Re-identification of <i>Colletotrichum gloeosporioides</i> Species Complex Isolates in Korea and Their Host Plants

Le Dinh Thao; Hyorim Choi; Yunhee Choi; Anbazhagan Mageswari; Daseul Lee; Dong-Hyun Kim; Hyeon-Dong Shin; Hyowon Choi; Ho-Jong Ju; Seung-Beom Hong

doi:10.5423/PPJ.OA.09.2023.0133

Plant Pathol J > Volume 40(1); 2024 > Article

Thao, Choi, Choi, Mageswari, Lee, Kim, Shin, Choi, Ju, and Hong: Re-identification of Colletotrichum gloeosporioides Species Complex Isolates in Korea and Their Host Plants

Research Article

The Plant Pathology Journal 2024;40(1):16-29.

Published online: February 1, 2024

DOI: https://doi.org/10.5423/PPJ.OA.09.2023.0133

Re-identification of Colletotrichum gloeosporioides Species Complex Isolates in Korea and Their Host Plants

Le Dinh Thao^1,², Hyorim Choi¹, Yunhee Choi¹, Anbazhagan Mageswari¹, Daseul Lee¹, Dong-Hyun Kim¹, Hyeon-Dong Shin³, Hyowon Choi⁴, Ho-Jong Ju^5,^*, Seung-Beom Hong^1,^*

¹Korean Agricultural Culture Collection, Agricultural Microbiology Division, National Institute of Agricultural Sciences, Rural Development Administration, Wanju 55365, Korea

²Plant Pathology and Phyto-immunology, Plant Protection Research Institute, Duc Thang, Bac Tu Liem, Ha Noi 143315, Vietnam

³Division of Environmental Science and Ecological Engineering, Korea University, Seoul 02841, Korea

⁴Crop Protection Division, National Institute of Agricultural Sciences, Rural Development Administration, Wanju 55365, Korea

⁵Department of Agricultural Biology, Jeonbuk National University, Jeonju 54896, Korea

^*Co-corresponding authors. S.-B. Hong, E-mail) funguy@korea.kr, Phone) +82-63-238-3025, FAX) +82-63-238-3845. H.-J. Ju, E-mail) juhojong@jbnu.ac.kr, Phone) +82-63-270-2519, FAX) +82-63-270-2531

Handling Editor : Sook-Young Park

(Received on September 27, 2023; Revised on November 16, 2023; Accepted on December 3, 2023)

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted noncommercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Abstract

The Colletotrichum gloeosporioides species complex includes many phytopathogenic species, causing anthracnose disease on a wide range of host plants and appearing to be globally distributed. Seventy-one Colletotrichum isolates in the complex from different plants and geographic regions in Korea were preserved in the Korean Agricultural Culture Collection (KACC). Most of them had been identified based on hosts and morphological features, this could lead to inaccurate species names. Therefore, the KACC isolates were re-identified using DNA sequence analyses of six loci, comprising internal transcribed spacer, gapdh, chs-1, his3, act, and tub2 in this study. Based on the combined phylogenetic analysis, KACC strains were assigned to 12 known species and three new species candidates. The detected species are C. siamense (n = 20), C. fructicola (n = 19), C. gloeosporioides (n = 9), C. aenigma (n = 5), C. camelliae (n = 3), C. temperatum (n = 3), C. musae (n = 2), C. theobromicola (n = 2), C. viniferum (n = 2), C. alatae (n = 1), C. jiangxiense (n = 1), and C. yulongense (n = 1). Of these, C. jiangxiense, C. temperatum, C. theobromicola and C. yulongense are unrecorded species in Korea. Host plant comparisons showed that 27 fungus-host associations are newly reported in the country. However, plant-fungus interactions need to be investigated by pathogenicity tests.

Keywords: anthracnose, Colletotrichum fructicola, Colletotrichum siamense

The Colletotrichum gloeosporioides species complex is the most diverse group in the genus Colletotrichum, causing anthracnose disease in many economically important crops worldwide and also occurring as asymptomatic endophytes in a broad range of host plants (Hyde et al., 2009; Liu et al., 2022; Lu et al., 2004). In Korea, the species in this complex have been reported as destructive diseases on widely cultivated crops such as C. siamense and C. fructicola on apple (Kim et al., 2018a; Park et al., 2018), peach (Lee et al., 2020a, 2020b) and strawberry (Nam et al., 2013, 2022), and C. gloeosporioides on citrus, apple, grape, persimmon and strawberry (Korean Society of Plant Pathology, 2022).

