Research Article |
Deecksha Gomdola‡§|, Rajesh Jeewon¶#, Eric H.C. McKenzie¤, Ruvishika S. Jayawardena‡, Fatimah Al-Otibi#, Xia Tang‡, Yong Wang§, Kevin D. Hyde‡#, Li Fu|
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Corresponding author: Li Fu ( qianyier@sina.com ) Academic editor: Xin-Cun Wang
© 2025 Deecksha Gomdola, Rajesh Jeewon, Eric H.C. McKenzie, Ruvishika S. Jayawardena, Fatimah Al-Otibi, Xia Tang, Yong Wang, Kevin D. Hyde, Li Fu.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gomdola D, Jeewon R, McKenzie EHC, Jayawardena RS, Al-Otibi F, Tang X, Wang Y, Hyde KD, Fu L (2025) Phylogenetic diversity of Colletotrichum species (Sordariomycetes, Glomerellales, Glomerellaceae) associated with plant diseases in Thailand. MycoKeys 119: 137-195. https://doi.org/10.3897/mycokeys.119.152323
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Abstract
Colletotrichum is a cosmopolitan fungal genus, and its species are known for their important role as plant pathogens. They also occur as endophytes and saprobes. The genus comprises species complexes, many of which include cryptic species that are morphologically similar but phylogenetically distinct. Colletotrichum is characterized by globose to subglobose ascomata and acervular conidiomata and hyaline, aseptate ascospores and conidia, although a few species produce septate ascospores. Ascospores are typically cylindrical, oval, or fusiform, while conidia are mostly ellipsoidal to ovoid. Many Colletotrichum species also produce appressoria, which are formed at the tips of germ tubes or directly from the mycelium. In this study, we collected diseased leaves and pods with spots, blight, and blotches from Chiang Mai, Chiang Rai, and Tak provinces in Thailand. For species identification and delimitation, we employed a biphasic approach incorporating morphological data and multigene phylogenetic analyses of the locus of the internal transcribed spacer (ITS, nuclear rDNA consisting of ITS1-5.8S-ITS2) and the genes encoding for glyceraldehyde-3-phosphate dehydrogenase (GAPDH), chitin synthase 1 (CHS1), actin (ACT), beta-tubulin 2 (TUB2), histone (H3), and calmodulin (CAM). We obtained 20 Colletotrichum isolates representing eight species, among which we propose two new species: Colletotrichum dendrobii (from Dendrobium sp.) and C. musichiangmaiense (from Musa sp.). We also establish 13 new host records (for six species): C. castaneae (on Jasminum grandiflorum), C. chrysophilum (on Terminalia sp.), C. endophyticum (on Artocarpus heterophyllus, Begonia luxurians, Cassia sp., Castanopsis sp., Ficus auriculata, and Schefflera sp.), C. fructicola (on Castanea sp., Hedychium sp., and Rhododendron sp.), C. jiangxiense (on Artocarpus sp.), and C. schimae (on Jasminum sp.). In addition, this is the first study to present the sexual morph of C. endophyticum.
Key words:
2 novel species, 13 new host records, asexual, plant pathogens, sexual, species complexes, taxonomy
Introduction
Colletotrichum was introduced by
Colletotrichum has a cosmopolitan distribution with a wide host range, colonizing a myriad of plants and displaying remarkable adaptability to different environments (
There are over 1000 epithets in
In this study, 20 Colletotrichum isolates representing eight species were obtained from diseased leaves and pods collected in Thailand. Detailed morphological examinations were conducted, and their taxonomic placement was confirmed using phylogenetic analyses based on concatenated gene trees incorporating the locus of the internal transcribed spacer (ITS, nuclear rDNA consisting of ITS1-5.8S-ITS2) and the genes encoding for glyceraldehyde-3-phosphate dehydrogenase (GAPDH), chitin synthase 1 (CHS1), actin (ACT), beta-tubulin 2 (TUB2), histone (H3), and calmodulin (CAM). Two new species are proposed: Colletotrichum dendrobii, associated with pod blight of Dendrobium sp. and placed in the C. spaethianum species complex, and C. musichiangmaiense, a singleton species associated with leaf blight of Musa sp. Additionally, 13 new host records are established for six species: C. castaneae, C. chrysophilum, C. endophyticum, C. fructicola, C. jiangxiense, and C. schimae. The sexual morph of C. endophyticum is herein reported for the first time. Furthermore, an updated phylogenetic tree is provided, incorporating all Colletotrichum taxa with DNA sequence data across species complexes, alongside a revised count of species distributed within each complex and the number of singleton species.
