverse fuscous white.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH320167 holotype, JZBH320168–9
paratype), and living cultures ZBH320167 ex–holotype; JZBH320168–9 ex–paratype.
Notes – In the phylogenetic analysis, four isolates obtained in this study formed a well–
supported clade with 70% ML and 68% MP bootstrap values and 0.97 BYPP. These taxa show
particular neighbour relation to Diaporthe amygdali (CBS 126679). Compared to the sister species,
Diaporthe sinensis develops oval and shorter alpha conidia whereas conidia of Diaporthe amygdali
are fusiform, and biguttulate (Gomes et al. 2013). A comparison of the ITS (497bp), tef1 (492bp),
and Cal (300bp) between our species (JZBH3340167) and closely associated Diaporthe amygdali
(CBS 126679) revealed 2%, 2.4% and 14% base pair differences respectively. Therefore, based on
both morphological and phylogenetic evidence we identified these isolates as a novel Diaporthe
species associated with tea.
Figure 17 – Diaporthe sinensis (JZBH3340167 Holotype) a Diseased leaf. b Upper view of
mycelium on PDA five days. c Reverse view of mycelium on PDA five days. d Pycnidia on PDA.
e–f Alpha conidia. Scale bars: e = 10 µm, f = 5 µm.
Diaporthe unshiuensis F. Huang, K.D. Hyde & Hong Y. Li, in Huang et al., Fungal Biology
119(5): 344 (2015)
Fig. 18
455
Index Fungorum: IF 810845; Facesoffungi number: FoF 09422
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: Not observed.
Asexual morph: Conidiomata 100–300 μm in diam., globose to subglobose, dark brown to black,
cream conidial drops exuded from the ostioles. Conidiophores not observed. Conidiogenous cells
cylindrical, hyaline. Alpha conidia 6–8 × 2–3 μm (x̅ = 6 × 3 μm, n = 30), unicellular, aseptate,
fusiform, hyaline, biguttulate and tapering towards both ends. Beta conidia not observed.
Culture characteristics – Cultures incubated on PDA at 25°C reach 90 mm., after seven days.
Colony at first white, becoming pale brownish, reverse pale yellowish at the centre with age. Aerial
mycelium white, cottony, with slightly fringed margin and conidiomata visible at maturity.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li (dried cultures JZBH320160– JZBH320164), and
living cultures JZB320160– JZB320164.
Notes – Five isolates obtained in the present study clustered together with the ex–type of
Diaporthe unshiuensis (ZJUD52) with 100% ML, 100% MP and 1.00 BYPP values.
Morphologically isolates from this study are similar to the type description of Diaporthe unhuensis
(Huang et al. 2015). This species was reported on Citrus unshiu in China (Huang et al. 2015) and
this is the first report of D. unshiuensis on Camellia sinensis (Farr & Rossman 2020).
Figure 18 – Diaporthe unhuensis (JZBH3340163) a Diseased leaves and shoot. b Upper view of
mycelium on PDA after seven days. c Reverse view of mycelium on PDA after seven days.
d–e Pycnidia on PDA. f Pycnidial wall. g Conidiogenus cells attached to the pycnidial wall.
h Alpha conidia. Scale bars: d, e = 100 µm, f–g = 20 µm, h= 10 µm.
Diaporthe viniferae Dissanayake, X.H. Li & K.D. Hyde, in Manawasinghe et al., Frontiers in
Microbiology 10: 21 (2019)
Fig. 19
Index Fungorum: IF552002; Facesoffungi number: FoF05981
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: not observed; Asexual
morph: Pycnidia on PDA 400–900 μm (x̄ = 500 µm, n = 20) superficial, scattered, dark brown to
black, globose, solitary in most. Conidiophores not observed. Conidiogenous cells not observed.
Alpha conidia 5–8 × 1–2.5 μm (x̅ = 6 × 2 μm, n = 40), biguttulate, hyaline, fusiform or oval, both
ends obtuse, Beta conidia 20–30 ×1–1.5 μm (x̅ = 27 × 1 μm, n = 40), aseptate, hyaline, filiform.
Culture characteristics – Colonies on PDA reach 90 mm diam., after five days at 25°C,
producing abundant white aerial mycelia and reverse fuscous white.
456
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried culture JZBH320148), and living culture JZB320148.
Notes – In the phylogenetic analysis, the single isolate clustered together with Diaporthe
viniferae (JZBH3340148), 85% ML and 76% MP bootstrap values and 1.00 BYPP.
Morphologically these two isolates are similar with no differences in sequence data. This species
was first introduced by Manawasinghe et al. (2019) as a pathogen associated with grapevine
dieback in China. This is the first report of Diaporthe viniferae on Camellia sinensis (Farr &
Rossman 2020).
Figure 19 – Diaporthe viniferae (JZBH3340148). a Diseased shoot. b Upper view on PDA after
five days. c Reverse view on PDA after five days. d An alpha conidium. Scale bar: d = 10 µm.
Subclass Hypocreomycetidae O.E. Erikss. & Winka, Myconet 1: 6 (1997)
Notes – Currently there are seven orders; Coronophorales, Falcocladiales, Glomerellales,
Hypocreales,
Microascales,
Parasympodiellales
and
Torpedosporales associated
with
Hypocreomycetidae with 37 families (Hyde et al. 2020b).
Glomerellales Chadef. ex Réblová, W. Gams & Seifert, Studies in Mycology 68: 170
Notes – Chadefaud (1960) introduced Glomerellales. This order is composed of endophytic
fungi and phytopathogens with ascomata varying from endostromatal to apostromatal and
ascospores that are often unicellular and hyaline. Currently, five families are accepted in the
Glomerellales: Glomerellaceae, Australiascaceae, Malaysiascaceae, Plectosphaerellaceae and
Reticulascacea (Hyde et al. 2020b).
Glomerellaceae Locq., Mycol. gén. struct. (Paris): 175 (1984).
Notes – Almost all species identified in this family are well–known plant pathogens on a
wide range of hosts (Jayawardena et al. 2016a). Type genus of this family is Colletotrichum (Hyde
et al. 2014, Maharachchikumbura et al. 2016).
Colletotrichum Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(12): 41 (1831)
Notes – Species in this genus are known as pathogens on a wide range of crops and some
species are endophytes or saprotrophs (Hyde et al. 2014, Jayawardena et al. 2016a, 2020). Species
delimitation based on morphology alone is difficult in Colletotrichum due to overlapping
morphological characters in the asexual morphs (Hyde et al. 2009, Cannon et al. 2012). Therefore,
polyphasic approaches including multi–locus sequence analyses are essential (Jayawardena et al.
2016a). Currently, 14 species complexes (Jayawardena et al. 2016a, Damm et al. 2019, Bhunjun et
al. 2021) are accepted in this genus. In the present study, we obtained two isolates belonging to two
known species of Colletotrichum (Fig. 20).
Colletotrichum camelliae Massee, Bull. Misc. Inf., Kew: 91 (1899)
Fig. 21
Index Fungorum: IF176099; Facesoffungi number: FoF09388
Pathogenic or saprobic on Camellia sinensis leaves, Sexual morph: Ascomata on PDA
perithecia, globose, ovoid, obpyriform, aggregated or scattered, immersed, single ostiole. Ascomata
457
wall thick, the outer wall of ascomata composed of flattened angular cells, Asci clavate, 60–80 ×
10–14 µm (x̅ = 60 × 12 µm, n = 40) long, 8 spored, apex truncated and a small apical point. Asci
covered with a thick sheath. Ascospores hyaline 13–18 × 4–5 µm (x̅ = 16 × 4.5 n = 20), one–celled,
allantoid or fusiform. Asexual morph: not observed.
Culture characteristics – Colonies reach <90 mm diam., in 10 days, flat with an entire edge,
aerial mycelium white, cottony, sparse; reverse white at first, then grey to black at the centre.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH330153), and living culture JZB330153.
Figure 20 – Phylogenetic tree generated by ML analysis of combined ITS, glyceraldehyde–3–
phosphate dehydrogenase (GAPDH), chitin synthase (CHS–1), actin ACT and tub2 sequence data
of Colletotrichum species. In the phylogenetic tree, Colletotrichum boninense (CBS123755) and
Colletotrichum catinaense (CBS 142417) used as outgroup. Tree topology of the ML analysis was
similar to the BI. The best scoring RAxML tree with a final likelihood value of –10102.441238 is
presented. The matrix had 812 distinct alignment patterns, with 15.39% of undetermined characters
458
or gaps. Estimated base frequencies were as follows: A 0.229139, C = 0.296735, G = 0.244809, T =
0.229316; substitution rates AC = 1.065846, AG = 2.974226, AT = 0.978030, CG = 0.858886, CT
= 4.711667, GT = 1.000000; gamma distribution shape parameter α = 1.663370. RAxML bootstrap
support values ≥50% and Bayesian posterior probabilities ≥0.95 (BYPP) are shown near the nodes.
The scale bar indicates 0.04 changes per site. Ex–type/ex–epitype) strains are in bold and new
isolates recovered in the the present study are in red.
Figure 21 – Colletotrichum camelliae (JZB330153). a–b Ascomata on PDA. c Ascomatal wall
(surface view). d–f Developing and mature asci. g Ascospores. h Upper view of the colony on PDA
after 10 days. i Reverse view of the colony on PDA after 10 days. Scale bars: a–c = 100 µm.
d–f = 20 µm, g = 10 µm.
Notes – A strain isolated in the present study clustered together with the C. camelliae
(CGMCC 3.14925) within the gloeosporioides complex with 83% ML bootstrap value and 0.90
BYPP. The species isolated in this study was confirmed as C. camelliae based on both
morphological characters and phylogenetic placement. Colletotrichum camelliae was introduced as
Glomerella cingulata ‘f. sp. camelliae’ Dickens & R.T.A. Cook., which has been reported as
causing twig blight and brown blight of Camellia. This species can be observed in many tea
growing regions (Liu et al. 2015, Wang et al. 2016b).
Colletotrichum fructicola Prihast., L. Cai & K.D. Hyde, in Prihastuti et al., Fungal Diversity 39: 96
(2009)
Fig. 22
Index Fungorum: IF515409; Facesoffungi number: FoF06767
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: not observed. Asexual
morph: Conidiophore reduced to a conidiogenous cell. Conidiogenous cell hyaline, thick
ampliform, Conidia 10–14 × 3–4 μm (x̅ = 10 × 3 μm, n = 40), common in mycelium, one–celled,
smooth–walled with a large guttule at the centre and surrounded by smaller guttules, hyaline,
cylindrical with obtuse to slightly rounded ends, sometimes oblong.
Culture characteristics – colonies on PDA reaches 90 mm at mm diam., in seven days at
25°C. Colonies are white initially then become grey to dark grey with age. Reverse greyish to
black. Aerial mycelium pale grey, dense, cottony, without visible conidial masses.
Material(s) examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH330154), and living culture JZB330154.
459
Notes – In the present study an isolate obtained from tea leaves (out of two Colletotrichum
isolates) clustered together with Colletotrichum fructicola (ICMP 18581) with 97% ML bootstrap
and 0.97 PP. Colony characters and morphology (conidial shape and sizes) are similar to the ex–
type isolate (Prihastuti et al. 2009). Based on morphology and phylogeny we confirmed our isolate
as Colletotrichum fructicola. This species was introduced by Prihastuti et al. (2009) as a taxon
associated with Coffea arabica. However, this species has been reported on C. sinensis from China
and Indonesia (Liu et al. 2015).
Figure 22 – Colletotrichum fructicola (JZB330154). a–b Conidiogenus cells with developing
conidia. c–e Conidia. f Hyphae. g Upper view of the colony on PDA after 10 days. h Reverse view
of the colony on PDA after 10 days. Scale bars: a–b = 20 µm, c–f = 10 µm.
Hypocreales Lindau, Natürl. Pflanzenfam.: 343 (1897)
Notes – Species belonging to Hypocreales are highly diverse in the tropics, subtropics and
temperature regions (Põldmaa 2011). Hypocreales accepted with family Bionectriaceae,
Calcarisporiaceae, Clavicipitaceae, Cocoonihabitaceae, Cordycipitaceae, Flammocladiellaceae,
Hypocreaceae, Myrotheciomycetaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae,
Sarocladiaceae, Stachybotryaceae, and Tilachlidiaceae (Maharachchikumbura et al. 2016, Hyde et
al. 2020b).
Hypocreaceae De Not., [as ‘Hypocreacei’], G. bot. ital. 2(1): 48 (1844).
Notes – Species belonging to this family are diverse are biotrophic, hemibiotrophic,
saprobic or hypersaprobic on a wide range of hosts. For recent taxonomic treatments, we follow
Hyde et al. (2020b).
Trichoderma Pers., Neues Magazin für die Botanik 1: 92 (1794)
Notes – Species belonging to Trichoderma have a wide range of life modes that includes
hypersaprobic on Basidiomycetes (Schuster & Schmoll 2001, Chen & Zhuang 2017). Some of the
taxa are important as they produce industrially important enzymes (cellulases and hemicellulases),
antibiotics, and some are used in biocontrol agents (Sivasithamparam & Ghisalberti 1998). In this
study, we isolated eight strains belonging to three species including one novel species (Fig. 23).
460
Figure 23 – Phylogenetic tree generated by ML analysis of combined ITS, rpb2 sequence data
Trichoderma species. 65 strains are included in the analyses. The tree is rooted with Nectria
eustromatica (CBS 125578). Tree topology of the ML analysis was similar to BI. The best scoring
RAxML tree with a final likelihood value of – 24349.980578 is presented. The matrix had 1172
distinct alignment patterns, with 9.91% of undetermined characters or gaps. Estimated base
frequencies were as follows: A = 0.251668, C = 0.245757, G = 0.259668, T = 0.242908;
substitution rates AC = 1.353890, AG = 4.605576, AT = 1.059439, CG = 0.801610, CT =
9.121730, GT = 1.000000; gamma distribution shape parameter α = 0.944898. RAxML bootstrap
support values ≥50% and Bayesian posterior probabilities ≥0.95 (BYPP) are shown near the nodes.
The scale bar indicates 0.02 changes per site. Ex–type/ex–epitype strains are in bold. New isolates
recovered in this study are in red.
Trichoderma atroviride P. Karst. 1892
Index Fungorum: IF451289; Facesoffungi number: FoF09389
Fig. 24
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Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: Undetermined. Asexual
morph Conidiophores tree–like, comprising a main axis with second branches, second branches
paired, sometimes second branches branched again, main axis and branches terminating in whorls
of up to five phialides. Conidiogenous cells phialidic, lageniform or ampulliform, arising singly,
non–equilateral when curved. Conidia 4–5 × 3–3.5 μm (x̅ = 4 ×3 μm, n = 30), ovoid, verrucose.
Culture characteristics – On PDA mycelium covers plate after three days at 25°C. Margin
conspicuous and radial. Aerial hyphae, hairy to floccose, dense internal zone, but relative sparse on
margin, abundantly and flat in a large green disc around the inoculum, turning green after 24 h of
conidiation.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried culture JZBH3360001), and living cultures JZB3360001.
Figure 24 – Trichoderma atroviride (JZB3360001). a–c Conidiomata on PDA. d–f Branched
conidiogenous cells. g–i Conidia. j, k Septate mycelia. l Upper view of the colony on PDA after
three days. m Reverse view of the colony on PDA after three days. Scale bars: a–c, j, k = 100 µm,
d–j = 10 µm.