C. gloeosporioides was first introduced by Penzig (1882), and then became a common phytopathogenic fungus with a global distribution. Prior to the availability of DNA sequence data, C. gloeosporioides and other taxa in the genus Colletotrichum were identified based on host species and morphological characteristics. However, micro-morphological features such as the size and shape of conidia, appressoria and setae or teleomorph characters overlapped in many species and these features could change under different growing conditions (Weir et al., 2012). For instance, cylindrical conidia with round ends were typical characteristics of species of the C. gloeosporioides species complex, but these characteristics were also observed in many members in the C. dracaenophilum, C. magnum, and C. orchidearum species complexes, or in some singleton species (Damm et al., 2019; Liu et al., 2022). Cultural characteristics of different strains of a species in the C. gloeosporioides species complex were often significant variations (Weir et al., 2012). On the other hand, most members of the C. gloeosporioides species complex have a broad host range and a plant could be associated with more than one Colletotrichum species (Jayawardena et al., 2021; Johnston, 2000; Johnston et al., 2005). Therefore, the host reference and morphology are insufficient to distinguish species members in this species complex.

A large number of Colletotrichum species have been re-identified based on DNA sequence analyses of multiple loci such as the nuclear ribosomal internal transcribed spacer (ITS) region, glyceraldehyde-3-phosphate dehydrogenase (gapdh), chitin synthase 1 (chs-1), histone-3 (his3), actin (act), and beta-tubulin 2 (tub2) (Cannon et al., 2012; Jayawardena et al., 2016; Liu et al., 2022; Weir et al., 2012). To date, at least 61 accepted species have been introduced in the C. gloeosporioides species complex (Liu et al., 2022; Zhang et al., 2023; Zheng et al., 2022). They shared high similarity of ITS sequences but could separate using multi-locus analyses of informative loci, especially gapdh and tub2 genes (Weir et al., 2012).

Isolates of the C. gloeosporioides species complex have been collected in Korea since the 1990s and stored in the Korean Agricultural Culture Collection (KACC), National Institute of Agricultural Sciences. However, these isolates had been identified based mainly on the host species and morphological characteristics by depositors. Hence, this study aims to (1) reassess the identification of the C. gloeosporioides species complex isolates in KACC using multi-locus analyses of six loci (ITS, gapdh, chs-1, his3, act, and tub2); (2) re-arrange the combination between host plants and fungal species.

Materials and Methods

Fungal isolates

Seventy-one isolates in the C. gloeosporioides species complex that originated in Korea and stored in KACC were used in this study. Details of the isolates are listed in Table 1. The living cultures preserved in liquid nitrogen were retrieved on potato dextrose agar (PDA; Difco Laboratories, Detroit, MI, USA) and were used for DNA extraction and phylogenetic analyses.

DNA extraction, polymerase chain reaction amplification, and sequencing

The mycelium of each isolate was taken from a 5-day-old culture on a PDA medium. Genomic DNA extraction was performed with the DNeasy plant mini kit (Qiagen, Hilden, Germany), following the manufacturer’s instructions. Six loci of Colletotrichum isolates, including ITS, gapdh, chs-1, his3, act, and tub2 were amplified with the primer pairs ITS1/ITS4 (White et al., 1990), GDF1/GDR1 (Guerber et al., 2003), CHS-79F/CHS-345R (Carbone and Kohn, 1999), CYLH3F/CYLH3R (Crous et al., 2004), ACT-512F/ACT-783R (Carbone and Kohn, 1999), and T1/BT2b (Glass and Donaldson, 1995; O’Donnell and Cigelnik, 1997), respectively. Each polymerase chain reaction (PCR) was performed in a volume of 25 μl containing 12.5 μl PCR Master Mix (2×), 8.5 μl nuclease-free water, 1 μl (4.5 pMol) of each primer, and 2 μl DNA template (100 ng/μl). PCR conditions of six loci were set up as described by Thao et al. (2023). PCR amplifications were carried out using a MJ Research PTC-200 Thermal Cycler (MJ Research, Ramsey, MN, USA). PCR products were purified with the QIAquick PCR Purification Kit (Qiagen) and checked by gel electrophoresis before being sent to the Macrogen Company (Seoul, Korea) for sequencing with the amplification primers.

Phylogenetic analysis

Raw sequences generated from forward and reverse primers in this study were trimmed and paired by MEGA 11 (Tamura et al., 2021). Assemble sequences were deposited to RDA-GeneBank (http://genebank.rda.go.kr) with accession numbers in Table 1. The closest reference sequences in GenBank were obtained by BLASTN search and were used for phylogenetic analyses with sequences from 71 KACC isolates and a species in the different species complex as an outgroup. The dataset of each locus was separately aligned using the multiple alignment program MAFFT version 7 (https://mafft.cbrc.jp/alignment/server/) with the G-INS-1 option. The poor alignments at both ends were cut and the alignments were improved by visual inspection in MEGA 11 and concatenated afterward in this software. A maximum likelihood (ML) phylogenetic tree was inferred based on the combined six datasets (ITS, gapdh, chs-1, his3, act, and tub2), using IQ-TREE with the best-fit model “TN+F+G4” and 1,000 ultrafast bootstrap replicates. The substitution model options were auto-evaluated according to the provided alignment files. The phylogenetic trees were viewed in MEGA11 and depicted in Adobe Illustrator.