This research enhances the understanding of plant pathogens that impact crop yield and food security through the development of accurate fungal diagnostics to support sustainable agriculture and by improving knowledge of plant-fungal interactions, biodiversity, and ecosystem health.
Materials and methods
Sample collection, examination, and material deposition
Diseased leaves and pods displaying symptoms such as spots, blight, and blotches were collected from Chiang Mai, Chiang Rai, and Tak Provinces in Thailand. Specimens were stored in plastic bags and brought to the laboratory for further examination. Micro- and macro-morphological characteristics were examined with a Motic SMZ 168 Series stereomicroscope. Single spore and tissue isolation techniques following the guidelines outlined in
Dried cultures are stored in the Mae Fah Luang University Herbarium (
DNA extraction, PCR amplification, and sequencing
Genomic DNA was extracted from axenic cultures using the BIOMIGA Fungus Genomic DNA Extraction Kit (San Diego, CA, USA), following the manufacturer’s protocols. Seven gene regions, the locus of the internal transcribed spacer (ITS, nuclear rDNA consisting of ITS1-5.8S-ITS2), and the genes encoding for glyceraldehyde-3-phosphate dehydrogenase (GAPDH), chitin synthase 1 (CHS1), actin (ACT), beta-tubulin 2 (TUB2), histone (H3), and calmodulin (CAM) were amplified and sequenced to identify the Colletotrichum isolates at the species level. The following primer pairs were used: ITS1/ITS4 for ITS (
The polymerase chain reaction (PCR) mixture was prepared to a final volume of 20 µL, which included 10 µL of PCR master mix, 1 µL each of forward and reverse primers (10 µM stock concentration), 7 µL of double-distilled water, and 1 µL of template DNA. The PCR program consisted of an initial denaturation at 95 °C for 3 min, followed by 40 cycles of denaturation at 95 °C for 45 s, annealing at 55 °C for 50 s for ITS, 52 °C for 1 min for GAPDH and ACT, and 58 °C for 1 min and 30 s for CHS1, TUB2, CAM, and H3, with an extension step at 72 °C for 2 min and a final extension at 72 °C for 10 min. Purification and sequencing of the PCR amplicons for the forward and reverse directions were performed by Sangon Biotech (Shanghai) Co., China.
Phylogenetic analysis
The quality of sequences was verified using DNA Baser Assembler, and unreadable bases at each end were trimmed manually. Sequences were subjected to BLAST searches in NCBI, and sequence data of other species were retrieved from GenBank (Suppl. material
Maximum likelihood (ML) phylogenetic analysis was performed in the IQ-TREE web server (https://iqtree.cibiv.univie.ac.at) by applying the default parameters and 1000 ultrafast bootstrap replicates (
The alignments are deposited in Figshare (https://doi.org/10.6084/m9.figshare.28377707). Phylogenetic trees were visualized in FigTree version 1.4.4 (
Results
Phylogenetic analyses
Dataset 1 consisted of 2920 characters (ITS = 1–524, GAPDH = 525–777, CHS1 = 778–1059, ACT = 1060–1315, TUB2 = 1316–1813, H3 = 1814–2189, and CAM = 2190–2920). Dataset 2 consisted of 3137 characters (ITS = 1–534, GAPDH = 535–797, CHS1 = 798–1087, ACT = 1088–1354, TUB2 = 1355–2022, H3 = 2023–2406, and CAM = 2407–3137). Dataset 3 consisted of 2920 characters (ITS = 1–519, GAPDH = 520–773, CHS1 = 774–1024, ACT = 1025–1261, TUB2 = 1262–1751, H3 = 1752–2122, and CAM = 2123–2920). Dataset 4 consisted of 2981 characters (ITS = 1–538, GAPDH = 539–788, CHS1 = 789–1070, ACT = 1071–1316, TUB2 = 1317–1806, H3 = 1807–2190, and CAM = 2191–2981). The log-likelihood of the consensus trees was -83865.668, -18946.853, -7997.184, and -11551.837 for trees 1–4, respectively (Figs
Maximum likelihood analysis (IQ-tree) based on a combined dataset of ITS, GAPDH, CHS1, ACT, TUB2, H3, and CAM sequences from dataset 1, which includes all Colletotrichum species. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with Monilochaetes infuscans (CBS 869.96) and M. melastomae (CBS145059). Type, ex-type, and reference species are denoted with T. Our isolates are in bold red font, while our new species are in bold blue font. Different color blocks represent distinct species complexes, with a single color used to denote singleton species.