462
Notes – The single isolate obtained in the present study clustered together with the
Trichoderma atroviride (GAOM 222144) with 99% ML and 1.0 BYPP. Morphologically the strain
isolated in the present study is similar to the species description of the type specimen (Brunner et
al. 2005). Trichoderma atroviride is commonly isolated from soil and it is a well–known biocontrol
agent (Brunner et al. 2005). This species has been reported on Betula papyrifera, Morus sp.,
Triticosecale sp., Vitis vinifera, and Zea mays (Farr & Rossman 2020). This is the first report of T.
atroviride on Camellia species (Farr & Rossman 2020).
Trichoderma camelliae Jayaward., Manawas., X.H. Li, J.Y. Yan, & K. D. Hyde, sp. nov.
Fig. 25
Index Fungorum: IF558000, Facesoffungi Number: FoF09390
Etymology – refers the host genus
Holotype – JZBH3360002
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: Not observed.
Asexual morph Mycelia aseptate, branched, effused Conidiophores scattered, dark green to
greyish–green, tree–like, comprising a main axis. Conidiogenous cells ampulliform, arising singly
as clusters. Conidia developed at the hyphal end also observed. Conidia 1.5 –2× 1–2 μm (x̅ = 2×2
μm, n = 40) ovoid to short ellipsoidal, verrucose.
Culture characteristics – On PDA mycelium covers plate after three days at 25°C. Aerial
hyphae, hairy dense internal zone, initially white mycelium with time become pale yellow. Develop
abundant, and flat large green disc around the inoculum, turning green.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li (dried cultures JZBH3360002 holotype;
JZBH3360003 – JZBH3360006 paratypes), and living JZB3360002 ex–holotype; JZB3360003 –
JZB3360006 ex–paratypes.
Notes – The isolates obtained in the present study fit well morphologically within the
Trichoderma. The present species, Trichoderma camelliae, developed a strongly supported
monophyletic clade with 100% ML and 1.0 BYPP values. Morphologically this species differs
from the type species of Trichoderma viride, by developing ellipsoidal and larger conidia, whereas
conidia of the type species are mostly ovoid and smaller than the species identified in this study
(0.7 µm long and 1 µm diam.) (Lieckfeldt et al. 1999).
Figure 25 – Trichoderma camelliae (JZB3360002 ex–holotype). a Diseased leaf. b Upper view of
the colony on PDA after three days. c Reverse view of the colony on PDA after three days.
d–e Conidiomata on PDA. f Pycnidal wall. g Conidiogenous cell. h–i Conidia. j Germinating
conidium. Scale bars: d, e = 100 µm, f, g = 100 µm, h–j = 10 µm.
463
Trichoderma lixii (Pat.) P. Chaverri, in Chaverri et al., Mycologia 107(3): 578 (2015)
Fig. 26
Index Fungorum: IF 809999; Facesoffungi number: FoF09391
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Undetermined. Asexual
morph: Mycelium aseptate, less branched, developed effused. Conidiomata pycnidial, black,
superficial. Conidiophores less branched, branches arise horizontally from the main axis initially
yellow later turning grey. Conidiogenous cells phialidic ampulliform, arising solitary, haline thin–
walled, smooth, Conidia 2–4 × 1–2 μm (x̅ = 3–1.5 μm, n = 30), ovoid, verrucose Clamydospres
developed at the terminals of the hyphal tips, ovoid, various in size, develop single germination
tube.
Culture characteristics – On PDA mycelium covers the plate after three days at 25°C. Colony
layered distinctly, margin conspicuous and radial. Arial hyphae, hairy to the floccose, dense
internal zone. Pycnidia appear as concentric rings, dense near the edge of the plate. Initially white
and become olivaceous yellow. Reverse olivaceous brown.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH3360007 and JZBH3360008), living
cultures JZB3360007 and JZB3360008.
Notes – The species identified in the present study clusters together with Trichoderma lixii
(GJS 97.96) with 44% ML, and 1.0 BYPP values. This isolate is morphologically similar to the
type species description of T. lixii (Chaverri et al. 2015). In pairwise nucleotide comparison of ITS
region (534bp) between our species (JZB3360008) and closely associated Trichoderma lixii (GJS
97.96) revealed 0.37% base pair differences. However, rpb2 sequence is available for only one
strain isolated in this study. This might be the reason the three isolates obtained in this study
develop a distinct cluster. Based on these we identified the strains in this study as T. lixii. This is
the first report of T. lixii on Camellia species (Farr & Rossman 2020).
Figure 26 – Trichoderma lixii (ZB3360007). a–c Conidiomata on PDA. d–f Conidiogenous cells
e–f Branched conidiogenous cells. g–i Conidia. j Septate mycelia. j Upper view of the colony on
464
PDA after three days. k Reverse view of the colony on PDA after three days. Scale bars: a–c = 100
µm. d–f, j = 20 µm. g–i = 10 µm.
Nectriaceae Tul. & C. Tul., Selecta Fungorum Carpologia: Nectriei–Phacidiei– Pezizei 3: 3 (1865)
Notes – Nectriaceae species are commonly found as saprobes, plant endophytes or pathogens,
mycopathogen or pathogens on insects (Hyde et al. 2020b). For taxonomic treatments, we follow
Hyde et al. (2020b).
Fusarium Link, Magazin der Gesellschaft Naturforschenden Freunde Berlin 3 (1): 10 (1809
Notes – Fusarium species are well–known plant pathogens, saprobes and some species
produce mycotoxins that can contaminate food (Perincherry et al. 2019). For the species level
characterisation of Fusarium morphological characters together with molecular data are required.
In this study, we isolated nine strains that belong to three species representing three new host
records (Fig. 27).
Fusarium asiaticum O’Donnell, T. Aoki, Kistler & Geiser, in O’Donnell, Ward, Geiser, Kistler &
Aoki, Fungal Genetics Biol. 41(6): 619 (2004)
Fig. 28
Index Fungorum: IF809999; Facesoffungi number: FoF09392
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: not observed. Asexual
morph: Conidia develop in the aerial mycelium. Conidiophores not observed. Conidia conidia 30–
40 × 2–4 μm (x̅ = 30 × 3 μm, n = 30), sporodochial conidia gradually curved and frequently widest
above the mid–region, septate, smooth and thin–walled. Chlamydospores not seen.
Culture characteristics – Colonies on PDA covers the entire plate within five days. Entire
margin, aerial mycelium reddish–white velvety to lanose. Pigmentation in reverse, sclerotia absent
later becomes dark purple.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li dried cultures JZBH110019–23), and living cultures
JZB110019–23.
Notes – Five isolates from in this study cluster together with the Fusarium asiaticum
(CBS110257) with 99% ML and 87% MP bootstrap values. These isolates share similar
morphology to the type description of F. asiaticum (Leslie & Summerell 2008). Fusarium
asiaticum is a well–known pathogen causing Fusarium head blight (Qiu et al. 2019). This species
has been reported on Bletilla striata, Glycine max, Hordeum vulgare, Lolium multiflorum, Oryza
sativa, Triticum aestivum and Zea mays, which are all monocot plants (Farr & Rossman 2020). This
is the first report of Fusarium asiaticum associated with Camellia sinensis (Farr & Rossman 2020).
Fusarium concentricum Nirenberg & O’Donnell Mycologia 90(3): 442 (1998)
Fig. 29
Index Fungorum: IF 809999; Facesoffungi number: FoF 09423
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: Not observed.
Asexual morph: Sporulation is starting early in the aerial mycelium. Conidia develop in false
heads, the aerial conidiophores. Conidiogenous cells monophialides and polyphialides cylindrical
flask–shaped. Conidia 8–12 × 3–4 μm (x̅ = 10 × 3 μm, n = 30), develop in the aerial mycelium,
oval, obovoid to allantoid, aseptate, smooth– and thin–walled, Chlamydospores not observed.
Sporodochial conidia not observed.
Culture characteristics – Colonies on PDA grow 45mm diam., after five days. Entire margin,
aerial mycelium white velvety. Pigmentation in reverse initiates after 10–14 days, pale orange and
reddish grey concentric rings, later becoming dark purple.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li dried cultures JZBH3110010– JZBH3110014), and
living cultures JZB3110010– JZB3110014.
Notes – Six isolates obtained in the present study cluster together with the Fusarium
concentricum in the phylogenetic analysis forming strongly supported clade with 100% ML and
465
100% MP bootstrap values. Morphologically species identified in the present study share similar
characters to those of the Fusarium concentricum type species (Leslie & Summerell 2008).
However, we did not observe sporodochial conidia after 10 days of incubation. This species has
been reported on several different hosts including Capsicum annum (Wang et al. 2013), Musa sp.
(Sandoval–Denis et al. 2018), Nilaparvata lugens (Nirenberg & O’Donnell 1998), Oryza sativa
(Aoki et al. 2002, Choi et al. 2019), Paris polyphylla var. chinensis (Xiao et al. 2019), Triticum
aestivum (Aoki et al. 2002) and Vanilla sp. (Koyyappurath et al. 2016). This is the first report of
Fusarium concentricum on Camellia sinensis (Farr & Rossman 2020).
Figure 27 – Phylogenetic tree generated by MP analysis of combined tef1 and rpb2 sequence data
of Fusarium species. Eighty strains are included in the analyses. Fusarium buharicum (CBS
796.70) and Fusarium sp. NRRL 66182 were used to root the tree. Tree topology of the ML
analysis was similar to the MP and BI. The best scoring RAxML tree was with a final likelihood
466
value of –7573.307178 is presented. The matrix had 358 distinct alignment patterns, with 2.33% of
undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.280950,
C = 0.235343, G = 0.263312, T = 0.220394; substitution rates AC = 2.323636, AG = 8.480494,
AT = 2.016299, CG = 1.445800, CT = 22.415448, GT = 1.000000; gamma distribution shape
parameter α = 1.499621. Maximum parsimony analysis of 864 constant characters and 302
informative characters resulted in 62 equally most parsimonious tree of 462 steps (CI = 0.381, RI =
0.836, RC = 0.318, HI = 0.619). RAxML bootstrap support and maximum parsimony bootstrap
support values ≥50% are shown near the nodes. The scale bar indicates 0.05 changes per site. Ex–
type (ex–epitype) strains are in bold. New isolates recovered in this study are given in red.
Figure 27 – Continued.
Fusarium fujikuroi Nirenberg, Mitt. biol. BundAnst. Ld– u. Forstw. 169: 32 (1976)
Fig. 30
Index Fungorum: IF 809999; Facesoffungi number: FoF09393
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: not observed.
Asexual morph: Sporulation is starting early in the aerial mycelium. Aerial conidiophores
cynlindrical mono and polyphialidic. Conidia 8–26 × 2–5 μm (x̅ = 16 × 3 μm, n = 40), develop in
the aerial mycelium obovoid and oval to allanoid, asepatae, smooth– and thin–walled,
chlamydospores absent.
Culture characteristics – Colonies on PDA grows covers the entire plate within five days.
Colony margin entire, aerial mycelium reddish–white velvety to lanose. Pigmentation in reverse
consisting of the concentric pink ring the middle and pale orange ring at the margin. Later
becoming dark purple.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li dried cultures JZBH110017 and JZBH110018), and
living cultures JZB110017 and JZB110018.
467
Notes – Two isolates obtained in this study cluster with Fusarium fujikuroi (NRRL13566)
representative strain by forming a strongly supported clade with 100% ML, and 100% MP
bootstrap values. In a comparison of morphology and sequence data, these two strains did not show
any significant differences. Therefore, we confirmed these two strains as Fusarium fujikuroi. This
species has been reported causing Fusarium wilt of soybean, rice and barnyard grass in Korea (Choi
et al. 2019). This the first report of this species associated with Camellia sinensis in China (Farr &
Rossman 2020).
Figure 28 – Fusarium asiaticum (JZB110020). a Diseased shoot. b Upper view of the colony on
PDA after five days. c Reverse view of the colony on PDA after five days. d Conidiophores of
aerial mycelium. e Conidia. Scale bars: d 20 µm. e = 10 µm.
Figure 29 – Fusarium concentricum (JZB110013). a Diseased leaves and shoots. b Upper view of
the colony on PDA after five days. c Reverse view of the colony on PDA after five days.
d Conidiophores of aerial mycelium. e Conidia. Scale bars: d = 20 µm, e = 10 µm.
Fusarium proliferatum (Matsush.) Nirenberg ex Gerlach & Nirenberg, Mitt. biol. BundAnst. Ld–
u. Forstw. 169: 38 (1982)
Fig. 31
Index Fungorum: IF 809999; Facesoffungi number: FoF09394
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: not observed. Asexual
morph: Sporulation is starting early in the aerial mycelium. Conidia develop in false heads of
mycelia, the aerial conidiophores cynlindrical mono and pluphyialidic, phialides flask–shaped.
Conidia 12–21 × 2–5 μm (x̅ = 16 × 3 μm, n = 30), develop in the aerial mycelium obovoid and oval
468
to allanoid, mostly asepatae occasionally one septate, smooth and thin–walled, Sporodochial
conidia rare, septate, smooth and thin–walled. Chlamydospores not observed.
Culture characteristics – Colonies on PDA grows 45mm diam., after five days. Colony
margin is entire aerial mycelium white velvety to lanose. Pigmentation not observed. Colony
surface dry, white becoming livid pink towards the margin, turning completely light pink with age.
Material examined – CHINA, Fujian Province, Zhangzhou County, pathogenic on dead
Camellia sinensis shoot, June 2015, H.L. Li dried cultures JZBH110016, and living cultures
JZB110016.
Notes – The single isolate obtained in this study clustered together with Fusarium
proliferatum (CBS 217.76) representative strain by developing a strong clade with 98% ML and
96% MP bootstrap values. In a comparison of morphology and sequence data, these two strains
share the same characters. This species is a well–known pathogenic species causing diseases in
Maize (Visentin et al. 2009). There are 199 records under this species in Farr & Rossman (2020)
database. This the first report of this species associated with Camellia sinensis in China (Farr &
Rossman 2020).
Figure 30 – Fusarium fujikuroi (JZB110018). a Diseased leaf. b Upper view of the colony on PDA
after five days. c Reverse view of the colony on PDA after five days. d Hyphae. e conidiophores.
f–h conidia. Scale bars: d–h = 10 µm.
Figure 31 – Fusarium proliferatum (JZB110016). a Sporocadial conidia. b–d conidia. e Septate
mycelia. f Upper view of the colony on PDA after five days. g Reverse view of the colony on PDA
after five days. Scale bars: a–e = 10 µm.
469
Subclass Sordariomycetidae O.E. Erikss. & Winka, Myconet 1: 10 (1997)
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Systema Ascomycetum 5: 182 (1986)
Notes – Sordariales was introduced by Hawksworth & Eriksson (1986) and consists of three
families Chaetomiaceae, Sordariaceae and Lasiosphaeriaceae. The Sordariales species are
characterized by membranous or coriaceous ascomata, and hyaline or brown ascospores often with
appendages or sheaths (Zhang et al. 2006). The taxonomic treatment follows Maharachchikumbura
et al. (2016) and Hyde et al. (2020b).
Chaetomiaceae G. Winter [as ‘Chaetomieae’], Rabenh. Krypt.–Fl., Edn 2 (Leipzig) 1.2: 153 (1885)
Notes – Chaetomiaceae belongs to Sordariales (Wijayawardene et al. 2020). The species in
this family are mostly opportunistic fungi in both animals and plants (Plumlee et al. 2017). Some
species are commonly found in plant debris and they play a significant role in degradation of plant
debris (Plumlee et al. 2017).
Chaetomium Kunze, Mykologische Hefte (Leipzig) 1: 15 (1817)
Notes – Chaetomium was established by Kunze (Kunze & Schmidt 1817). Since then, this
genus has undergone several taxonomic reassignments (Wang et al. 2016a). For taxonomic
treatments we refer to Wang et al. (2016a). In the present study, we identified two strains that
belong to a novel species based on both morphology and phylogeny (Fig. 32).