Results

The phylogenetic analysis included 71 KACC isolates that originated in Korea (Table 1), 58 reference strains from 56 previously accepted species and C. arecacearum (MH0003) as the outgroup (Supplementary Table 1). The concatenated dataset alignment consisted of 2,504 characters (1,419 constant, 1,002 variable, and 502 parsimony-informative characters), including gaps. Of these, 565 characters were from ITS, 282 characters from gapdh, 251 characters from chs-1, 376 characters from his3, 264 characters from act, and 766 characters from tub2.

The multi-locus phylogenetic analysis resolved all KACC isolates into 15 clades. Each of the 12 clades was well clustered with an ex-type or holotype strain of the previously described species, supported by 86% to 100% ML bootstrap values (Fig. 1). Of which, 20 isolates clustered with C. siamense, 19 isolates with C. fructicola, nine isolates with C. gloeosporioides sensu stricto, five isolates with C. aenigma, three isolates with C. camelliae, three isolates with C. temperatum, two isolates with C. musae, two isolates with C. theobromicola, two isolates with C. viniferum, one isolate with C. alatae, one isolate with C. jiangxiense and one isolate with C. yulongense. The isolate KACC 47776 formed a sister clade to C. makassarense and C. tropicale. However, this isolate was distinguished from C. makassarense at gapdh (93.97% identity), act (98.76% identity), chs-1 (98.8% identity), and ITS (98.97% identity), and differentiated from C. tropicale by gapdh (94.03% identity), act (98.77% identity), and chs-1 (97.61% identity). The isolate KACC 47035 was phylogenetically related to C. endophyticum and C. artocarpicola, but shared low sequence similarity at gapdh (92.13% and 90.75%, respectively), act (97.93% and 98.35%), his3 (97.2% with C. endophyticum), chs-1 (98.01% with both species), and tub2 (98.34% with both species). Exception of ITS, all other loci of KAC 48700 shared low sequence similarity with the most closely related species, consisting of gapdh (91.3% identity shared with C. siamense), his3 (92.2% identity with C. siamense), act (95.45% identity with C. conoides, C. aenigma and C. siamense), tub2 (95,71% identity with C. endophyticum), and chs-1 (97.61% identity with C. siamense). According to phylogenetic analysis and BLASTN search results, each of the isolates, KACC 47776, KACC 47035, and KACC 48700, were genetically distinct from the other taxa.

Host plants of fungal species in this study

Twenty combinations between hosts and accepted fungal species found in this study have not been reported in any countries. However, the fungi were just isolated from the hosts and their pathogenicity was not confirmed. They are C. aenigma on Ampelopsis glandulosa f. citrulloides, Boehmeria japonica, Ilex rotunda, Laurus nobilis, and Zanthoxylum schinifolium; C. alatae on Dioscorea polystachya; C. camelliae on Cayratia japonica; C. fructicola on Pinus koraiensis, Pinus strobus, and Nepeta tenuifolia; C. gloeosporioides on Amaranthus hybridus, Boehmeria japonica, and Rhododendron micranthum; C. jiangxiense on Atractylodes ovata; C. siamense on Limonium sinuatum and Malus coronaria; C. temperatum on Malus domestica and Tricyrtis macropoda; C. theobromicola on Actinidia arguta; and C. yulongense on Camellia japonica. Seven other combinations, including C. fructicola on Capsicum annuum, Castanea sp., and Limonium sinuatum; C. gloeosporioides on Euonymus japonicus, Morus alba, and Portulaca oleracea; and C. siamense on Vitis vinifera are first reported in Korea (Table 2).

Discussion

Based on DNA sequence analyses of six loci, 43 KACC isolates were renamed, three isolates were newly identified at the species level, 21 strains remained as original names and three isolates were considered as three new species candidates. Nineteen isolates in this study had previously been identified as C. gloeosporioides sensu lato by Kim et al. (2006) and one isolate had been identified as C. acutatum by Lee et al. (2007) using morphological features, ITS and tub2 sequence analyses. However, these isolates were resolved by Kim et al. (2020) and by the present study into four different species, C. siamense (n = 10), C. fructicola (n = 7), C. theobromicola (n = 2), and C. temperatum (n = 1), based on multi-locus analyses of ITS, gapdh, chs-1, tub2 and (his3, act) or (cal and ApMat). The results of this study also revealed four fungal species, C. jiangxiense, C. temperatum, C. theobromicola, and C. yulongense, were first recorded in Korea.

C. siamense (n = 20), C. fructicola (n = 19), and C. gloeosporioides sensu stricto (n = 9) are the most dominant species among KACC isolates in the C. gloeosporioides species complex. According to previous studies and this work, C. gloeosporioides, C. siamense, C. fructicola, C. aenigma, C. jiangxiense, C. theobromicola, C. viniferum, and C. temperatum have a wide range of hosts, whereas C. musae has a narrow host range, on Musa spp. and Mangifera indica, and C. alatae is a host-specific fungus, on the genus Dioscorea (Farr and Rossman, 2021; Jayawardena et al., 2021; Li et al., 2019; Liu et al., 2022; Weir et al., 2012; Zhang et al., 2023). In addition, C. yulongense was originally described by Wang et al. (2019a) on Vaccinium dunalianum (Ericaceae family) and this fungus was found on Camellia japonica (Theaceae family) in this study as the second host plant. Another species, C. camelliae was also reported only on the genus Camellia of the family Theaceae (Liu et al., 2022), but this fungal species was herein from the different family Vitaceae (Cayratia japonica).