Maximum likelihood analysis (IQ-tree) based on a combined dataset of ITS, GAPDH, CHS1, ACT, TUB2, H3, and CAM sequences from dataset 2, which includes all species in the C. gloeosporioides species complex. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with C. beeveri (CBS 128527) and C. boninense (CBS 123755), belonging to the C. boninense species complex. Type, ex-type, and reference species are denoted with T. Our isolates are in bold red font. Different color blocks represent distinct species complexes.
Maximum likelihood analysis (IQ-tree) based on a combined dataset of ITS, GAPDH, CHS1, ACT, TUB2, H3, and CAM sequences from dataset 3, which includes all species in the C. spaethianum species complex. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with C. luanense (CNUCC 157A-4-4 and CNUCC 157A-4-4-2), belonging to the C. destructivum species complex. Type, ex-type, and reference species are denoted with T. Our new species are in bold blue font. Different color blocks represent distinct species complexes.
Maximum likelihood analysis (IQ-tree) based on a combined dataset of ITS, GAPDH, CHS1, ACT, TUB2, H3, and CAM sequences from dataset 4, which includes all species in the Colletotrichum acutatum species complex. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with C. dematium (CBS 125.25) and C. parthenocissicola (
Taxonomy
Colletotrichum acutatum species complex
Colletotrichum schimae
Description.
Associated with leaf spots of Jasminum sp. Leaf spots circular, pale brown to brown. Sexual morph: Not observed. Asexual morph on PDA: Conidiomata 200–1000 µm diam. (x̄ = 400 µm, n = 10), semi-immersed, scattered or segregated, globose to subglobose, dark brown to black, exuding glistening yellowish to orange conidial mass. Setae not observed. Conidiophores formed directly from mycelium, hyaline, sometimes septate. Conidiogenous cells 16–45 × 2–4.5 µm (x̄ = 32 × 3.5 µm, n = 10), formed terminally or laterally on hyphae, hyaline, cylindrical, solitary or branched, straight or flexuous, tapering towards the apex. Conidia 9–15 × 2.5–4 µm (x̄ = 11.5 × 3.5 µm, n = 30; L/W ratio = 3.3), hyaline, cylindrical to fusiform, smooth-walled, guttulate, aseptate, mostly with acute ends. Appressoria not observed.
Culture characteristics.
Colonies on PDA reaching approximately 40 mm diam. after 7 d of incubation at 25 °C; mycelium initially white, becoming yellowish to olivaceous brown with age; elevation flat, with an entire margin.
Specimens examined.
Thailand • Chiang Mai Province, Doi Inthanon National Park, Kew Mae Pan nature trail, associated with leaf spots of Jasminum sp. (Oleaceae), 20 Oct 2021, D. Gomdola DGJas1(L2)-A (
GenBank accession numbers.
Known hosts, distributions, and lifestyles.
Endophytic
on Schima sp. in China (
Pathogenic
on leaves of Camellia sinensis var. assamica and Ilex chinensis in China (
Associated with leaf spots of Jasminum sp. in Thailand (this study).
Notes.
Our isolates (
Based on phylogenetic and morphological species concepts, we identify our isolates as Colletotrichum schimae. This study represents the first report of C. schimae associated with leaf spots on Jasminum sp. and establishes a new geographical record in Thailand.
Colletotrichum gloeosporioides species complex
Colletotrichum castaneae
Description.