Figure 32 – Phylogenetic trees generated by MP analysis of combined LSU, ITS, tub2, tef1 and
rpb2 sequence data of Chaetomium species. The tree is rooted with Achaetomium strumarium (CBS
333.67). Tree topology of the ML analysis was similar to the MP and BI. The best scoring RAxML
tree was with a final likelihood value of –16838.646969. The matrix had 853 distinct alignment
patterns, with 4.88% of undetermined characters or gaps. Estimated base frequencies were as
follows: A = 0.229473, C = 0.292519, G = 0.272427, T = 0.205580; substitution rates AC =
0.805009, AG = 3.277592, AT = 0.803238, CG = 1.189987, CT = 5.975471, GT = 1.000000;
gamma distribution shape parameter α = 0.564760. Maximum parsimony analysis of 3388 constant
470
characters and 639 informative characters resulted in five equally most parsimonious tree (TL =
2253, CI = 0.525, RI = 0.828, RC = 0.435, HI = 0.475). RAxML bootstrap support values ≥75%
and MP bootstrap support values ≥50% are shown near the nodes. Nodes with BYPP ≥0.95 are
thicked. The scale bar indicates 0.02 changes per site. Ex–type/ ex–epitype strains are in bold and
new isolates recovered in the present study are in red.
Figure 32 – Continued.
Chaetomium camelliae Jayaward., Manawas., X.H. Li, J.Y.Yan, & K. D. Hyde, sp. nov.
Fig. 33
Index fungorum: IF558001; Facesoffungi number: FoF03512
Etymology – The specific epithet is derived from that of the host plant
Holotype – JZBH3340001
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Ascomata superficial,
ostiolate, yellowish to greenish olivaceous subglobose, 165–315 μm diam. Ascomatal wall brown,
471
composed of hypha–like cells, textura intricata in surface view. Asci fasciculate, clavate, 20–30 ×
10–15 μm (x̅ = 20 × 10 μm, n = 20), stalks 20–40 μm long, with 6–7 ascospores, Ascospores 10–12
× 6–8 μm (x̅ = 10 × 7 μm, n = 40), hyaline at the begin become olivaceous brown when mature,
limoniform, bilaterally flattened slightly with age, with an apical germ pore. Asexual morph: not
observed.
Culture characteristics – Colonies on PDA grow 95 mm diam., within five days, yellowish
floccose aerial hyphae, and greenish exudates; reverse light brown.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures; JZBH3340001 holotype; JZBH3340002
isotype–2) and living cultures JZB3340001 ex–holotype; JZB3340002 ex–isotype.
Notes – Preliminary data analysis of ITS region revealed the taxon isolated in the present
study belongs to Chaetomium. According to the phylogenetic analysis based on LSU, ITS, tef1 and
tub2, the isolates obtained from the current study developed a clade sister to Chaetomium
pseudoglobosum (CBS 574.71) with 100% ML bootstrap, 99% MP bootstrap and 1.0 BYPP. In a
pairwise sequence comparison between the sequences of the type of the present study and
Chaetomium pseudoglobosum (CBS 574.71), there was 8% nucleotide difference in LSU along
with the 584 nucleotides and 4% nucleotide difference in ITS along the 521 nucleotides. In
comparisons of protein-coding regions; there were 3% differences in tef1 (out of 926 nucleotides),
1% differences in tub2 (465 nucleotides) and 7% differences in rpb2 (565 nucleotides). Based on
both morphological and molecular data the strains isolated in the present study were identified as a
new species. There is only one record of species of Chaetomium associated with Camellia flowers
(Watson 1950).
Figure 33 – Chaetomium camelliae (JZB340001 Ex-holotype). a Diseased leaf. b Upper view of
the colony on PDA after five days. c Reverse view of the colony on PDA after five days.
d ascomata on PDA. e–f Asci. g–h Ascospores. Scale bars: d = 1000 µm, e–f = 20 µm,
g, h = 10 µm.
Subclass Xylariomycetidae O.E. Erikss & Winka, Myconet 1: 12 (1997)
Amphisphaeriales D. Hawksw. & O.E. Erikss., Systema Ascomycetum 5: 177 (1986)
Notes – Currently there are 17 families and 88 genera in this order (Hyde et al. 2020b). For
recent taxonomic treatment we follow Hyde et al. (2020b).
472
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor & M.E. Barr., in Hyde, Fröhlich & Taylor,
Sydowia 50(1): 23 (1998)
Notes – Apiosporaceae was introduced by Hyde et al. (1998). After several years of
taxonomic conflicts, it is now accepted under Xylariales (Smith et al. 2003, Daranagama et al.
2018). The type genus of this family is Apiospora Sacc. Apiosporaceae species are endophytes
pathogens and saprobes on a wide range of hosts (Hyde et al. 1998).
Arthrinium Kunze., in Kunze & Schmidt, Mykologische Hefte (Leipzig) 1: 9 (1817)
Notes – Species in Arthrinium are found in a wide range of hosts as plant pathogens (Chen et
al. 2014), lichens (He & Zhang 2012) marine algae (Suryanarayanan 2012), soil (Singh et al. 2012)
and human pathogens (de Hoog et al. 2000). The current study identified one strain of Arthrinium
jiangxiense (Fig. 34).
Figure 34 – Phylogenetic tree generated by ML analysis of combined ITS, tub2 and tef1 sequence
data of Arthrinium species. Eighty strains are included in the analyses. The tree is rooted with
Nigrospora gorlenkoana (CBS 480.73). Tree topology of the ML analysis was similar to the BI.
The best scoring RAxML tree with a final likelihood value of –17931.476388 is presented. The
matrix had 1327 distinct alignment patterns, with 35.59% of undetermined characters or gaps.
Estimated base frequencies were as follows: A = 1.219297, C = 3.435344, G = 1.309116, T =
1.132730; substitution rates AC = 1.219297, AG = 3.435344, AT = 1.309116, CG = 1.132730, CT
= 4.469358, GT = 1.000000; gamma distribution shape parameter α = 1.241093. RAxML bootstrap
support values ≥50% and Bayesian posterior probabilities ≥0.95 (BYPP) are shown near the nodes.
The scale bar indicates 0.02 changes per site. Ex–type/ex–epitype strains are in bold and new
isolates recovered in this study are in red.
473
Figure 34 – Continued.
Arthrinium jiangxiense M. Wang & L. Cai., in Wang, Tan, Liu & Cai, MycoKeys 34(1): 14
(2018)
Fig. 35
Index Fungorum: IF824910; Facesoffungi number: FoF09395
Pathogenic or saprobic on dead Camellia sinensis leaves. Sexual morph: not observed.
Asexual morph: Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect,
scattered or aggregated in clusters on hyphae, hyaline to pale brown, smooth, ampulliform. Conidia
6–10 µm (x̄ = 8 µm, n = 40) diam., brown, smooth to finely roughened, granular, globose to
ellipsoid in surface view.
Culture characteristics – Colonies on PDA reaching 85 mm diam., in five days at 25°C.
Initially white and later become greyish–yellow, woolly, circular margin, with sparse aerial mycelia
reaching, hyphae hyaline, branched, septate.
474
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH3260001), and living culture JZB3260001.
Notes – In phylogenetic tree constructed using ITS, tub2 and tef1 sequences, two isolates from
the present study clustered together with the Arthrinium jiangxiense (LC4494) with 99% ML
bootstrap and 0.99 BYPP supports. Arthrinium jiangxiense was introduced in 2018 by Wang et al.
(2018). This species has been isolated from several different hosts including C. sinensis, Imperata
cylindrica, Machilus sp., Maesa sp., Phyllostachys sp. However, the status of the pathogenicity of
Arthrinium jiangxiense is understudied. In addition to the taxa identified in this study, there are
three Arthrinium species A. arundinis, A. camelliae–sinensis, and A. xenocordella associated with
C. sinensis (Farr & Rossman 2020).
Figure 35 – Arthrinium jiangxiense (JZB3260001). a Diseased leaf. d Upper view of the colony on
PDA after five days. e Reverse view of the colony on PDA after five days. d Conidiogenus cells
with conidia. e–g Conidia. Scale bars: d–f = 10 µm.
Nigrospora Zimm., in Centbl. Bakt. ParasitKde, Abt. I 8: 220 (1902)
Notes – This genus is a cosmopolitan fungal group that comprises endophytes, saprobes,
plant pathogens and opportunistic fungal pathogens in human (Wang et al. 2017a). Nigrospora
spores are one of the more dominant groups in the atmosphere (Wu et al. 2004). The present study
isolated and identified one strain that belongs to Nigrospora camellia–sinensis (Fig. 36).
Nigrospora camelliae–sinensis Mei Wang & L. Cai, in Wang, Liu, Crous & Cai, Persoonia 39: 127
(2017)
Fig. 37
Index Fungorum: IF820731; Facesoffungi number: FoF09396
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: not observed. Asexual
morph: Conidiophores, reduced to conidiogenous cells and aggregated in clusters on hyphae.
Conidiogenous cells hyaline to pale brown, globose to ampulliform, sometimes appearing as a
bulge directly from the mycelia without septa, Conidia 3–20 μm (x̄ = 16 μm, n = 40) diam., solitary
spherical, black, shiny, smooth, aseptate.
Culture characteristics – Colonies on PDA reach 80 mm diam. within five days at 25°C.
Initially white, later becoming grey, reverse black.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH3230016), and living culture JZB3230016.
Notes – Multi–locus phylogenetic analysis of ITS, tef1 and tub2 placed the isolate in the
present study together with Nigrospora camellia–sinensis supported by 86% ML bootstrap values
and 0.95 BYPP. The colony characters and morphology of the current species are similar to
N. camellia–sinensis (Wang et al. 2017a). So far eight species of Nigrospora have been reported on
Camellia sinensis and all those records are from Chinese tea plants (Farr & Rossman 2020).
475
Figure 36 – Phylogenetic tree generated by ML analysis of combined ITS, tub2 and tef1sequence data
of Nigrospora species. The tree is rooted with Arthrinium malaysianum (CBS 102053). Tree
topology of the ML analysis was similar to the BI. The best scoring RAxML tree with a final
likelihood value of –1480.382026 is presented. The matrix had 107 distinct alignment patterns,
with 2.77% of undetermined characters or gaps. Estimated base frequencies were as follows: A =
0.232659, C = 0.262146, G = 0.226065, T = 0.279129; substitution rates AC = 1.499597, AG =
0.544505, AT = 0.620143, CG = 0.815512, CT = 4.394781, GT = 1.000000; gamma distribution
shape parameter α = 0.641856. RAxML bootstrap support values ≥50% and Bayesian posterior
probabilities ≥0.95 (BYPP) are shown near the nodes. The scale bar indicates 0.02 changes per site.
Ex–type/ex–epitype strains are in bold and isolates recovered in the present study are in red.
476
Figure 37 – Nigrospora camellia–sinensis (JZB3230016) a Diseased leaf. d Upper view of the
colony on PDA after five days. e Reverse view of the colony on PDA after five days. d–f conidia.
Scale bars: d–f = 10 µm.
Sporocadaceae Corda, Icones fungorum hucusque cognitorum 5: 34 (1842)
Notes – Sporocadaceae consists of the pestalotioid fungi, which are typically appendaged
coelomycetes (Nag Raj 1993). They are characterised by multiseptate conidia with more or less
fusiform appendages at one or both ends. Many species belonging to this family are well–known
pathogens, but they can also be found as endophytes and saprobes (Maharachchikumbura et al.
2014).
Pestalotiopsis Steyaert, Bulletin du Jardin Botanique de l'État à Bruxelles 19 (3): 300 (1949)
Notes – Pestalotiopsis is a species–rich asexual genus with appendage bearing conidia
(Maharachchikumbura et al. 2013) that is widely distributed throughout tropical and temperate
regions. The species belong to this genus are well–known phytopathogens causing various diseases
in economically important crops (Maharachchikumbura et al. 2013, 2014). In the present study,
four Pestalotiopsis species were identified associated with leaf and shoot blights on Camellia
sinensis. (Fig. 38).
Pestalotiopsis camelliae Yan M. Zhang, Maharachch. & K.D. Hyde, in Zhang et al., Sydowia
64(2): 337 (2012)
Fig. 39
Index Fungorum: IF800980; Facesoffungi number: FoF09351
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Not observed. Asexual
morph: Conidiomata pycnidial on PDA, globose, scattered, semi–immersed, black, conidial masses
globose, black conidial masses. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells subcylindrical, hyaline, smooth, proliferating, Conidia 27–30 × 7–10 µm (x̅ = 30 × 9 µm, n =
40), fusoid, straight to slightly curved, 4 septate. Basal cell 4–7 µm (x̅ = 5 µm, n = 40), obconic,
hyaline, smooth, thin–walled, Median cells 20–22 µm (x̅ = 20.5 µm, n = 40), three, doliiform to
subcylindrical, walls thick verruculose, slightly constricted at the septa, concolourous, olivaceous,
septa and periclinal walls darker than the rest of the cell, Apical appendages three, tubular, arising
from the upper portion of the apical cell, various in length. Basal appendages not observed.
Culture characteristics – Colonies on PDA attaining up to 40 mm diam., after seven days at
25°C, with an undulate edge, whitish, with medium dense aerial mycelium on the surface with
black, gregarious conidiomata; reverse similar in colour.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH340062–3), and living cultures
JZB340062–3.
Notes – In the present study two Pestalotiopsis isolates obtained from tea leaves clustered
together with the Pestalotiopsis camelliae (MFLUCC 12–0277) type species with 58% ML
bootstrap and less than 0.90 BYPP. These isolates are similar to the ex–type isolate. Hence, we
identified two isolates from our study as Pestalotiopsis camelliae. This species was introduced by
Liu et al. (2017) from Camellia sinensis leaves in China. Pathogenicity of this species was proven
by (Wang et al. 2019b).
477
Figure 38 – Phylogenetic tree generated by ML analysis of combined ITS, tub2 and tef1 sequence data
of Pestalotiopsis species. Eighty strains are included in the analyses. Pseudopestalotiopsis
longiappendiculata (LC3013) and Pseudopestalotiopsis cocos (CBS27229) used as the out–group.
Tree topology of the ML analysis was similar to the BI. The best scoring RAxML tree with a final
likelihood value of – 24349.980578 is presented. The matrix had 1172 distinct alignment patterns,
with 9.91% of undetermined characters or gaps. Estimated base frequencies were as follows: A =
0.251668, C = 0.245757, G = 0.259668, T = 0.242908; substitution rates AC = 1.353890, AG =
478
4.605576, AT = 1.059439, CG = 0.801610, CT = 9.121730, GT = 1.000000; gamma distribution
shape parameter α = 0.944898. RAxML bootstrap support values ≥50% and Bayesian posterior
probabilities ≥0.95 (BYPP) are shown near the nodes. The scale bar indicates 0.05 changes per site.
Ex–type/ex–epitype strains are in bold and isolates recovered in this this study are in red.
Figure 38 – Continued.
479
Figure 39 – Pestalotiopsis camelliae (JZB340062). a Diseased leaf. b Upper view of the colony on
PDA after seven days. c Reverse view of the colony on PDA after seven days. d–e Pycnida on
PDA. f–g Conidiogenus cells with conidia. h–i Conidia. Scale bars: d, e = 100 µm. f–i = 10 µm.
Pestalotiopsis kenyana K.D. Hyde & Crous, in Maharachchikumbura et al., Studies in Mycology
79: 166 (2014)
Fig. 40
Index fungorum: IF809741; Facesoffungi number: FoF06981
Pathogenic or saprobic on Camellia sinensis leaves. Sexual morph: Not observed. Asexual
morph: Conidiomata pycnidial in culture on PDA, pycnidial globose, scattered, semi–immersed,
black, conidial masses black, globose, Conidiophores, reduced to conidiogenous cells.