C. aenigma was introduced by Weir et al. (2012) and this fungus later became a very popular species, causing anthracnose on the grapevine, Asian pear, and walnut in China (Fu et al., 2019; Wang et al., 2021; Yan et al., 2015), on dragon fruits in Thailand (Meetum et al., 2015), on avocado in Israel (Sharma et al., 2017), on miracle in Japan (Truong et al., 2018), on grape and orange stonecrop in Korea (Choi et al., 2017; Kim et al., 2021). In our study, this fungus was isolated from five new hosts (Ampelopsis glandulosa f. citrulloides, Boehmeria japonica, Ilex rotunda, Laurus nobilis, and Zanthoxylum schinifolium) belonging to the five different families.

Apple (Malus domestica) is one of the most economically important crops in Korea. Kim et al. (2006) and Lee et al. (2007) implicated that C. gloeosporioides and C. acutatum were serious pathogens causing bitter rots of apple fruits in Korea. However, the identified species were unreliable as mentioned above. Later, causal agents of apple bitter rot disease in the country were authentically identified as C. siamense, C. fructicola, C. nymphaeae, C. fioriniae, C. gloeosporioides, and C. aenigma based on multi-locus sequence analyses (Cheon et al., 2016; Kim et al., 2020; Lee et al., 2021; Oo et al., 2018). The Colletotrichum isolates from apple in this work and in our earlier study (Thao et al., 2023) have been collected since 1997 from many different geographic locations in Korea, and the most prevalent species on apple were C. siamense (10 isolates), followed by C. fructicola (seven), C. nymphaeae (five) and C. fioriniae (four), while only one isolate of C. gloeosporioides sensu stricto was from apple and no C. acutatum sensu stricto isolates were found in KACC. Additionally, a fungal species C. temperatum, originally described by Doyle et al. (2013) on Vaccinium macrocarpon, was first detected on apple in this study.

Grape (Vitis vinifera) is an extensively cultivated crop worldwide and in Korea. The crop was seriously damaged by the infection of C. acutatum and C. gloeosporioides (Hong et al., 2008), C. viniferum (Oo and Oh, 2017), and C. aenigma (Kim et al., 2021). In this study, two isolates on grape (Vitis sp.) were identified as C. viniferum, whereas the other two isolates on Vitis vinifera were identified as C. siamense.

Kiwiberry (Actinidia spp.) has been widely cultivated in the country and worldwide (Kim et al., 2017; Latocha, 2017). The anthracnose symptoms caused by C. nymphaeae on kiwiberry fruits were observed in Jeonnam province (Kim et al., 2018b). Many other Colletotrichum species also infected kiwi plants worldwide such as C. aenigma on Actinidia arguta in China (Wang et al., 2019b), C. fioriniae, C. karsti, C. gloeosporioides, C. fructicola, and C. siamense on Actinidia spp. in Japan (Poti et al., 2023). A new combination of kiwiberry and C. theobromicola was found in this study.

The main causal fungi of chili pepper (Capsicum annuum) anthracnose in Korea were known as C. gloeosporioides sensu lato, C. acutatum s. lat., C. dematium s. lat., and C. trucatum s. lat. (Oo and Oh, 2016, 2020; Oo et al., 2017; Park and Kim, 1992). However, only one strain of C. fructicola in the C. gloeosporioides species complex from chili pepper is stored in KACC. While 16 strains of the C. acutatum species complex from chili pepper are preserved in KACC (Thao et al., 2023). It suggests that C. gloeosporioides s.c. is not a main causal fungus on chili pepper anthracnose in Korea.

Machilus thunbergii is a medicinal plant and it was widely distributed in the west-southern islands of Korea and some other Asian countries (Park et al., 1990). Only Glomerella cingulate and C. fructicola were formerly known as pathogens on this plant in Japan (Kobayashi, 2007) and China (Liu et al., 2022), respectively. A new species candidate (KACC47776) from Machilus thunbergii in Jeju, Korea was identified in this study. The second new species candidate (KACC48700) was from Paederia foetida, this host was also infected by many different Colletotrichum species (Liu et al., 2022). The third new species candidate (KACC47035) was the first report of a Colletotrichum fungus on coastal hogfennel (Peucedanum japonicum), an edible wild vegetable and medicinal resource.

A reassessment of members of the C. gloeosporioides species complex in Korea revealed new records of four Colletotrichum species and 27 combinations between plants and fungi in Korea. Of which, new findings of C. fructicola on pepper, C. siamense on grape, C. temperatum on apple, and C. theobromicola on kiwiberry could play an important role in the agricultural sciences and practices of Korea. However, the new combinations found in this study need to be clarified by pathogenicity tests in further studies.

Notes

Conflicts of Interest

No potential conflict of interest relevant to this work was reported.