Associated with leaf spots of Jasminum grandiflorum. Leaf spots irregular, pale brown to brown. Sexual morph: Not observed. Asexual morph on PDA: Conidiomata 150–500 µm diam. (x̄ = 250 µm, n = 5), semi-immersed, scattered or segregated, globose to subglobose, dark brown to black, exuding glistening yellowish to orange conidial mass. Setae not observed. Conidiophores hyaline. Conidiogenous cells 6.5–28 × 2–5 µm (x̄ = 18.6 × 3.1 µm, n = 10), hyaline, cylindrical to ampulliform, solitary or branched, straight or flexuous, tapering towards the apex. Conidia 13.5–19.5 × 4–6 µm (x̄ = 16.1 × 5 µm, n = 30; L/W ratio = 3.2), hyaline, cylindrical, straight, smooth-walled, guttulate, aseptate, mostly with rounded ends. Appressoria not observed.
Culture characteristics.
Colonies on PDA reaching approximately 65 mm diam. after 7 d of incubation at 25 °C; mycelium initially white, becoming yellowish to olivaceous brown with age; elevation flat, aerial, and filamentous with an entire or undulate margin.
Specimens examined.
Thailand • Chiang Mai Province, Doi Inthanon National Park, Kew Mae Pan nature trail, associated with leaf spots of Jasminum grandiflorum (Oleaceae), 20 Oct 2021, D. Gomdola DGJas3(L6)-C (
GenBank accession numbers.
Known hosts, distributions, and lifestyles.
Associated with leaf spots of Castanea mollissima in China (
Notes.
Based on the phylogenetic analyses, our isolates (
Based on phylogenetic and morphological species concepts, we identify our isolates as Colletotrichum castaneae. This study represents the first report of C. castaneae associated with leaf spots on Jasminum grandiflorum and establishes a new geographical record in Thailand.
Colletotrichum chrysophilum
Description.
Associated with leaf spots of Terminalia sp. Leaf spots irregular or oval, pale brown, surrounded with a dark brown margin. Sexual morph: Not observed. Asexual morph on substrate: Conidiomata 70–120 × 60–120 µm (x̄ = 94.5 × 96 µm, n = 10), semi-immersed, scattered, globose to subglobose, black, sometimes erumpent. Setae not observed. Conidiomatal wall 8.5–21 µm thick (x̄ = 14.6 µm, n = 10), consisting of 3–5 layers of thick-walled pseudoparenchymatous cells of textura angularis, outer layers dark brown, inner layer pale brown to hyaline. Asexual morph on PDA: Conidiophores hyaline, smooth-walled, aseptate, unbranched. Conidiogenous cells 16–22 × 3–4.5 µm (x̄ = 18.9 × 3.6 µm, n = 10), hyaline, cylindrical to ampulliform, straight or flexuous, tapering towards the apex. Conidia 13–19 × 4.5–6 µm (x̄ = 15.8 × 5.1 µm, n = 30; L/W ratio = 3.1), hyaline, cylindrical or oblong, smooth-walled, guttulate, aseptate, mostly with rounded ends. Appressoria 7–12 × 7–13 µm (x̄ = 10.9 × 10.5 µm, n = 5), hyaline, single-celled, globose to subglobose or irregular, smooth-walled.
Colletotrichum chrysophilum. a. Leaf spot (Terminalia sp.); b. Close up of a leaf spot; c. Conidiomata on substrate; d, e. Top and reverse of colony on PDA after 3 d f Section through a conidioma; g. Conidiomatal wall; h, i. Conidiogenesis and developing conidia; j, k. Appressoria; l, m. Conidia. Scale bars: 1 mm (b); 200 μm (c); 50 μm (f); 10 μm (g–m).
Culture characteristics.
Colonies on PDA reaching approximately 45 mm diam. after 7 d of incubation at 25 °C; mycelium greyish white, elevation flat or raised, aerial and filamentous with an entire or undulate margin.
Specimen examined.
Thailand • Chiang Rai Province, around the vicinity of Central Plaza, associated with leaf spots of Terminalia sp. (Combretaceae), 11 Jul 2019, D. Gomdola DG01-SM (
GenBank accession numbers.