Conidiogenous cells discrete, lageniform to subcylindrical, hyaline, smooth, proliferating 1–3 times
percurrently. Conidia 20 – 40 × 7–10 μm (x̅ = 25 × 8 μm, n = 40), fusoid, subcylindrical, straight to
slightly curved, 4–septate Basal cell conic to obconic, truncate base, hyaline, and thin–walled.
Median cells 15–20 μm (x̅ = 16 μm, n = 40), three, doliform, concolourous, brown, septa darker
than the rest of the cell. Apical appendages mostly 3 arising from the apical crest, unbranched,
filiform.
Culture characteristics – Colonies on PDA attaining 30–40 mm diam., after seven days at
25°C, with an undulate edge, whitish, medium dense aerial mycelium on the surface with black,
gregarious conidiomata. Reverse white.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried culture JZBH340062 and JZBH340063), and living
culture JZB340062– JZB340063.
Notes – In the present study, two species isolated from tea leaves developed a sister clade
with Pestalotiopsis kenyana (CBS 442.67 and OP068) with 81% ML bootstrap and 0.95 BYPP.
Based on phylogeny and morphology these isolates were identified as Pestalotiopsis kenyana. This
species was introduced by Maharachchikumbura et al. (2014) from a branch of Coffea sp. in Kenya.
Liu et al. (2016a) first reported this species from tea plants in China. There are no other hosts
reported for this species (Farr & Rossman 2020).
Pestalotiopsis lushanensis F. Liu & L. Cai, in Liu et al., Scientific Reports (2017)
Fig. 41
Index Fungorum: IF818919; Facesoffungi number: FoF09397
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: Not observed. Asexual
morph: Conidiomata pycnidial in culture on PDA, globose, aggregated or scattered, semi–
immersed, black, exuding conidial masses. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells discrete or integrated, ampulliform, clavate or subcylindrical, hyaline,
480
smooth–walled. Conidia 20–30 × 7–10 μm (x̅ = 20 × 8 μm, n = 40), fusoid, ellipsoid, straight to
slightly curved, 4 septate Basal cell obconic truncate base, hyaline, verruculose, thin–walled, 3.5–
6 μm long. Median cells 10–20 μm (x̅ = 15 μm, n = 40) three, doliiform, long, pale brown to brown,
septa darker than the rest of cell. Apical appendages 20–25 μm (x̅ = 20 μm, n = 40), 2–3 tubular,
unbranched, filiform, Basal appendage single, tubular, unbranched, filiform. Basal appendage
single, tubular, and unbranched.
Culture characteristics – Colonies on PDA attaining 30–40 mm diam., after seven days at
25°C, with undulate edge, whitish, with medium dense aerial mycelium on the surface with black,
gregarious conidiomata; reverse similar in colour.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoot, June 2015, H.L. Li (dried culture JZBH340059), and living culture JZB340059.
Notes – In the present study, a single isolate clustered together with the Pestalotiopsis
lushanensis with 94% ML and 0.99 BYPP values. According to the type species description given
by Liu et al. (2017), the current isolate is morphologically similar to P. lushanensis species. This
species was introduced by Liu et al. (2017) as a pestaloid species associated with Camellia sinensis
China.
Figure 40 – Pestalotiopsis kenyana (JZB340062) a Diseased leaf. b Upper view of the colony on
PDA after seven days. c Reverse view of the colony on PDA after seven days. c Pycnida on PDA.
d Conidia. Scale bars: c = 100 µm, c = 10 µm.
Figure 41 – Pestalotiopsis lushanensis (JZB340059). a Diseased shoot. b Upper view of the colony
on PDA after seven days. c Reverse view of the colony on PDA after seven days. d–e Pycnida on
PDA. f–g Conidia. Scale bars: d–e = 100 µm. f–g = 10 µm.
481
Pestalotiopsis rhodomyrtus Song, K. Geng, K.D. Hyde & Yong Wang bis, in Song et al. Phytotaxa
126(1): 27 (2013)
Fig. 42
Index Fungorum: IF804968; Facesoffungi number: FoF09398
Pathogenic or saprobic on Camellia sinensis shoots. Sexual morph: not observed. Asexual
morph: Conidiomata pycnidial in culture on PDA, globose, scattered, semi–immersed, conidial
mass black, Conidiophores reduced to conidiogenous cells. Conidiogenous cells discrete, hyaline,
filiform. Conidia 20–25 × 5–6 μm (x̅ = 24 × 5 μm, n = 40), fusoid, straight to slightly curved, 4–
septate. Basal cell 3–6 μm (x̅ = 5 μm, n = 40), conic, pale brown, smooth, thin–walled. Median
cells 12–20 μm (x̅ = 16 μm, n = 30) three, brown, thin septa, septa darker than cells, milled cell
dark brown than the other cells. Apical appendages 7.5–15 μm (x̅ = 11 μm, n = 30), three, tubular
unequal. Basal appendage one and filiform.
Culture characteristics – Colonies on PDA reaching 90 mm diam., after seven days at 28°C.
White mycelium, crenate edge, whitish, surface aerial mycelium, fruiting bodies start to appear
after 7 days, black, reverse of pinkish–white become black when old.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis shoots, June 2015, H.L. Li (dried culture JZBH340060), and living culture JZB340060.
Notes – Pestalotiopsis rhodomyrtus was previously isolated from Rhodomyrtus tomentosa in
China (Song et al. 2013). In the present study, a single strain obtained from a diseased tea shoot
clustered together with the P. rhodomyrtus (LC4458 type species) with 90% ML and 0.98 BYPP
values. The taxon identified in the present study is similar to the type specimen. This is the first
report of P. rhodomyrtus on Camellia sinensis (Farr & Rossman 2020).
Figure 42 – Pestalotiopsis rhodomyrtus (JZB340060). a Diseased shoot. b Upper view of the
colony on PDA after seven days. c Reverse view of the colony on PDA after seven days.
d–e Pycnida on PDA. f–h Conidia. Scale bars: d, e = 100 µm, f–h = 10 µm.
Pseudopestalotiopsis Maharachch., K.D. Hyde & Crous, in Maharachchikumbura et al., Studies in
Mycology 79: 180 (2014)
Notes – This genus was introduced by Maharachchikumbura et al. (2014) to accommodate
pestaloid species with dark concolourous median cells and knobbed apical appendages. Combined
gene phylogenetic analysis of ITS, tub2 and tef1, showed that taxa from current study belong to two
species. The phylogenetic placements of those taxa are given in Fig. 43.
482
Pseudopestalotiopsis camelliae–sinensis F. Liu & L. Cai in Liu et al., Scientific Reports 7(no.
866): 12 (2017)
Fig. 44
Index Fungorum: IF818924; Facesoffungi number: FoF09351
Pathogenic or saprobic on Camellia sinensis leaves and shoots. Sexual morph: not observed.
Asexual morph: Conidiomata pycnidial in culture on PDA, globose, scattered, semi–immersed,
black, exuding globose, dark brown to black conidial masses. Conidiophores not observed.
Conidiogenous cells 10–20 × 2–5 μm (x̅ 20 × 4 μm, n = 30), discrete, subcylindrical, hyaline,
smooth, proliferating 1–3 times percurrently. Conidia 20–30 × 7–10 μm (x̅ = 25 × 8 μm, n = 40),
fusoid, ellipsoid, straight, 4–septate, Basal cell conic, truncate base, hyaline, minutely verruculose
and thin. Median cells 15–20 μm (x̅ 16 μm, n = 40), three, doliiform, middle cell darker than the
other two. Apical appendages 8–20 (x̅ 15 μm, n = 40), three, arising from the apical crest,
unbranched, filiform. Basal appendages two, centric, tubular and flexuous.
Culture characteristics – Colonies on PDA attaining 30–40 mm diam., after seven days at
25°C, with an undulate edge, whitish, with medium dense aerial mycelium on the surface with
black, gregarious conidiomata; reverse similar in colour.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves and shoots, June 2015, H.L. Li (dried cultures JZBH340040–JZBH340054), and
living cultures JZB340040–JZB340054.
Notes – In the present study 14 strains isolated from diseased leaf and shoots samples
clustered with Pseudopestalotiopsis camelliae–sinensis. Morphologically both cultural and
structural characters such as conidial shape and dimensions of the isolated taxa were similar to the
type description of Pseudopestalotiopsis camelliae–sinensis (Liu et al. 2017). All isolates in the
present study share 98–100% nucleotide similarities at three gene regions. This species was
introduced by Liu et al. (2017) associated with Camellia sinensis in China. In addition, the only
other host reported so far is Vitis vinifera (Farr & Rossman 2020). Pseudopestalotiopsis camelliae–
sinensis was the most isolated species in the present study.
Pseudopestalotiopsis chinensis F. Liu & L. Cai Liu et al., Scientific Reports 7(no. 866): 12 (2017)
Fig. 45
Index Fungorum: IF818923; Facesoffungi number: FoF09399
Pathogenic or saprobic on Camellia sinensis. Sexual morph: not observed. Asexual morph:
Conidiomata pycnidial in culture on PDA, globose, scattered, semi–immersed, black, exuding
globose, dark brown to black conidial masses. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells discrete, lageniform to subcylindrical, hyaline, smooth, proliferating. Conidia
20–30 × 7–10 μm (x̅ = 25 × 8 μm, n = 40), fusoid, ellipsoid to subcylindrical, straight to slightly
curved, 4–septate, Pigmentation occurs while attached to the conidiogenous cell. Basal cell 15–20
μm (x̅ = 16 μm, n = 40). Median cells three, doliiform, wall verruculose concolourous, brown, septa
darker than the cells. Apical cell 4–6 μm long, hyaline, subcylindrical, rugose and thin–walled.
Apical appendages 2–3 tubular, initiate from the apical crest, unbranched, filiform. Basal
appendages two, centric, tubular, flexuous.
Culture characteristics – Colonies on PDA attaining 80–90 mm diam., after seven days at
25°C. Medium dense aerial mycelium, undulate, whitish, surface with black, gregarious
conidiomata. Reverse white and become darker with age.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH340055–JZBH340058), and living cultures
JZB340055–JZB340058.
Notes – In the present study, the four isolates clustered with the ex–type isolate of Ps.
chinensis (LC3011) with 81% ML and 0.95 BYPP values. These strains are similar to the type
species description of Pseudopestalotiopsis chinensis (Chen et al. 2018a). This species was
introduced by Chen et al. (2018a) from Camellia sinensis leaves. Other than Camellia sinensis
there are no other host records for this species (Farr & Roseman 2020).
483
Figure 43 – Phylogenetic tree generated by ML analysis of combined ITS, tub2 and tef1 sequence
data of Pseudopestalotiopsis species. The tree is rooted with Neopetalotiopsis clavispora
(MFLUCC 12–0277). Tree topology of the ML analysis was similar to the MP and BI. The best
scoring RAxML tree with a final likelihood value of – 24349.980578 is presented. The matrix had
354 distinct alignment patterns, with 12.95% of undetermined characters or gaps. Estimated base
frequencies were as follows: A = 0.251912, C = 0.252891, G = 0.233751, T = 0.261446;
substitution rates AC = 1.202821, AG = 5.554634, AT = 2.143706, CG = 1.053081, CT =
6.705276, GT = 1.000000; gamma distribution shape parameter α = 0.440726. RAxML bootstrap
support values ≥50% and Bayesian posterior probabilities ≥0.95 (BYPP) are shown near the nodes.
The scale bar indicates 0.03 changes per site. Ex–type/ex–epitype strains are in bold. New isolates
recovered in the present study are in red.
484
Figure 44 – Pseudopestalotiopsis camelliae–sinensis (JZBH340040). a Pycnidial wall with
developing conidiogenous cells and developing conidia. b–c Conidia. d–e Pycnidia on PDA.
f Upper view of culture on PDA after seven days. g Reverse view of culture on PDA after seven
days. Scale bars: a = 20 µm, b–c = 10 µm, d–e = 100 µm.
Figure 45 – Pseudopestalotiopsis chinensis (JZB340058). a Pycnidial wall with developing
conidiogenous cells and developing conidia. b Conidiogenous cells and developing conidia.
c Conidia. d–e Different shapes of pycnidia. f Front view of culture on PDA after seven days.
g Reverse view of culture on PDA after seven days. Scale bars: a = 20 µm, b–c = 10 µm, d–e = 100
µm.
485
Xylariales Nannf., Nova Acta Regiae Societatis Scientiarum Upsaliensis 8 (2): 66 (1932)
Notes – In recent taxonomic treatments by Hyde et al. (2020b) 15 families are accepted in
Xylariales; Barrmaeliaceae, Cainiaceae, Clypeosphaeriaceae, Coniocessiaceae, Diatrypaceae,
Graphostromataceae, Hansfordiaceae, Hypoxylaceae, Induratiaceae, Lopadostomataceae,
Microdochiaceae, Polystigmataceae, Requienellaceae, Xylariaceae and Zygosporiaceae with 160
genera (Hyde et al. 2020b).
Xylariaceae Tul. & C. Tul., Selecta Fungorum Carpologia, Tomus Secundus. Xylariei – Valsei –
Sphaeriei 2: 3 (1863)
Notes – Up to now 32 genera are accepted in Xylariaceae (Hyde et al. 2020b). Xylariaceae
species are saprobic, pathogenic, or endophytic on a wide range of hosts, some are important
producers of bioactive compounds and secondary metabolites (Stadler & Hellwig 2005, Helaly et
al. 2018).
Nemania Gray, A natural arrangement of British plants 1: 516 (1821)
Notes – Nemania consists of xylariaceous species that are more or less carbonaceous, dark
brown to black stromata that do not release coloured pigments in 10% potassium hydroxide (KOH)
(Ju & Rogers 2002). They are mostly reported as endophytes on different hosts. In the present
study, we isolated three strains belonging to Nemania diffiusa (Fig. 46).
Nemania diffusa (Sowerby) Gray, Nat. Arr. Brit. Pl. (London) 1: 517 (1821)
Fig. 47
Index Fungorum: IF477312; Facesoffungi number: FoF09400
Pathogenic or Saprobic on Camellia sinensis leaves. Sexual morph: not observed. Asexual
morph: Conidiomata pycnidial in culture on PDA, scattered or aggregated, irregular black.
Conidiophores not observed. Conidiogenus cells not observed. Conidia 8–10 × 3–4 µm (x̅ = 8 × 3
µm, n = 40), hyaline, ellipsoidal, guttulate, single germination tube.
Culture characteristics – Colonies on PDA, reaching 50 mm diam., after seven days at 28°C.
White fluffy mycelium, entire, smooth margin, reverse become dark brown with age.
Material examined – CHINA, Fujian Province, Zhangzhou County, on dead Camellia
sinensis leaves, June 2015, H.L. Li (dried cultures JZBH3370001– JZBH3370003), and living
cultures JZB3370001– JZB3370003.
Notes – In the present study we obtained three isolates belonging to Nemania. In the
phylogenetic analysis, these isolates clustered together with the Nemania diffusa (type strain
NC0608 and other representative strains) with 100% ML and 1.0 BYPP values. Based on
morphological and phylogenetic analyses we confirmed the isolates obtained in this study as
Nemania diffusa. This species has been reported in tea plantations causing soft rot in shoots in Sri
Lanka (Balasuriya & Adikaram 2008). This species has also been reported on Alnus glutinosa,
Betula sp., Fagus sp., Fraxinus sp., Metrosideros polymorpha, Nothofagus menziesii, Nothofagus
solandri, Nothofagus sp., Quercus robur and Ulmus suberosa (Farr & Rossman 2020). However,
this is the first report of Nemania diffusa in Chinese tea cultivations.