Acknowledgments

This study was supported by a grant (PJ017286) from the National Institute of Agricultural Sciences and was a part of the “2023 KoRAA Long-term Training Program”, Rural Development Administration, Republic of Korea.

Electronic Supplementary Material

Supplementary materials are available at The Plant Pathology Journal website (http://www.ppjonline.org/).

ppj-oa-09-2023-0133-Supplementary-Table-1.pdf

Fig. 1

Maximum likelihood tree based on multi-locus sequences of ITS, tub2, his3, gapdh, chs-1, and act. Species names are followed by strain accession numbers and their hosts (blue). Isolates in this study are in bold and listed by the original names. Bootstrap values below 70% are not present. Ex-type strains are emphasised by asterisks (*). The tree is rooted to Colletotrichum arecacearum (MH003).

Table 1

Isolates of the Colletotrichum gloeosporioides species complex used in this study

Species (this study)	Isolate no.	Location	Collection date	Host^a	Previous identifications by depositors	Re-identifications in previous studies	RDA-GeneBank accession no.

							ITS	gapdh	chs-1	his3	act	tub2
C. aenigma	KACC 48663	Wando	2018	Laurus nobilis	C. siamense	-	RDA0062101	RDA0062486	RDA0067036	RDA0067150	RDA0066965	RDA0062292
	KACC 48701	Seogwipo	2018	Ampelopsis glandulosa f. citrulloides	C. aenigma	-	RDA0062110	RDA0062500	RDA0067035	RDA0067151	RDA0066966	RDA0062306
	KACC 48698	Jeju	2018	Boehmeria japonica	C. aenigma	-	RDA0062108	RDA0062498	RDA0067038	RDA0067148	RDA0066964	RDA0062304
	KACC 48896	Yeosu	2019	Zanthoxylum schinifolium	C. fructicola	-	RDA0062113	RDA0062504	RDA0067039	RDA0067147	RDA0066944	RDA0062309
	KACC 48654	Wando	2018	Ilex rotunda	C. siamense	-	RDA0062099	RDA0062484	RDA0067037	RDA0067149	RDA0066987	RDA0062290
C. alatae	KACC 47605	Gongju	2013	Dioscorea polystachya	Colletotrichum sp.	-	RDA0062083	RDA0062456	RDA0067092	RDA0067176	RDA0067034	RDA0062252
C. camelliae	KACC 43131	Boseong	2006	Camellia sinensis	C. gloeosporioides	-	RDA0062064	RDA0062421	RDA0067095	RDA0067180	RDA0067025	RDA0062216
	KACC 48695	Jeju	2018	Camellia japonica	C. camelliae	-	RDA0062105	-	RDA0067100	RDA0067182	RDA0067024	RDA0062301
	KACC 48694	Jeju	2018	Cayratia japonica	C. camelliae	-	RDA0062104	RDA0062495	RDA0067099	RDA0067183	RDA0067027	RDA0062300
C. fructicola	KACC 43014	Daejeon	2006	Pinus koraiensis	C. gloeosporioides	-	RDA0062063	RDA0062408	-	-	RDA0067015	RDA0062202
	KACC 40003	Daejeon	1993	Capsicum annuum	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0062019	RDA0062330	-	-	RDA0067017	RDA0062123
	KACC 42251	Daejeon	Unknown	Pinus strobus	C. gloeosporioides	-	RDA0062049	RDA0062377	RDA0067062	RDA0067117	RDA0067012	RDA0062173
	KACC 47033	Eumseong	2012	Nepeta tenuifolia	C. fructicola	-	RDA0062078	RDA0062449	RDA0067073	RDA0067111	RDA0067009	RDA0062244
	KACC 42502	Seoul	2006	Malus domestica	C. acutatum	C. acutatum (Lee et al., 2007)	RDA0062057	RDA0062396	RDA0067066	RDA0067115	RDA0067011	RDA0062191
	KACC 46566	Gyeryong	2009	Fragaria sp.	C. fructicola	-	RDA0062072	RDA0062439	RDA0067057	RDA0067119	RDA0067020	RDA0062232
	KACC 40695	Geochang	1999	Fragaria × ananassa	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006 )	RDA0060939	RDA0062351	RDA0067045	RDA0067143	RDA0067021	RDA0062142
	KACC 40812	Damyang	1998	Fragaria × ananassa	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0062041	RDA0062364	-	-	RDA0067016	RDA0062162
	KACC 40692	Namwon	1998	Limonium sinuatum	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0060934	RDA0062349	RDA0067058	RDA0067118	RDA0067013	RDA0062141
	KACC 42507	Jeongeup	2006	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007)	RDA0062061	RDA0062401	RDA0067053	RDA0067121	RDA0067014	RDA0062196