Discussion
This study revealed the diversity of fungi associated with diseased leaves and shoots of tea in
a plantation in China. The 110 isolates obtained comprised 32 species in 13 genera in 11 families.
Of these 32 species, five were determined to represent hitherto unknown species and thus were
introduced as new. In addition, nine new host records were reported. These taxa were associated
with typical symptoms of leaf necrosis and shoots blights on C. sinensis. Moreover, some of these
taxa belong to genera well–established as pathogenic on tea, namely Arthrinium, Botryosphaeria
(Jayawardena et al. 2016b) Colletotrichum (Liu et al. 2015), Diaporthe (Gao et al. 2016),
Pestalotiopsis, Pseudopestalotiopsis (Maharachchikumbura et al. 2013), Nigrospora and
Trichoderma (Dutta et al. 2015). However, this study reported several genera, Chaetomium
Epicoccum and Setophoma for which pathogenicity has not been confirmed on tea.
486
Figure 46 – Phylogenetic tree generated by ML analysis of combined ITS and rpb2 sequence data
of Nemania species. Podosordaria muli (WSP 167) and P. mexicana (WSP 176) were used as the
outgroup taxa. Tree topology of the ML analysis was similar to the MP. The best scoring RAxML
tree with a final likelihood value of – 24349.980578 is presented. The matrix had 1172 distinct
alignment patterns, with 9.91% of undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.251668, C = 0.245757, G = 0.259668, T = 0.242908; substitution rates AC
= 1.353890, AG = 4.605576, AT = 1.059439, CG = 0.801610, CT = 9.121730, GT = 1.000000;
gamma distribution shape parameter α = 0.944898. RAxML bootstrap support values ≥50% and
Bayesian posterior probabilities ≥0.95 (BYPP) are shown near the nodes. The scale bar indicates
0.02 changes per site. Ex–type/ex–epitype strains are in bold and isolates recovered in the present
study are in red.
487
Figure 47 – Nemania diffusa (JZBH3370001). a Diseased leaf. b Upper view of the colony on PDA
after seven days. c Reverse view of the colony on PDA after seven days. d–e Pycnidia on PDA.
f–i Conidia. j Germinating conidia. Scale bars: d, e = 100 μm, f–i = 10 μm.
In the present study we isolated five Botryosphaeria dothidea strains associated with shoot
blights on tea. Botryosphaeriaceae species are normally regarded as opportunistic pathogens. Even
though the exact underlying mechanism is unknown, it is thought that these fungi become
pathogenic when the environmental conditions are unfavourable for the host (Chethana et al. 2016,
Manawasinghe et al. 2016). In addition to Botryosphaeria dothidea, Lasiodiplodia theobromae and
L. pseudotheobromae have been reported causing leaf necrosis on Camellia sinensis in China (Li et
al. 2019). In comparison of disease symptoms caused by these Botryosphaeriaceae taxa, all species
induce brown lesions on young and mature leaves that become necrotic with age. However, in this
study we also isolated Botryosphaeria dothidea from dead shoots. Twig die–back caused by
Macrophoma theicola Petch, (Botryosphaeriaceae) is considered to be one of the major stem
diseases of C. sinensis (Mareeswaran et al. 2015). The disease symptoms associated with this
species are identical to the dieback caused by Botryosphaeria dothidea observed in the present
study (Mareeswaran et al. 2015). Furthermore, colony morphology and conidial characters of these
two species are quite similar (Phillips et al. 2013). Therefore, it is important to identify these
species using molecular data to avoid misidentifications. Furthermore, considering the confused
nature of Macrophoma (Sutton 1980) there is a need to re-collect and epitypify M. theicola to
establish its phylogenetic position. Other than opportunistic pathogens, in this study, we also
identified species belonging to well established phytopathogenic genera in Camellia sinensis.
Two Colletotrichum species were isolated from tea leaves with leaf necrosis symptoms. Up
until now, 24 Colletotrichum species have been associated with tea worldwide (Farr & Rossman
2020). From these, 17 species have been observed in China, namely: C. alienum, C. boninense,
C. camelliae, C. cliviae, C. fioriniae, C. fructicola, C. gloeosporioides, C. karstii, C. Siamense,
C. henanense and C. jiangxiense (Farr & Rossman 2020), C. acutatum (Chen et al. 2017b),
C. aenigma (Chen et al. 2019), C. endophyticum (Wang et al. 2016b), C. plurivorum (Damm et al.
2019), C. truncatum, C. wuxiense (Wang et al. 2016b). The study conducted by Chen et al. (2017b)
showed that C. camelliae is the most dominant taxon occurring on Camellia. In this study, we
isolated C. fructicola and C. camelliae associated with tea leaf necrosis. However, the isolation rate
of C. Camelliae was lower in this study compared to the other phytopathogenic genera such as
Diaporthe. This might be due to selective sampling. Our sampling area was limited to Fujian
488
province and the only cultivar was cv purple rose. In addition, within a small range our sampling
rate was high and the present study focused on different symptoms rather looking at particular
symptoms or specific genera or species.
The greatest numbers of species isolated in this study were in Diaporthe (ten of the 32
species). These includes three novel species and four new host records. So far 21 Diaporthe species
have been reported as associated with C. sinensis (Farr & Rossman, 2020). Among them,
D. amygdali, D. apiculata, D. compacta, D. discoidispora, D. eres, D. hongkongensis, D. oraccinii,
D. penetriteum, D. tectonigena (Gao et al. 2016), D. incompleta, D. masirevicii, D. ueckerae,
D. velutina, D. xishuangbanica (Gao et al. 2017) and Diaporthe nobilis (Li et al. 2017) have been
reported in China. Among this pathogenicity has been proven only for Diaporthe penetriteum
(Table 3). All species isolated in this study were associated with either shoot blight or leaf necrosis
on tea. Therefore, further studies are necessary to understand the pathogenicity of each species on
Camellia sinensis.
Grey blight of tea is one of the most destructive foliar diseases in tea worldwide including
China (Chen et al. 2017c, Wang et al. 2019c) and southern India (Joshi et al. 2009). The symptoms
associated with this disease are pale yellow–green leaf spots that initially are small, oval and
surrounded by a narrow yellow zone. With age the spots become brown or grey with concentric
rings and scattered, tiny black dots can be observed. When the disease becomes severe it can result
in defoliation (Chen et al. 2017c). This disease is caused by Pestalotiopsis–like species in many tea
cultivation regions including China (Chen et al. 2017c). There are 220 records of Pestalotiopsis
species associated with C. sinensis (Farr & Rossman 2020). In this study we observed symptoms on
leaves similar to grey blight. However, most of the taxa isolated in this study associated with shoot
blight appeared in necrotic regions on young leaves. These isolates belong to two species;
Pseudopestalotiopsis camelliae–sinensis and Pseudopestalotiopsis chinensis. Pseudopestalotiopsis
camelliae–sinensis is one of the main causal organisms associated with grey blight in China (Chen
et al. 2018a). Since Pseudopestalotiopsis camelliae–sinensis was the most isolated species from
diseased samples, it might be the prominent phytopathogenic species in Fujian tea plantations. In
addition to that, Ps. ampullacea and Ps. theae also have been reported on tea (Chen et al. 2018a).
Nigrospora camelliae-sinensis is the only Nigrospora species isolated in this study. The
pathogenicity of this species has not been confirmed on tea. Nigrospora includes well–known plant
pathogens on economically important crops, fruits and ornamentals (Wang et al. 2019b).
Nigrospora sphaerica has been reported causing leaf blight on C. sinensis in China (Liu et al.
2015). Apart from being plant pathogens, the species in this genus are important allergenic fungi
and some also produce useful natural by–products (Saha & Bhattacharya 2015, Chen et al. 2016).
In addition to this species, we isolated a single Arthrinium species associated with leaf necrosis.
Arthrinium species are widely distributed on a range of hosts as endophytes, pathogens or saprobes
(Hong et al. 2015). Moreover, they have been reported as the causal organisms of cutaneous
infections of humans (Crous et al. 2012). They are known to produce bioactive compounds as well
(Hong et al. 2015). Wang et al. (2018) identified Arthrinium camelliae–sinensis from tea plants.
However, the pathogenicity of Arthrinium camelliae–sinensis has not been confirmed.
Setophoma yingyisheniae is one of the Pleosporaceae species identified in this study. A recent
study conducted by Liu et al. (2019) introduced four new species belonging to Setophoma, namely
S. antiqua, S. longinqua, S. yingyisheniae and S. yunnanensis associated with leaf spots on tea from
seven provinces in China. Until now, S. yingyisheniae has been isolated from five provinces in
China (Liu et al. 2019). However, pathogenicity of this species is unknown. In the present study,
several genera were identified for the first time associated with Camellia sinensis. In addition, we
isolated one species belonging to Didymellaceae, Epicoccum layuense. So far three species
belonging to this genus (E. camelliae E. latusicollum and E. sorghinum) have been reported on tea
plants (Chen et al. 2017a). This is the first report of Epicoccum layuense associated with tea.
A novel species belonging to Chaetomium based on morphological characters and molecular
data was identified. Species in Chaetomium are not common on Camellia sinensis. The only record
489
of this host–fungus relationship is reported by Watson (1950) who did not mention the species name. In the present study, our isolates of this genus
represent a novel taxon Chaetomium camelliae. Therefore, this is the first report of Chaetomium camelliae on tea pants in China. In addition, three
Fusarium species were identified and these are novel host records on Camellia sinensis. Seven other Fusarium species have been reported on
C. sinensis (Farr & Rossman 2020).
By comparing the results of this study and the checklist, it is clear that Camellia sinensis supports a high diversity of fungal species. These
fungal communities might have different effects on the plants, most importantly to increase host fitness to tolerate biotic and abiotic stresses. In
addition, these taxa play different roles as endophytes, saprobes and pathogens, possibly interacting. Thus, it it is possible that a small ecosystem exists
within a single host in nature. In the present study we found that fungal species with potential biocontrol ability co–exist with pathogenetic taxa on tea
bushes. Some species belonging to Trichoderma have potential to attack or inhibit the growth of other fungi through their production of inhibitory
secondary metabolites (Degenkolb et al. 2008, Lopes et al. 2012). A recent study conducted by Del Frari et al. (2019) has shown the potential of
Epicoccum species to act against Esca disease on grapevines.. In this way they may be acting as natural biocontrol agents keeping the diseases under
natural control when the conditions are favourable for plant and fungus (De Silva et al. 2019). In almost all the tea–growing regions, blister blight,
horse–hair blight, and twig dieback/stem canker have become the most destructive diseases (Keith et al. 2006). Therefore, many plantation practices
focus of the control these pathogens often via addition of excessive amounts of fungicides. This might provide a chance for other species to develop
into more aggressive or pathogenic strains unnoticed. Human–mediated factors, such as application of the excessive amounts of fungicides together
with environmental changes, provide both challenging and opportunistic environments for pathogenic species. Since fungi have potential for rapid
adaptation, they might either switch their host or emerge as novel taxa (Manawasinghe et al. 2018). Therefore, it is important to understand the
diversity of fungi, the roles they play in this small ecosystem and their interactions with one another. This will provide new insights into the
development of new management strategies by enhancing the antagonists and thus suppress severity of the diseases.
Table 3 Checklist of fungi associated with Tea. The checklist includes species names, family, life modes, disease name (if any), locality and
references. The current name is used according to Index Fungorum (2020) and the classification follows Wijayawardene et al. (2020). Genera and
species are listed in alphabetical order.
Species
Acremoniella atra (Corda) Sacc.,
Alternaria alternata (Fr.) Keissl*
Alternaria sp.
Annulohypoxylon michelianum (Ces. & De
Not.) Y.M. Ju, J.D. Rogers & H.M. Hsieh
Athelia rolfsii (Curzi) C.C. Tu & Kimbr
Family
Incertae sedis
Ascomycota
Pleosporaceae
Hypoxylaceae
Atheliaceae
Life mode
Disease caused
locality
Japan
References
Kobayashi (2007)
S, P**
Leaf spots
China, India, Japan
Greece
Kenya
Tai (1979), Chakraborty et al. (2006),
Kobayashi (2007), Zhou et al. (2014),
Chen et al. (2018b), Farr & Rossman
(2020)
Pantidou (1973)
Nattrass (1961)
Japan, Malawi Taiwan
(China)
Kobayashi (2007) Farr & Rossman
(2020)
490
Table 3 Continued.
Disease caused
Species
Armillaria mellea (Vahl) P. Kumm
Family
Physalacriaceae
Life mode
N/A
locality
Japan, Kenya, Malawi
Malay, Peninsula,
Papua New Guinea,
Tanzania, Zimbabwe
Brazil, Kenya,
Zimbabwe
China
References
Thompson & Johnston (1953), Wiehe
(1953), Riley (1960), Nattrass (1961),
Whiteside (1966), Shaw (1984),
Kobayashi (2007)
Mendes et al. (1998), Perez Sierra et al.
(2003), Jimu et al. (2015)
Thangaraj et al. (2019)
Armillaria sp.
Physalacriaceae
OP, S
Arthrinium arundinis (Corda) Dyko & B.
Sutton*
Arthrinium camelliae–sinensis M. Wang, F.
Liu & L. Cai*
Arthrinium jiangxiense M. Wang & L. Cai*
Arthrinium xenocordella Crous *
Aschersonia eugeniae Koord.
Ascochyta sp.
Ascochyta theae Hara
Asterina theae W. Yamam.
Athelia rolfsii (Curzi) C.C. Tu & Kimbr
Beltrania rhombica Penz.,
Beltraniella japonica Matsush.
Bifusella camelliae C.L. Hou*
Botryosphaeria dothidea (Moug.: Fr.) Ces. &
De Not*
Apiosporaceae
P**
Apiosporaceae
S
N/A
China
S
S
N/A
N/A
P, S
P
Branch rot
Dieback**
China
China
India
Papua New Guinea
Japan
China
Papua New Guinea
Malaysia
Japan
China
Australia, China,
Japan
Wang et al. (2019b), Yan et al. (2019),
This study
Wang et al. (2019b), Yan et al. (2019)
Wang et al. (2019b)
Mathur (1979)
Farr & Rossman (2020)
Kobayashi (2007)
Tai (1979), Farr & Rossman (2020)
Thompson & Johnston (1953)
Johnston (1960), Heredia–Abarca (1994)
Matsushima (1975), Kobayashi (2007)
Hou (2000), Chen et al. (2011)
Cunnington et al. (2007), Kobayashi
(2007), Dissanayake et al. (2016),
Jayawardena et al. (2016b), Burgess et
al. (2019), This study
Farr & Rossman (2020)
Tai (1979)
Watson (1950), Kobayashi (2007)
Richardson (1990)
Kobayashi (2007)
Crous (2002), Lombard et al. (2014,
2016), Liu & Chen (2017), Wang et al.
(2019a)
Thompson & Johnston (1953), Tai
(1979), Crous (2002), Lombard et al.
(2016)
Crous (2002)
Apiosporaceae
Apiosporaceae
Clavicipitaceae
Didymellaceae
Didymellaceae
Asterinaceae
Atheliaceae
Beltraniaceae
Amphisphaeriaceae
Rhytismataceae
Botryosphaeriaceae
Botryosphaeria microspora Petch
Botryosphaeria sp.
Botrytis cinerea Pers
Botryotinia sp.
Byssosphaeria rhodomphala (Berk.) Cooke
Calonectria colhounii Peerally*
Botryosphaeriaceae
Botryosphaeriaceae
Sclerotiniaceae
Sclerotiniaceae
Melanommataceae
Nectriaceae
Calonectria indusiata (Seaver) Crous
Nectriaceae
China Germany, Sri
Lanka, Thailand
Calonectria kyotensis Terash
Nectriaceae
Mauritius, Sri Lanka
S
Sri Lanka
China
Japan, USA
India
Japan
Indonesia, Mauritius,
USA
491
Table 3 Continued.
Species
Calonectria reteaudii (Bugnic.) C. Booth
Calonectria spathiphylli El–Gholl, J.Y.
Uchida, Alfenas, T.S. Schub., Alfieri & A.R.
Chase*
Calonectria brassicae (Panwar & Bohra) L.
Lombard, M.J. Wingf. & Crous
Calycellina camelliae Dennis
Capnodium sp.
Cephaleuros sp.
Ceratobasidium sp.
Ceratocystis fimbriata Ellis & Halst*
Family
Nectriaceae
Nectriaceae
Life mode
Cercospora chaae Hara
Ceriporiopsis hypolateritia (Berk. ex Cooke)
Ryvarden
Chaetomium camelliae Jayaward., Manawas.,
X.H. Li, J.Y.Yan, & K. D. Hyde
Chaetothyrium javanicum (Zimm.) Boedijn
Chaetothyrium spinigerum (Höhn.)
W. Yamam
Chaetothyrium setosum (Zimm.) Hansf
Cladosporium herbarum (Pers.) Link*
Cladosporium sp.
Clonostachys rosea (Link) Schroers,
Samuels, Seifert & W. Gams
Clypeolella camelliae (Syd., P. Syd. & E.J.
Butler) Hansf
Colletotrichum acutatum J.H. Simmonds*
Mycosphaerellaceae
Phanerochaetaceae
Glomerellaceae
P**
Colletotrichum aenigma B.S. Weir & P.R.
Johnst*
Colletotrichum alienum B.S. Weir & P.R.
Johnst.
Colletotrichum boninense Moriwaki, Toy.
Sato & Tsukib
Glomerellaceae
P**
Glomerellaceae
OP**, S
Glomerellaceae
OP**, S
Disease caused
locality
Mauritius
Mauritius
References
Crous (2002)
Risede & Simoneau (2001), Crous (2002)
Nectriaceae
Mauritius
Crous (2002)
Pezizellaceae
Capnodiaceae
Trentepohliaceae
Ceratobasidiaceae
Ceratocystidaceae
Papua New Guinea
Fiji
Thailand
Japan
China
Shaw (1984)
Firman (1972), Dingley et al. (1981)
Giatgong (1980)
Kobayashi (2007)
Xu et al. (2019)
Japan
Thailand
Kobayashi (2007)
Thompson & Johnston (1953)
S
P**
Wilt and
Canker
Chaetomiaceae
S or P
China
This study
Chaetothyriaceae
Chaetothyriaceae
S
China, Taiwan
China
Tai (1979)
Tai (1979), Farr & Rossman (2020)
China
Japan, Korea
Thailand
Japan
Tai (1979), Farr & Rossman (2020)
Cho & Shin (2004), Kobayashi (2007)
Thompson & Johnston (1953)
Kobayashi (2007)
Thailand
Thompson & Johnston (1953)
Brown blight
China
Anthracnose
China
Arzanlou & Torbati (2013), Chen et al.
(2016, 2017b)
Jayawardena et al. (2016a), Wang et al.
(2016b), Chen et al. (2019)
Liu et al. (2015)
Chaetothyriaceae
Cladosporiaceae
Cladosporiaceae
Bionectriaceae
P
Englerulaceae
China
Anthracnose
New Zealand
Vieira et al. (2014), Hou et al. (2016), Liu
et al. (2016a), Chen et al. (2017b), Diao et
al. (2017), Douanla–Meli et al. (2018)
492
Table 3 Continued.
Species
Colletotrichum camelliae Massee*
Family
Glomerellaceae
Life mode
E, P**
Disease caused
Leaf spots
locality
China Jamaica,
Korea, Thailand, USA
Colletotrichum cliviicola Damm & Crous*
Glomerellaceae
S, P
Anthracnose
Brazil, China
Colletotrichum endophyticum Manamgoda,
Udayanga, L. Cai & K.D. Hyde*
Colletotrichum fioriniae R.G. Shivas & Y.P.
Tan*
Colletotrichum fructicola Prihast., L. Cai &
K.D. Hyde*
Glomerellaceae
OP, P**
China
References
Larter & Martyn (1943), Thompson &
Johnston (1953), Tai (1979), Alfieri et
al. (1984), Cho & Shin (2004), Alizadeh
et al. (2015), Liu et al. (2015),
Jayawardena, et al. (2016a), Wang et al.
(2016b), De Silva et al. (2017), Chen et
al. (2017b), This study
Liu et al. (2015), Jayawardena et al.
(2016a), Wang et al. (2016b)
Wang et al. (2016b)
Glomerellaceae
OP**, S
China
Liu et al. (2015)
Glomerellaceae
E, P**
China, Indonesia
Colletotrichum gigasporum Rakotonir. &
Munaut*
Colletotrichum gloeosporioides (Penz.) Penz.
& Sacc*
Glomerellaceae
E, P
Weir et al. (2012), Liu et al. (2015),
Jayawardena et al. (2016a), Wang et al.
(2016b), This study
Alizadeh et al. (2015)
Glomerellaceae
E, P**
Leaf spots
Brazil, China, Fiji,
Japan, Kenya, Korea,
Malaysia, Papua New
Guinea, Taiwan,
Tanzania, USA,
Zimbabwe
Colletotrichum henanense Liu & L. Cai*
Glomerellaceae
P**
Leaf spots
China
Colletotrichum jiangxiense F. Liu & L. Cai*
Glomerellaceae
E, P**
Leaf spots
China
Colletotrichum karsti You L. Yang, Zuo Y.
Liu, K.D. Hyde & L. Cai*
Colletotrichum plurivorum Damm, Alizadeh
& Toy. Sato*
Glomerellaceae
OP**, S
China
Riley (1960), Nattrass (1961), Whiteside
(1966), Turner (1971), Firman (1972),
Williams & Liu (1976), Tai (1979),
Dingley et al. (1981), Alfieri et al.
(1984), Shaw (1984), Mendes et al.
(1998), Cho & Shin (2004), Kobayashi
(2007), Liu et al. (2015), Chen et al.
(2017b)
Alizadeh et al. (2015), Liu et al. (2015),
Jayawardena et al. (2016a), Wang et al.
(2016b), De Silva et al. (2017)
Liu et al. (2015), Jayawardena et al.
(2016a)
Wang et al. (2016b)
Glomerellaceae
S, P
China
Damm et al. (2019)
Leaf spots
Iran
493
Table 3 Continued.
Species
Colletotrichum pseudomajus F. Liu, L. Cai,
Crous & Damm*
Family
Glomerellaceae
Life mode
S, P
Disease caused
locality
Taiwan
Colletotrichum siamense Prihast., L. Cai &
K.D. Hyde*
Colletotrichum sp.*
Glomerellaceae
E, P**
Leaf spots
China
Glomerellaceae
E, P, S
Colletotrichum truncatum (Schwein.) Andrus
& W.D. Moore*
Colletotrichum wuxiense Yu Chun Wang,
X.C. Wang & Y.J. Yang*
Coniothyrium sp.
Corticium sp.
Corynespora polyphragmia (Syd. & P. Syd.)
M.B. Ellis
Corallomycetella repens (Berk. & Broome)
Rossman & Samuels
Cryptomyces theae Sawada
Cylindrocladiella novae–zelandiae (Boesew.)
Boesew
Cylindrocladium peruvianum Bat., J.L.
Bezerra & M.P. Herrera
Cylindrocladiella parva (P.J. Anderson)
Boesew
Cylindrocarpon lichenicola (Massal.) D.
Hawksw
Cylindrocladium sp.
Clypeolella camelliae (Syd., P. Syd. & E.J.
Butler) Hansf *
Cytospora ceratosperma (Tode) G.C. Adams
& Rossman
Dematophora necatrix R. Hartig*
Glomerellaceae
P**
Glomerellaceae
OP**, S
Xylariaceae
P**
Diaporthe amygdali (Delacr.) Udayanga,
Crous & K.D. Hyde*
Diaporthaceae
E, S
China, India, Fiji,
Thailand
Anthracnose
China
China
References
Liu et al. (2014), Alizadeh et al. (2015),
Jayawardena et al. (2016a), Costa et al.
(2019)
Jayawardena et al. (2016a), Wang et al.
(2016b).
Giatgong (1980), Dingley et al. (1981),
Sharma et al. (2015), Chen et al.
(2017b), Liu et al. (2015).
Wang et al. (2016b)
Coniothyriaceae
Corticiaceae
Corynesporascaceae
Malawi
Papua New Guinea
Japan
Jayawardena et al. (2016a), Wang et al.
(2016b)
Corbett (1964)
Shaw (1984)
Kobayashi (2007)
Nectriaceae
Thailand
Thompson & Johnston (1953)
Rhytismataceae
Nectriaceae
Taiwan
New Zealand
Farr & Rossman (2020)
Crous et al. (2006)
Nectriaceae
USA
Alfieri et al. (1984), Crous (2002)
Nectriaceae
Malawi
Wiehe (1953)
Nectriaceae
Papua New Guinea
Shaw (1984)
Nectriaceae
Englerulaceae
Brazil, Japan
India, Java
Mendes et al. (1998), Kobayashi (2007).
Hosagoudar (2003)
Valsaceae
Japan
Kobayashi (2007)
China, Japan
Tai (1979), Kobayashi (2007), Sun et al.
(2008)
Gao et al. (2016)
White rot
China
494
Table 3 Continued.
Disease caused
Species
Diaporthe apiculata Y.H. Gao & L. Cai*
Family
Diaporthaceae
Life mode
E, S
locality
China
Diaporthe biguttulata F. Huang, K.D. Hyde
& Hong Y. Li
Diaporthe compacta Y.H. Gao & L. Cai*
Diaporthaceae
S or P
China
Diaporthaceae
E
China
Diaporthe discoidispora F. Huang, K.D.
Hyde & Hong Y. Li*
Diaporthe eres Nitschke*
Diaporthaceae
E
China
Diaporthaceae
E, P
China, Japan
Diaporthe eucalyptorum Crous & R.G.
Shivas.
Diaporthe foeniculacea Niessl*
Diaporthaceae
S or P
China
Diaporthe fujianensis Jayaward., Manawas.,
X.H. Li, J.Y.Yan, & K. D. Hyde
Diaporthe fusiformis Jayaward., Manawas.,
X.H. Li, J.Y.Yan, & K. D. Hyde
Diaporthe hongkongensis R.R. Gomes,
Glienke & Crous*
Diaporthe incompleta Y.H. Gao & L. Cai*
Diaporthe masirevicii R.G. Shivas, L. Morin,
S.M. Thomps. & Y.P. Tan*
Diaporthe nobilis Sacc. & Speg*
Diaporthe oraccinii Y.H. Gao, F. Liu & L.
Cai*
Diaporthaceae
S or P
China
Udayanga et al. (2012), Gomes et al.
(2013), Chen et al. (2014), Lombard et
al. (2014)
This study
Diaporthaceae
S or P
China
This study
China
Gao et al. (2016), Dissanayake et al.
(2017a)
Gao et al. (2017), Yang et al. (2018b).
Gao et al. (2017)
Diaporthe penetriteum Y.H. Gao & L. Cai*
Italy
Diaporthaceae
Diaporthaceae
Diaporthaceae
Diaporthaceae
S
S
China
Diaporthaceae
Diaporthaceae
P, S
P, S
China
China
Diaporthaceae
E, P**
China
References
Du et al. (2016), Gao et al. (2016), Yang
et al. (2017), Gao et al. (2017), Yang et
al. (2018), Dissanayake et al. (2017a),
Yang et al. (2017), Fan et al. (2018)
This study
Yang et al. (2015), Dissanayake et al.
(2017a, b), Gao et al. (2017), Yang et al.
(2018)
Gao et al. (2016), Dissanayake et al.
(2017a), Gao et al. (2017)
Kobayashi (2007), Gao et al. (2016),
Dissanayake et al. (2017a), Gao et al.
(2017)
This study
Li et al. (2017)
Du et al. (2016), Gao et al. (2016, 2017),
Yang et al. (2017), Dissanayake et al.
(2017a, b), Yang et al. (2017, 2018a, b)
Du et al. (2016), Dissanayake et al.
(2017 a, b), Gao et al. (2017), Yang et
al. (2017, 2018)
495
Table 3 Continued.
Species
Diaporthe sackstonii R.G. Shivas, S.M.
Thomps. & Y.P. Tan
Diaporthe sennae C.M. Tian & Qin Yang
Diaporthe sinensis Jayaward., Manawas.,
X.H. Li, J.Y.Yan, & K. D. Hyde
Diaporthe sp*
Family
Diaporthaceae
Life mode
S or P
Diaporthaceae
Diaporthaceae
Disease caused
locality
China
References
This study
S or P
S or P
China
China
This study
This study
Diaporthaceae
E, S, P
Diaporthe tectonigena Doilom, Dissan. &
K.D. Hyde*
Diaporthe ueckeri Udayanga & Castl. *
Diaporthaceae
S
China, India, Papua
New Guinea
China
Gao et al. (2017), Mathur (1979), Farr &
Rossman (2020)
Gao et al. (2017)
Diaporthaceae
S
China
Diaporthe unshiuensis F. Huang, K.D. Hyde
& Hong Y. Li
Diaporthe velutina Y.H. Gao & L. Cai*
Diaporthe viniferae Dissanayake, X.H. Li &
K.D. Hyde
Diaporthe xishuangbanica Y.H. Gao & L.
Cai*
Diaporthe theae (Petch) Rossman &
Udayanga
Diatrype conferta Petch
Diatrype falcata (Syd. & P. Syd.) Sacc
Diatrype stigma (Hoffm.) Fr
Diatrype theae Hara
Dictyochaeta assamica (Agnihothr.) Aramb.,
Cabello & Mengasc
Dimeriellopsis theicola Sawada & W.
Yamam
Dimerina nantoensis (Sawada) W. Yamam
Dinemasporium neottiosporoides
(Agnihothr.) W.P. Wu
Discosia artocreas (Tode) Fr.
Discosia strobilina Lib
Discula theae–sinensis (I. Miyake) Moriwaki
& Toy*
Discosiella longiciliata Agnihothr
Diaporthaceae
S or P
China
Gao et al. (2016), Dissanayake et al.
(2017a), Gao et al. (2017)
This study
Diaporthaceae
Diaporthaceae
S
S or P
China
China
Gao et al. (2017)
This study
Diaporthaceae
S
China
Diaporthaceae
S
Diatrypaceae
Diatrypaceae
Diatrypaceae
Diatrypaceae
Chaetosphaeriaceae
Japan, Papua New
Guinea, Tanzania
Sri Lanka
Japan
Japan
Japan
New Zealand
Dissanayake et al. (2017a), Gao et al.
(2017), Yang et al. (2018a)
Ebbels & Allen (1979), Kobayashi
(2007)
Rappaz (1987)
Kobayashi (2007)
Kobayashi (2007)
Kobayashi (2007)
Hughes & Kendrick (1968)
Pseudoperisporiaceae
China
Tai (1979)
Valsariaceae
Xylariomycetidae
China
India
Farr & Rossman (2020)
Duan et al. (2007)
Discosiaceae
Discosiaceae
Gnomoniaceae
Southeastern states
Japan
Japan, China
Watson (1950)
Kobayashi (2007)
Tai (1979), Kobayashi (2007), Moriwaki
& Sato (2009)
Mathur (1979), Nag Raj (1993)
Ascomycota
P** S
Anthracnose
India
496
Table 3 Continued.