	KACC 49840	Sangju	2020	Atractylodes ovata	C. fructicola	-	RDA0062116	RDA0062511	RDA0067048	RDA0067122	RDA0067018	RDA0062315
	KACC 47034	Cheongyang	2012	Lycium sp.	C. fructicola	-	RDA0062079	RDA0062450	RDA0067074	RDA0067110	RDA0067008	RDA0062245
	KACC 49923	Yeongdong	2016	Malus domestica	C. gloeosporioides	-	RDA0062121	RDA0062521	RDA0067046	RDA0067142	RDA0067019	RDA0062325
	KACC 47512	Gongju	2013	Castanea sp.	C. gloeosporioides	-	RDA0062082	RDA0062455	RDA0067054	RDA0067120	RDA0066989	RDA0062251
	KACC 46964	Gunwi	2012	Malus domestica	C. gloeosporioides	-	RDA0062076	RDA0062444	RDA0067072	RDA0067112	RDA0066988	RDA0062239
	KACC 47685	Gunwi	2013	Malus domestica	C. gloeosporioides	-	RDA0062084	RDA0062458	RDA0067043	RDA0067144	RDA0066990	RDA0062263
	KACC 47827	Iksan	2014	Mangifera indica	C. fructicola	-	RDA0062088	RDA0062465	RDA0067063	RDA0067116	RDA0066991	RDA0062271
	KACC 46963	Gimcheon	2012	Malus domestica	C. gloeosporioides	-	RDA0062075	RDA0062443	RDA0067070	RDA0067113	RDA0067010	RDA0062238
	KACC 42499	Cheongsong	2006	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007)	RDA0062055	RDA0062393	RDA0067068	RDA0067114	RDA0067022	RDA0062188
C. gloeosporioides	KACC 40892	Jeju	Unknown	Cactus sp.	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0062042	RDA0062367	RDA0067081	RDA0067101	RDA0067003	RDA0062163
	KACC 47686	Mungyeong	2013	Malus domestica	C. gloeosporioides	-	RDA0062085	RDA0067187	RDA0067085	RDA0067103	RDA0066997	-
	KACC 42354	Namyangju	2006	Euonymus japonica	Colletotrichum sp.	-	RDA0062050	RDA0062380	RDA0067083	RDA0067104	RDA0066999	RDA0062176
	KACC 48637	Goesan	2017	Rhododendron micranthum	C. gloeosporioides	-	RDA0062097	RDA0062482	-	-	RDA0067001	RDA0062288
	KACC 40961	Sancheong	2001	Portulaca oleracea	Glomerella cingulata	G. cingulata (Kim et al., 2006)	RDA0062047	RDA0062373	RDA0067086	RDA0067102	RDA0066996	RDA0062169
	KACC 48696	Jeju	2018	Boehmeria japonica	C. gloeosporioides	-	RDA0062106	RDA0062496	RDA0067079	RDA0067108	RDA0067004	RDA0062302
	KACC 49841	Mungyeong	2020	Atractylodes ovata	C. gloeosporioides	-	RDA0062117	RDA0062512	RDA0067080	RDA0067107	RDA0067002	RDA0062316
	KACC 49597	Jangseong	2019	Morus alba	C. gloeosporioides	-	RDA0062114	RDA0062506	RDA0067084	RDA0067109	RDA0066998	RDA0062311
	KACC 48691	Okcheon	2018	Amaranthus hybridus	C. gloeosporioides	-	RDA0062103	RDA0062493	RDA0067082	RDA0067105	RDA0067000	RDA0062298
C. jiangxiense	KACC 49842	Sangju		Atractylodes ovata	C. kahawae	-	RDA0062118	RDA0062513	RDA0067093	RDA0067179	RDA0067028	RDA0062317
C. musae	KACC 49894	Wanju	2021	Musa × paradisiaca	C. musae	-	RDA0062120	RDA0062515	RDA0067078	RDA0067171	RDA0067005	RDA0062319
C. musae	KACC 40947	Daejeon	2001	Musa × paradisiaca	C. musae	-	RDA0062046	RDA0062372	RDA0067077	RDA0067172	RDA0067006	RDA0062168
C. siamense	KACC 49924	Yeongju	2016	Malus domestica	C. gloeosporioides	-	RDA0062122	RDA0062522	RDA0067047	RDA0067166	RDA0066969	RDA0062326
	KACC 40897	Suwon	Unknown	Malus domestica	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0062045	RDA0062370	RDA0067044	RDA0067167	RDA0066970	RDA0062166
	KACC 46988	Chungju	2011	Juglans regia	C. gloeosporioides	-	RDA0062077	RDA0062445	RDA0067050	RDA0067164	RDA0066981	RDA0062240
	KACC 48478	Sangju	2015	Unknown	C. siamense	-	RDA0062096	RDA0062479	RDA0067060	RDA0067159	RDA0066986	RDA0062285
	KACC 48106	Chuncheon	2015	Malus domestica	Colletotrichum sp.	-	RDA0062092	RDA0062474	RDA0067067	RDA0067155	RDA0066963	RDA0062279
	KACC 48402	Hwaseong	2017	Vitis vinifera	C. fructicola	-	RDA0062095	RDA0062477	RDA0067056	RDA0067161	RDA0066976	RDA0062283
	KACC 40696	Miryang	1999	Fragaria × ananassa	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0060941	RDA0062352	RDA0067052	RDA0067169	RDA0066979	RDA0062151
	KACC 42506	Paju	2005	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007)	RDA0062060	RDA0062400	RDA0067075	-	RDA0067007	RDA0062195
	KACC 42390	Sangju	2004	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007)	RDA0062051	RDA0062381	RDA0067076	-	RDA0066980	RDA0062177
	KACC 49782	Miryang	2020	Carya sp.	C. siamense	-	RDA0062115	RDA0062508	RDA0067049	RDA0067165	RDA0066982	RDA0062312
	KACC 49844	Mungyeong	2020	Atractylodes ovata	C. siamense	-	RDA0062119	RDA0062514	RDA0067071	RDA0067153	RDA0066971	RDA0062318
	KACC 40896	Yeongi	2000	Prunus persica	C. gloeosporioides	-	RDA0062044	RDA0062369	RDA0067064	RDA0067157	RDA0066974	RDA0062165
	KACC 40694	Suwon	1998	Limonium sinuatum	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0060936	RDA0062350	RDA0067042	RDA0067168	RDA0066978	RDA0066985
	KACC 40299	Uiseong	1997	Malus domestica	Glomerella cingulata	G. cingulata (Kim et al., 2006)	RDA0062028	RDA0062344	RDA0067091	RDA0067152	RDA0066943	RDA0062136
	KACC 40300	Gunwi	1997	Malus domestica	Glomerella cingulata	G. cingulata (Kim et al., 2006)	RDA0062029	RDA0062345	RDA0067051	RDA0067163	RDA0066968	RDA0062137
	KACC 42497	Gunwi	2006	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007); C. siamense (Kim et al., 2020)	RDA0062053	RDA0062391	RDA0067059	RDA0067160	RDA0066967	RDA0062186
	KACC 42498	Yeongju	2006	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007); C. siamense (Kim et al., 2020)	RDA0062054	RDA0062392	RDA0067069	RDA0067154	RDA0066972	RDA0062187
	KACC 42494	Cheongsong	2006	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007); C. siamense (Kim et al., 2020)	RDA0062052	RDA0066942	RDA0067055	RDA0067162	RDA0066977	RDA0066983
	KACC 45482	Jinju	2010	Malus coronaria	C. gloeosporioides	-	RDA0062069	RDA0062434	RDA0067065	RDA0067156	RDA0066973	RDA0062227
	KACC 43520	Cheongju	2006	Vitis vinifera	C. gloeosporioides	-	RDA0062067	RDA0062424	RDA0067061	RDA0067158	RDA0066975	RDA0062219
C. temperatum	KACC 42503	Yeongju	2004	Malus domestica	C. gloeosporioides	C. gloeosporioides (Lee et al., 2007)	RDA0062058	RDA0062397	RDA0067096	RDA0067178	RDA0067023	RDA0062192
	KACC 46405	Osan	2011	Tricyrtis macropoda	C. tricyrtii	-	RDA0062071	RDA0062438	RDA0067097	RDA0067177	RDA0067030	RDA0062231
	KACC 46374	Gapyeong	2011	Tricyrtis macropoda	C. tricyrtii	-	RDA0062070	RDA0062437	RDA0067094	RDA0067181	RDA0067026	RDA0062230
C. theobromicola	KACC 40698	Muan	1998	Actinidia arguta	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0060942	RDA0062353	RDA0067089	RDA0067185	RDA0067032	RDA0062152
C. theobromicola	KACC 40699	Muan	1998	Actinidia arguta	C. gloeosporioides	C. gloeosporioides (Kim et al., 2006)	RDA0060943	RDA0062354	RDA0067088	RDA0067186	RDA0067033	RDA0062153
C. viniferum	KACC 43519	Cheonan	2006	Vitis sp.	C. gloeosporioides	-	RDA0062066	RDA0062423	-	-	RDA0066993	RDA0062218
C. viniferum	KACC 43132	Cheonan	2006	Vitis sp.	C. gloeosporioides	-	RDA0062065	RDA0062422	RDA0067040	RDA0067146	RDA0066994	RDA0062217
C. yulongense	KACC 48651	Wando	2018	Camellia japonica	C. siamense	-	RDA0062098	RDA0062483	RDA0067098	RDA0067184	RDA0067029	RDA0062289
Colletotrichum sp.	KACC 47776	Jeju	2014	Machilus thunbergii	Colletotrichum sp.	-	RDA0062087	RDA0066941	RDA0067041	RDA0067170	RDA0066992	RDA0066984
Colletotrichum sp.	KACC 47035	Eumseong	2012	Peucedanum japonicum	Colletotrichum sp.	-	RDA0062080	RDA0062451	RDA0067087	RDA0067173	RDA0066995	RDA0062246
Colletotrichum sp.	KACC 48700	Jeju	2018	Paederia foetida	Colletotrichum sp.	-	RDA0062109	RDA0062499	RDA0067090	RDA0067175	RDA0067031	RDA0062305