Species
Dyfrolomyces sinensis Samarak., Tennakoon
& K.D. Hyde*
Elsinoe theae Bitanc. & Jenkins*
Family
Dyfrolomycetaceae
Life mode
P, S
Elsinoaceae
Epicoccum camelliae Qian Chen, Crous & L.
Cai*
Epicoccum latusicollum Qian Chen, Crous &
L. Cai*
Epicoccum layuense
Epicoccum sorghinum (Sacc.) Aveskamp,
Gruyter & Verkley*
Erythricium salmonicolor (Berk. & Broome)
Burds
Exobasidium camelliae Shirai*
Exobasidium reticulatum S. Ito & Sawada*
Disease caused
locality
Thailand
References
Hyde et al. (2018)
P, S
Brazil, Japan, Korea
Tanzania
Didymellaceae
P, S
China
Didymellaceae
P, S
China
Didymellaceae
Didymellaceae
S or P
P, S
China
China
Riley (1960), Mendes et al. (1998), Cho
& Shin (2004), Kobayashi (2007), Fan et
al. (2018)
Chen et al. (2017a), Valenzuela-Lopez
et al. (2018)
Chen et al. (2017a), Valenzuela-Lopez
et al. (2018)
This study
Chen et al. (2017a), Bao et al. (2019)
Exobasidiaceae
Exobasidiaceae
P
P
Japan, Papua New
Guinea, Thailand
China, USA
China, Japan
Exobasidium vexans Massee*
Exobasidiaceae
P**
Exobasidium yunnanense Zhen Ying Li & L.
Guo
Fusarium asiaticum O’Donnell, T. Aoki,
Kistler & Geiser
Fusarium concentricum Nirenberg &
O’Donnell
Fusarium fujikuroi Nirenberg
Fusarium oxysporum Schltdl
Exobasidiaceae
P, S
China
Thompson & Johnston (1953),
Kobayashi (2007)
Alfieri et al. (1984)
Tai (1979), Chen (2002), Kobayashi
(2007)
Thompson & Johnston (1953), Tai
(1979), Giatgong (1980), Richardson
(1990), Chen (2002), Cho & Shin
(2004), Kobayashi (2007), Thaung
(2007), Silva et al. (2015).
Li & Guo (2009)
Nectriaceae
S or P
China
This study
Nectriaceae
S or P
China
This study
Nectriaceae
Nectriaceae
S or P
S
Fusarium proliferatum (Matsush.) Nirenberg
ex Gerlach & Nirenberg
Fusarium sp.
Nectriaceae
S or P
China
India, Kenya,
Southeast Asia
China
This study
Nattrass (1961), Sarbhoy & Agarwal
(1990), Lombard et al. (2019)
This study
Fusicladium theae Hara
Venturiaceae
Malaysia, Papua New
Guine, Sri Lanka
China, Japan
Liu (1977), Sinniah et al. (2017), Aoki et
al. (2018), Na et al. (2018)
Tai (1979), Kobayashi (2007).
Corticiaceae
Nectriaceae
Blister blight
China, India, Japan,
Korea, Myanmar,
Korea, Thailand
497
Table 3 Continued.
Species
Gliocladiopsis tenuis (Bugnic.) Crous & M.J.
Wingf.
Gliocladiopsis tenuis (Bugnic.) Crous & M.J.
Wingf.
Globisporangium debaryanum (R. Hesse)
Uzuhashi, Tojo & Kakish
Globisporangium mamillatum (Meurs)
Uzuhashi, Tojo & Kakish
Globisporangium spinosum (Sawada)
Uzuhashi, Tojo & Kakish
Gnomoniopsis fructicola (G. Arnaud)
Sogonov
Graphium rigidum (Pers.) Sacc
Guignardia abeana W. Yamam. & K. Konno
Guignardia theae (Racib.) C. Bernard
Helicobasidium longisporum Wakef
Helicobasidium purpureum (Tul.) Pat
Helicobasidium sp.
Hendersonia theae Hara
Family
Nectriaceae
Hypohelion durum Y.R. Lin, C.L. Hou & S.J.
Wang
Hypoxylon howeanum Peck
Rhytismataceae
Hypoxylon fuscopurpureum (Schwein.) M.A.
Curtis
Ilyonectria destructans (Zinssm.) Rossman,
L. Lombard & Crous
Julella vitrispora (Cooke & Harkn.) M.E.
Barr
Lasiodiplodia gonubiensis Pavlic, Slippers &
M.J. Wingf*
Lasiodiplodia pseudotheobromae A.J.L.
Phillips, A. Alves & Crous*
Lasiodiplodia theobromae (Pat.) Griffon &
Maubl*
Life mode
Disease caused
locality
Mauritius
References
Crous (2002)
Nectriaceae
Japan
Kobayashi (2007)
Pythiaceae
Philippines
Teodoro (1937)
Pythiaceae
Greece
Pantidou (1973)
Pythiaceae
Japan
Kobayashi (2007)
Gnomoniaceae
Malaysia
Liu (1977)
Microascaceae
Phyllostictaceae
Phyllostictaceae
Helicobasidiaceae
Helicobasidiaceae
Helicobasidiaceae
Phaeosphaeriaceae
Japan
Japan
China
Indonesia, China
Japan
Malawi
China, Japan, India
Matsushima (1975), Kobayashi (2007)
Kobayashi (2007)
Tai (1979)
Whiteside (1966)
Kobayashi (2007)
Wiehe (1953)
Mathur (1979), Tai (1979), Kobayashi
(2007)
Lin et al. (2004), Chen et al. (2011)
P, S
Branch rot
China
Hypoxylaceae
Japan
Hypoxylaceae
Japan
Kobayashi (2007) as Hypoxylon
coccinellu Sacc.
Kobayashi (2007)
Hypocreales
Japan
Kobayashi (2007)
Thelenellaceae
Japan
Kobayashi (2007)
Australia
Tan et al. (2019), Burgess et al. (2019)
Botryosphaeriaceae
E, P
Botryosphaeriaceae
P**
Leaf necrosis
China
Li et al. (2019)
Botryosphaeriaceae
P**
Leaf necrosis
China, Malawi,
Malaysia, Papua New
Wiehe (1953), Turner (1971), Liu
(1977), Shaw (1984), Whiteside (1966),
498
Table 3 Continued.
Species
Family
Life mode
Leptosphaerulina sp.
Lophodermium sinens Y.R. Lin, C.L. Hou &
Jiang L. Chen
Macrophoma sp.
Macrophoma theicola Petch
Macrophomina phaseolina (Tassi) Goid
Didymellaceae
Rhytismataceae
P
Marasmiellus scandens (Massee) Dennis &
D.A. Reid
Marasmius crinis–equi F. Muell. ex Kalchbr
Disease caused
locality
Guinea, Tanzania.
Zimbabwe
Malawi
China
References
Li et al. (2019)
Omphalotaceae
Thailand
Malawi
India, Malawi,
Malaysia, Tanzania
Malaysia
Thompson & Johnston (1953)
Wiehe (1953)
Wiehe (1953), Johnston (1960),
Riley (1960), Mathur (1979)
Turner (1971)
Marasmiaceae
Fiji, Japan
Marasmius sp.
Marasmius tenuissimus (Sacc.) Singer
Marssonina sp.
Meliola camelliae (Catt.) Sacc
Microcera coccophila Desm
Monilochaetes camelliae (Alcorn & Sivan.)
Réblová, W. Gams & Seifert
Mycosphaerella ikedae Hara
Marasmiaceae
Marasmiaceae
Dermateaceae
Meliolaceae
Nectriaceae
Australiascaceae
Thailand
Japan
Thailand
China
Papua New Guinea
Australia
Firman (1972), Dingley et al. (1981),
Kobayashi (2007)
Thompson & Johnston (1953)
Kobayashi (2007).
Giatgong (1980)
Tai (1979)
Shaw (1984)
Réblová et al. (2011)
Mycosphaerellaceae
Mycosphaerella sp.
Mycosphaerella theae Hara
Mycosphaerellaceae
Myriangium duriaei Mont. & Berk
Nectria bolbophylli Henn
Nectria cinnabarina (Tode) Fr
Nectria diversispora Petch
Nectria sp.
Myriangiaceae
Nectriaceae
Nectriaceae
Nectriaceae
Nectriaceae
Nectria pseudotrichia Berk. & M.A. Curtis
Nectriaceae
Nectricladiella viticola (Berk. & M.A.
Curtis) Hirooka, Rossman & P. Chaverri
Nectriaceae
China, Japan,
Malaysia
Mauritius, Tanzania
China, Japan, Samoa,
Zimbabwe
Japan
Japan
Japan
Taiwan
Malawi, Papua New
Guinea
Papua New Guinea,
Tanzania, Thailand
India
Botryosphaeriaceae
Botryosphaeriaceae
Botryosphaeriaceae
Corbett (1964)
Chen et al. (2011)
Johnston (1960), Tai (1979), Kobayashi
(2007)
Riley (1960), Orieux & Felix (1968)
Whiteside (1966), Tai (1979), Dingley et
al. (1981), Kobayashi (2007)
Kobayashi (2007)
Kobayashi (2007)
Kobayashi (2007)
Farr & Rossman (2020)
Wiehe (1953), Shaw (1984)
Thompson & Johnston (1953), Riley
(1960), Shaw (1984)
Crous (2002)
499
Table 3 Continued.
Family
Nectriaceae
Neocosmospora sp.*
Neocosmospora haematococca (Berk. &
Broome) Samuels, Nalim & Geiser
Neocosmospora ipomoeae (Halst.) L.
Lombard & Crous
Neocosmospora solani (Mart.) L. Lombard &
Crous
Neocapnodium tanakae (Shirai & Hara) W.
Yamam
Neofusicoccum ribis (Slippers, Crous & M.J.
Wingf.) Crous, Slippers & A.J.L. Phillips
Pyrrhoderma noxium (Corner) L.W. Zhou &
Y.C. Dai
Neocosmospora ambrosia (Gadd & Loos) L.
Lombard & Crous*
Neonectria ditissima (Tul. & C. Tul.)
Samuels & Rossman
Neopestalotiopsis clavispora (G.F. Atk.)
Maharachch., K.D. Hyde & Crous*
Neopestalotiopsis clavispora as
Pestalotiopsis clavispora (G.F. Atk.)
Steyaert*
Neopestalotiopsis ellipsospora (Maharachch.
& K.D. Hyde) Maharachch., K.D. Hyde &
Crous*
Neopestalotiopsis sp.*
Nectriaceae
Nectriaceae
Sri Lanka
Japan
References
Lombard et al. (2014), Guarnaccia &
Crous (2018), Sandoval–Denis et al.
(2018, 2019)
Sandoval–Denis et al. (2019)
Kobayashi (2007)
Nectriaceae
China
Tai (1979)
Nectriaceae
India, Japan
Trichomeriaceae
China
Sarbhoy & Agarwal (1990), Kobayashi
(2007), Aoki et al. (2018)
Tai (1979)
Botryosphaeriaceae
Malawi
Wiehe (1953)
China, Thailand
Thompson & Johnston (1953), Riley
(1960)
Freeman et al. (2013), Aoki et al.
(2018), Na et al. (2018)
Kobayashi (2007)
Nemania diffusa (Sowerby) Gray
Nigrospora camelliae–sinensis Mei Wang &
L. Cai*
Nigrospora chinensis Mei Wang & L. Cai*
Nigrospora guilinensis Mei Wang & L. Cai*
Nectriaceae
Life mode
P, S
Disease caused
Species
Neocosmospora ambrosia Gadd & Loos) L.
Lombard & Crous*
P, S
locality
India, Sri Lanka
India, Sri Lanka
Japan
Nectriaceae
Sporocadaceae
P**
Grey blight
China
Wei et al. (2005, 2007), Ge et al. (2009),
Wang et al. (2017b), Chen et al. (2018a)
Wei et al. (2005, 2007), Ge et al. (2009),
Wang et al. (2017b), Chen et al. (2018a)
Sporocadaceae
E, P**
Brown–black
spot
Sporocadaceae
P**
Grey blight
China
Wang et al. (2019b)
Sporocadaceae
P**, S
Grey blight
China, France
China
China
Maharachchikumbura et al. (2014),
Chen et al. (2018a)
This study
Wang et al. (2017b), This study
Xylariaceae
Apiosporaceae
S or P
P, S
Apiosporaceae
Apiosporaceae
P, S
P, S
China
China
Wang et al. (2017b)
Wang et al. (2017b)
500
Table 3 Continued.
Species
Nigrospora
Cai*
Nigrospora
Nigrospora
Nigrospora
lacticolonia Mei Wang & L.
Family
Apiosporaceae
Life mode
P, S
musae McLennan & Hoëtte*
oryzae (Berk. & Broome) Petch*
sphaerica (Sacc.) E.W. Mason*
Apiosporaceae
Apiosporaceae
Apiosporaceae
P, S
P, S
P**, S
Nigrospora pyriformis Mei Wang & L. Cai*
Nigrospora sp.*
Ophioirenina theae Sawada & W. Yamam*
Ophiognomonia setacea (Pers.) Sogonov
Ophiovalsa theae (Hara) Tak. Kobay
Paraconiothyrium fuckelii (Sacc.) Verkley &
Gruyter
Paraconiothyrium fuckelii (Sacc.) Verkley &
Gruyter*
Penicillium corylophilum Dierckx
Pestalotiopsis acaciae (Thüm.) K. Yokoy. &
S. Kaneko*
Pestalotiopsis aggestorum F. Liu & L. Cai*
Pestalotiopsis algeriensis (Sacc. & Berl.)
W.P. Wu*
Pestalotiopsis camelliae Yan M. Zhang,
Maharachch. & K.D. Hyde*
Apiosporaceae
Apiosporaceae
Meliolaceae
Gnomoniaceae
Gnomoniaceae
Didymosphaeriaceae
P, S
P, S
P, S
Pestalotiopsis chamaeropis Maharachch.,
K.D. Hyde & Crous*
Pestalotiopsis dilucida F. Liu & L. Cai*
Pestalotiopsis disseminata (Thüm.) Steyaert*
Pestalotiopsis funerea (Desm.) Steyaert*
Pestalotiopsis furcata Maharachch. & K.D.
Hyde*
Disease caused
locality
China
References
Wang et al. (2017b)
China
China
China, India
China
China
China, Taiwan
Japan
Japan
Japan
Wang et al. (2017b)
Wang et al. (2017b)
Dutta et al. (2015), Liu et al. (2016b).
Wang et al. (2017b)
Wang et al. (2017b)
Wang et al. (2017b)
Tai (1979), Hongsanan et al. (2015)
Kobayashi (2007)
Kobayashi (2007)
Kobayashi (2007)
Didymosphaeriaceae
Japan
Kobayashi (2007)
Aspergillaceae
Sporocadaceae
S
Kenya
China
Nattrass (1961)
Ge et al. (2009)
Sporocadaceae
Sporocadaceae
S
S
China
China
Liu et al. (2017)
Zhang et al. (2012)
Sporocadaceae
P**, S
Grey blight
China, Turkey
Sporocadaceae
P, S
Grey blight
China
Maharachchikumbura et al. (2014),
Moslemi & Taylor (2015), Chen et al.
(2017c), Liu et al. (2017), Wang et al.
(2017b), Solarte et al. (2017), This study
Liu et al. (2017), Wang et al. (2019b)
Sporocadaceae
Sporocadaceae
Sporocadaceae
Sporocadaceae
E, P, S
P, S
S
S
leaf blight
China
China
China
China, Thailand
Liu et al. (2017)
Zhang et al. (2012)
Ge et al. (2009)
Zhang et al. (2012),
Maharachchikumbura et al. (2013),
2014), Liu et al. (2017), Chen et al.
(2018a), Solarte et al. (2017)
501
Table 3 Continued.