ITS, internal transcribed spacer; gapdh, glyceraldehyde-3-phosphate dehydrogenase; chs-1, chitin synthase 1; his3, histone-3; act, actin; tub2, beta-tubulin 2; RDA, Rural Development Administration; KACC, Korean Agricultural Culture Collection.

^a The pathogenicity on host was not confirmed. The fungus was just isolated from the host.

Table 2

Comparison of host plants of KACC isolates and previous reports

Species	Host^a

	KACC	The List of Plant Diseases in Korea	Previous reports in Korea	Global reports	References
C. aenigma	*Ampelopsis glandulosa* f. *citrulloides**	-	-	-	-
	*Boehmeria japonica**	-	-	-	-
	*Ilex rotunda**	-	-	-	-
	*Laurus nobilis**	-	-	-	-
	*Zanthoxylum schinifolium**	-	-	-	-
C. alatae	*Dioscorea polystachya**	-	-	-	-
C. camelliae	Camellia sinensis	Camellia sinensis	Camellia sinensis	Camellia sinensis	Hassan et al. (2023)
	Camellia japonica	Camellia japonica	Camellia japonica	Camellia japonica	Korean Society of Plant Pathology (2022)
	*Cayratia japonica**	-	-	-	-
C. fructicola	Atractylodes ovata	-	Atractylodes ovata	Atractylodes ovata	Hassan et al. (2022)
	*Capsicum annuum*	-	-	Capsicum annuum	Liu et al. (2016)
	*Castanea* sp.	-	-	Castanea sp.	Jiang et al. (2021)
	Fragaria × ananassa	-	Fragaria × ananassa	Fragaria × ananassa	Nam et al. (2013)
	*Limonium sinuatum*	-	-	Limonium sinuatum	Zhang et al. (2023)
	Lycium sp.	-	Lycium chinense	Lycium chinense	Paul et al. (2014)
	Malus domestica	Malus domestica	Malus domestica	Malus domestica	Kim et al. (2018a)
	Mangifera indica	Mangifera indica	Mangifera indica	Mangifera indica	Joa et al. (2016)
	*Pinus koraiensis**	-	-	-	-
	*Pinus strobus**	-	-	-	-
	*Nepeta tenuifolia**	-	-	-	-
C. gloeosporioides	*Amaranthus hybridus**	-	-	-	-
	Atractylodes ovata	Atractylodes ovata	Atractylodes ovata	Atractylodes ovata	Hassan et al. (2022)
	*Boehmeria japonica**	-	-	-	-
	Cactus sp.	-	Cactus sp.	Cactus sp.	Kim et al. (2000)
	*Euonymus japonicus*	-	-	Euonymus japonicus	Huang et al. (2016)
	Malus domestica (syn. M. pumila)	Malus pumila	Malus pumila	Malus pumila	Cheon et al. (2016)
	*Morus alba*	-	-	Morus alba	Sharma et al. (2013)
	*Portulaca oleracea*	-	-	Portulaca oleracea	Simmonds (1966)
	*Rhododendron micranthum**	-	-	-	-
C. jiangxiense	*Atractylodes ovata**	-	-	-	-
C. musae	Musa × paradisiaca	Musa × paradisiaca	Musa × paradisiaca	Musa × paradisiaca	Lim et al. (2002)
C. siamense	Atractylodes ovata	-	Atractylodes ovata	Atractylodes ovata	Hassan et al. (2022)
	Carya sp.	-	Carya sp.	Carya sp.	Oh et al. (2021)
	Fragaria × ananassa	-	Fragaria × ananassa	Fragaria × ananassa	Nam et al. (2022)
	Juglans regia		Juglans regia	Juglans regia	Cho et al. (2023)
	*Limonium sinuatum**	-	-	-	-
	*Malus coronaria**	-	-	-	-
	Malus domestica (syn. M. pumila)	Malus pumila	Malus pumila	Malus pumila	Park et al. (2018)
	Prunus persica	Prunus persica	Prunus persica	Prunus persica	Lee et al. (2020a)
	*Vitis vinifera*	-	-	Vitis vinifera	Zhang et al. (2023)
C. temperatum	*Malus domestica**	-	-	-	-
C. temperatum	*Tricyrtis macropoda**	-	-	-	-
C. theobromicola	*Actinidia arguta**	-	-	-	-
C. viniferum	Vitis sp.	Vitis vinifera	Vitis vinifera	Vitis vinifera	Oo and Oh (2017)
C. yulongense	*Camellia japonica**	-	-	-	-
Colletotrichum sp. (KACC 47776, new species candidate)	*Machilus thunbergii**	-	-	-	-
Colletotrichum sp. (KACC 47035, new species candidate)	*Peucedanum japonicum**	-	-	-	-
Colletotrichum sp. (KACC 48700, new species candidate)	*Paederia foetida**	-	-	-	-

Hosts in “bold” and “*” are unreported in Korea and in the world, respectively.

KACC, Korean Agricultural Culture Collection.

^a The pathogenicity on host was not confirmed. The fungus was just isolated from the host.

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