Species
Pestalotiopsis gigas Steyaert
Pestalotiopsis jinchanghensis F. Liu & L.
Cai*
Pestalotiopsis kenyana Maharachch., K.D.
Hyde & Crous*
Pestalotiopsis longiappendiculata F. Liu &
L. Cai*
Pestalotiopsis longiseta (Speg.) K. Dai &
Tak. Kobay*
Pestalotiopsis lushanensis F. Liu & L. Cai *
Pestalotiopsis maculans (Corda) Nag Raj*
Family
Sporocadaceae
Sporocadaceae
Life mode
Disease caused
E, P, S
locality
Kenya
China
References
Nattrass (1961)
Liu et al. (2017)
Sporocadaceae
E, P, S
China
Liu et al. (2017), This study
Sporocadaceae
E, P, S
China
Liu et al. (2017)
Sporocadaceae
P, S
Japan, Korea
Kobayashi (2007)
Sporocadaceae
Sporocadaceae
P**
P, S
Pestalotiopsis menezesiana (Bres. &
Torrend) Bissett*
Pestalotiopsis microspora (Speg.) Bat. &
Peres*
Pestalotiopsis nattrassii Steyaert*
Sporocadaceae
P, S
China
China
Czechoslovakia,
France, Germany,
Japan, USA
China
Chen et al. (2018c), This study
Nag Raj (1993), Jeewon et al. (2002,
2003), Kobayashi (2007), Ge et al.
(2009), Maharachchikumbura et al.
(2011)
Zhang et al. (2012)
Sporocadaceae
E, P
China
Pestalotiopsis neglecta (Thüm.) Steyaert*
Pestalotiopsis palmarum (Cooke) Steyaert
Pestalotiopsis photiniae (Thüm.) Y.X. Chen*
Pestalotiopsis rhodomyrtus Yu Song, K.
Geng, K.D. Hyde & Yong Wang bis*
Pestalotiopsis sp.*
Sporocadaceae
Sporocadaceae
Sporocadaceae
Sporocadaceae
E, S
Sporocadaceae
E, P, S
Pestalotiopsis sydowiana (Bres.) B. Sutton*
Pestalotiopsis trachycarpicolaYan M. Zhang
& K.D. Hyde*
Pestalotiopsis versicolor (Speg.) Steyaert*
Pestalotiopsis virgatula (Kleb.) Steyaert*
Pestalotiopsis yanglingensis F. Liu & L. Cai*
Phacidium lauri (Sowerby) Crous & D.
Hawksw
Sporocadaceae
Sporocadaceae
P, S
China, Fiji, Papua
New Guinea, Samoa,
Thailand
China
China
Wei et al. (2005, 2007), Ge et al. (2009),
Zhang et al. (2012)
Nattrass (1961), Lu et al. (2000),
Zhuang (2001)
Wei et al. (2005, 2007), Ge et al. (2009)
Kobayashi (2007)
Tejesvi et al. (2009)
Liu et al. (2017), Wang et al. (2019b),
This study
Firman (1972), Giatgong (1980),
Dingley et al. (1981), Zhang et al.
(2012)
Zhuang (2001), Ge et al. (2009)
Liu et al. (2017)
China
China
China
Japan
Zhang et al. (2012)
Zhang et al. (2012)
Liu et al. (2017)
Ando et al. (1989), Kobayashi (2007)
Grey blight
China, Kenya
Sporocadaceae
Sporocadaceae
Sporocadaceae
Sporocadaceae
Phacidiaceae
China
Japan, Taiwan (China)
China
China
E
P, S
P, S
P, S
P**, S
Grey blight
502
Table 3 Continued.
Life mode
Disease caused
P**
Leaf spot
P, S
P**
Leaf spot
Species
Phaeodothis winteri (Niessl) Aptroot
Phaeoisaria clematidis (Fuckel) S. Hughes
Phaeosphaerella theae Petch
Phoma herbarum Westend*
Phoma sp.
Phyllosticta camelliae Westend*
Phyllosticta capitalensis Henn*
Phyllosticta citricarpa (McAlpine)
Phyllosticta erratica Ellis & Everh
Phyllosticta sp*
Family
Didymosphaeriaceae
Diatrypaceae
Venturiaceae
Didymellaceae
Didymellaceae
Phyllostictaceae
Phyllostictaceae
Phyllosticta theae Speschnew*
Phyllostictaceae
China, Fiji, Japan,
Tanzania, Thailand
Phyllosticta theicola Curzi
Pleospora theae Speschnew
Pseudocercospora camelliae (Deighton) U.
Braun*
Pseudocercospora camelliicola U. Braun &
C.F. Hill*
Pseudocercospora javanica Deighton
Phyllostictaceae
Pleosporaceae
Mycosphaerellaceae
P, S
Mycosphaerellaceae
P, S
Mycosphaerellaceae
P, S
Pseudocercospora ocellata (Deighton)
Deighton
Mycosphaerellaceae
P, S
Pseudocercospora theae (Cavara) Deighton
Mycosphaerellaceae
P**, S
Pestalotiopsidaceae
E, P, S
China, Japan
Japan
Georgia, New
Zealand
Mauritius, New
Zealand, Taiwan
Java, India, Indonesia,
Japan, Nigeria, Sri
Lanka, Tanzania
Brazil,
China, Ethiopia,
Japan, Kenya, Mauriti
us, Nigeria, Taiwan,
Tanzania
Argentina, China,
Hong Kong, Taiwan,
USA
India
China
Pestalotiopsidaceae
S, E
China
Pseudolachnea hispidula (Schrad.) B. Sutton
Pseudopestalotiopsis ampullacea F. Liu & L.
Cai*
Pseudopestalotiopsis camelliae Maharachch.,
L.D. Guo & K.D. Hyde*
Phyllostictaceae
Phyllostictaceae
P, S
Leaf spot
locality
Tanzania
Indonesia
Thailand
China
Florida
China, Japan
China
Papua New Guinea
Florida, Japan
Fiji, Hong Kong
References
Aptroot (1995)
Seifert (1990)
Thompson & Johnston (1953)
Thangaraj et al. (2019)
Alfieri et al. (1984)
Bai (2000), Kobayashi (2007)
Cheng et al. (2019)
Shaw (1984)
Alfieri et al. (1984), Kobayashi (2007)
Firman (1972), Dingley et al. (1981), Lu
et al. (2000), Zhuang (2001)
Thompson & Johnston (1953), Riley
(1960), Firman (1972), Tai (1979),
Dingley et al. (1981), Kobayashi (2007)
Tai (1979), Kobayashi (2007)
Kobayashi (2007)
Pennycook (1989), Gadgil (2005), Braun
et al. (2012)
Braun & Hill (2002), Gadgil (2005),
Kirschner et al. (2009)
Kobayashi (2007), Kamal (2010), Braun
et al. (2012)
Riley (1960), Tai (1979), Ragazzi &
Marino (1990), Mendes et al. (1998),
Crous & Braun (2003)
Deighton (1976), Alfieri et al. (1984),
Hsieh, & Goh (1990), Zhuang (2001),
Braun et al. (2012), Lu et al. (2000)
Mathur (1979)
Chen et al. (2018a), Liu et al. (2017),
Nozawa et al. (2018)
Maharachchikumbura et al. (2014),
Nozawa et al. (2018), Chen et al.
503
Table 3 Continued.
Disease caused
Species
Family
Life mode
locality
Pseudopestalotiopsis camelliae–sinensis F.
Liu & L. Cai*
Pseudopestalotiopsis chinensis F. Liu & L.
Cai
Pseudopestalotiopsis sp.
Pseudopestalotiopsis theae (Sawada)
Maharachch., K.D. Hyde & Crous*
Pestalotiopsidaceae
S, E
Pestalotiopsidaceae
P*
Grey blight
Pestalotiopsidaceae
Pestalotiopsidaceae
S, P
E, P**, S
Grey blight
Pyrenochaetopsis decipiens (Marchal)
Gruyter, Aveskamp & Verkley
Pyrrhoderma noxium (Corner) L.W. Zhou &
Y.C. Dai
Pythium sp.
Ramularia theicola Curzi
Cucurbitariaceae
China, Thailand
Brazil, China
(Thaiwan),
India, Japan,
Kenya, Korea,
Malawi, Malaysia,
Papua New Guinea,
Taiwan, Tanzania,
Thailand, Zimbabwe
India
Hymenochaetaceae
Sri Lanka
Adikaram & Yakandawal (2020)
Pythiaceae
Mycosphaerellaceae
Kobayashi (2007)
Farr & Rossman (2020)
Rigidoporus microporus (Sw.) Overeem
Meripilaceae
Rigidoporus vinctus (Berk.) Ryvarden
Rigidop
Rhizoctonia noxia (Donk) Oberw., R. Bauer,
Garnica & R. Kirschner
Rhizoctonia solani J.G. Kühn
Meripilaceae
Japan
Georgia, Italy,
Kazakhstan
Papua New Guinea,
Thailand
Thailand
Ceratobasidiaceae
Brazil
Mendes et al. (1998)
Ceratobasidiaceae
Rossmania aculeata (Petch) Lar.N.
Vassiljeva
Rossmania aculeata (Petch) Lar.N.
Vassiljeva
Rosellinia sp.
Xylariaceae
Japan, Malaysia,
Thailand
India, Sri Lanka
Xylariaceae
China
Thompson & Johnston (1953), Turner
(1971), Kobayashi (2007)
Agnihothrudu (1961),
Adikaram & Yakandawala (2020)
Tai (1979)
Xylariaceae
Brazil, Japan, Tanzani
a, Thailand
China
China
References
(2018a), This study
Chen et al. (2018a), Nozawa et al.
(2018), This study
Liu et al. (2017), Chen et al. (2018a),
Nozawa et al. (2018)
Liu et al. (2017), Wang et al. (2019b)
Zhuang (2001), Cho & Shin (2004),
Kobayashi (2007),
Maharachchikumbura et al. (2011),
Watanabe et al. (2012), Zhang et al.
(2012), Maharachchikumbura et al.
(2013, 2014), Nozawa et al. (2018)
Mathur (1979)
Thompson & Johnston (1953), Shaw
(1984)
Thompson & Johnston (1953)
Thompson & Johnston (1953), Riley
(1960), Alvarez (1976), Kobayashi
(2007)
504
Table 3 Continued.
Species
Sadasivanella indica Agnihothr
Sarocladium sp.
Scorias capitata Sawada
Scytalidium terminale G.V. Rao & de Hoog
Septobasidium acaciae Sawada
Septobasidium bogoriense Pat
Septobasidium pilosum Boedijn & B.A.
Steinm
Septobasidium sp.
Septobasidium tanakae (Miyabe) Boedijn &
B.A. Steinm
Setophoma antiqua F. Liu & L. Cai*
Setophoma endophytica F. Liu & L. Cai*
Setophoma longinqua F. Liu & L. Cai*
Setophoma yingyisheniae F. Liu & L. Cai*
Setophoma yunnanensis F. Liu & L. Cai*
Sillia theae Hara
Sporidesmium tropicale M.B. Ellis
Stagonospora theae Hara
Stilbum sp.
Taeniolella sp.
Terriera camelliicola (Minter) Y.R. Lin &
C.L. Hou*
Thozetellopsis tocklaiensis Agnihothr
Thelonectria lucida (Höhn.) P. Chaverri &
Salgado as Nectria lucida Höhn
Thelonectria mammoidea (W. Phillips &
Plowr.) C. Salgado & R.M. Sánchez
Tinctoporellus epimiltinus (Berk. & Broome)
Ryvarden
Trichoderma atroviride P. Karst
richoderma camelliae Jayaward., Manawas.,
X.H. Li, J.Y.Yan, & K. D. Hyde
Trichoderma lixii (Pat.) P. Chaverri
Trichoderma longibrachiatum Rifai
Trichoderma viride Pers
Family
Ascomycota
Hypocreales
Capnodiaceae
Hyaloscyphaceae
Septobasidiaceae
Septobasidiaceae
Septobasidiaceae
Life mode
Disease caused
locality
India
Fiji
Taiwan (China)
Netherlands
Taiwan (China)
Japan
Japan
References
Mathur (1979)
Dingley et al. (1981)
Tai (1979)
Rao & De Hoog (1975)
Kobayashi (2007)
Kobayashi (2007)
Kobayashi (2007)
Thailand
Japan
Thompson & Johnston (1953)
Kobayashi (2007)
China
China
China
China
China
Japan
Malaysia
Japan
Malawi
Papua New Guinea
China, India
Liu et al. (2019)
Liu et al. (2019)
Liu et al. (2019)
Liu et al. (2019), This study
Liu et al. (2019)
Senanayake et al. (2017)
Johnston (1960)
Kobayashi (2007)
Wiehe (1953)
Shaw (1984)
Zhang et al. (2015)
Chaetosphaeriaceae
Nectriaceae
India
Malaysia
Nectriaceae
Japan
Agnihothrudu (1961)
Thompson & Johnston (1953), Turner
(1971)
Kobayashi (2007)
Polyporaceae
Papua New Guinea
Shaw (1984)
Septobasidiaceae
Septobasidiaceae
Phaeosphaeriaceae
Phaeosphaeriaceae
Phaeosphaeriaceae
Phaeosphaeriaceae
Phaeosphaeriaceae
Sydowiellaceae
Pleosporomycetidae
Phaeosphaeriaceae
Chionosphaeraceae
Mytilinidiaceae
Rhytismataceae
P
P
P
P
P
S
P, S
Leaf
Leaf
Leaf
Leaf
Leaf
spot
spot
spot
spot
spot
Hypocreaceae
Hypocreaceae
S or P
S or P
China
China
This study
This study
Hypocreaceae
Hypocreaceae
Hypocreaceae
S or P
E
China
China
Kenya
This study
Wu et al. (2009)
Nattrass (1961)
505
Table 3 Continued.
Species
Family
Life mode Disease caused locality
References
Trichosphaeria corynephora (Cooke) Sacc
Trichosphaeriaceae
Japan
Kobayashi (2007)
Tripospermum sp.
Capnodiaceae
Malaysia
Turner (1971)
Valsaria insitiva (Tode) Ces. & De Not
Valsariaceae
Japan
Kobayashi (2007)
Identification confirmed by molecular data is marked with an asterisk (*). For the species, those with confirmed pathogenicity data are marked with a double asterisk (**).
The mode of life is given as (E) endophyte, (P) pathogen and (S) saprotroph.
Declarations
Funding
The research was funded by Beijing Talent Program for Dr Jiye Yan.
Conflicts of interest/Competing interests
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a
potential conflict of interest.
Availability of data and material
The sequence data generated in this study are deposited in NCBI GenBank (https://www.ncbi.nlm.nih.gov/genbank). All accession numbers are
given in Table 1. All isolates obtained in this study are deposited in culture collection and herbarium of Institute of Plant and Environmental
Protection, Beijing Academy of Agriculture and Forestry Sciences (JZB).
Authors’ contributions
JYY conceived the research. ISM and RSJ planned the basic research. HYL provided materials. ISM RSJ and YYZ conducted the experiments
and prepared the manuscript. WZ, AJLP, DNW, AJD, XHL, HLL, SB, RSJ, YHL, JYY and KH revised the manuscript. All authors read and approved
the final manuscript.
Acknowledgements
We would like to thank Dr Shaun Pennycook for the guidance with naming new species. Alan JL Phillips acknowledges the support from
UIDB/04046/2020 and UIDP/04046/2020 Centre grants from FCT, Portugal (to BioISI). Dhanushka Wanasinghe thanks CAS President’s International
Fellowship Initiative (PIFI) for funding his postdoctoral research (number 2021FYB0005) and the Postdoctoral Fund from Human Resources and
Social Security Bureau of Yunnan Province.
506
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