Known hosts, distributions, and lifestyles
(listed chronologically). Endophytic on Theobroma cacao and Genipa americana in Panama (
Pathogenic; anthracnose on fruits of Musa sp. in Minas Gerais, Pernambuco, Santa Catarina, and São Paulo states in Brazil and Mexico (
Bitter rot
of Malus domestica in New York (
Preharvest decay
of Malus domestica in Italy (
Leaf spots
on Euterpe oleracea and Malus domestica in Brazil and Uruguay (
Associated with leaf spots of Terminalia sp. in Thailand (this study).
Notes.
Our isolate (
Based on phylogenetic and morphological species concepts, we identify our isolate as Colletotrichum chrysophilum. This study represents the first report of C. chrysophilum associated with leaf spots on Terminalia sp. and establishes a new geographical record in Thailand.
Colletotrichum endophyticum
Description.
Associated with leaf spots, blight, and blotches. Leaf spots circular, brown, surrounded with a dark brown to black margin. Leaf blight and leaf blotches brown to dark brown. Sexual morph on substrate: Ascomata 100–150 × 100–120 µm (x̄ = 122 × 112 µm, n = 10), solitary or aggregated, semi-immersed, globose to subglobose, black, creamy at the center, ostiolate. Setae not observed. Peridium 9.5–18.5 µm thick (x̄ = 15.5 µm, n = 10), composed of 2–3 layers of thick-walled pseudoparenchymatous cells of textura angularis. Asci 54–88 × 12–21 µm (x̄ = 68.6 × 17.3 µm, n = 10), operculate, unitunicate, cylindrical to truncate or obovoid, 6–8-spored. Ascospores 15–17 × 4.5–6 µm (x̄ = 15.5 × 5.1 µm, n = 20; L/W ratio = 3.0), uniseriate or biseriate, hyaline, cylindrical to subellipsoidal or oblong, smooth-walled, guttulate, aseptate, mostly with rounded or obtuse ends. Asexual morph on substrate: Conidiomata 120–250 × 120–150 µm (x̄ = 200 × 140 µm, n = 10), semi-immersed, scattered, globose to subglobose, black, sometimes erumpent, exuding creamy orange conidial mass. Setae not observed. Asexual morph on PDA: Conidiomata 200–1000 µm diam. (x̄ = 400 µm, n = 10), semi-immersed, scattered or segregated, globose to subglobose, exuding orange conidial mass. Setae not observed. Conidiophores 20–25 µm long (x̄ = 21.6 µm, n = 10), formed directly from mycelium, hyaline, cylindrical, branched, or unbranched. Conidiogenous cells 6–9 × 3–4.5 µm (x̄ = 7 × 3.5 µm, n = 10), hyaline, cylindrical, or ampulliform, straight or flexuous, tapering towards the apex. Conidia 14–19 × 4–6 µm (x̄ = 15.9 × 4.9 µm, n = 30; L/W ratio = 3.2), hyaline, cylindrical to ovoid, smooth-walled, guttulate, aseptate, with rounded ends. Appressoria 8–11 × 7–8 µm (x̄ = 8.5 × 7.5 µm, n = 5), hyaline, single-celled, irregular, smooth-walled or verruculose.
Colletotrichum endophyticum. a, a1, a2. Schefflera sp. a. Diseased leaf with spots; a1. Close up of a leaf spot; a2. Ascomata on substrate; b, b1, b2. Artocarpus heterophyllus; b. Fallen dried leaf with spots; b1. Close up of a leaf spot; b2. Conidioma sporulating on substrate; c. Leaf blight (Ficus auriculata); d. Leaf spots (Begonia luxurians); e. Fallen dried leaf with spots (Castanopsis sp.); f. Leaf blotches (Cassia sp.); g. Section through an ascoma; h. Peridium; i. Asci; j. Ascospores; k–m. Conidiogenesis and developing conidia; n–r. Conidia; s. Germinated conidium; t. Appressorium formed on PDA; u, v. Top and reverse of colony on PDA after 7 d; w, x. Spore mass formation on PDA after 21 d. Scale bars: 3 mm (a1, b1); 50 μm (a2, b2, g, I, x); 10 μm (h, j–m, s, t); 5 μm (n–r).
Culture characteristics.
Colonies on PDA reaching approximately 70 mm diam. after 7 d of incubation at 25 °C; mycelium initially white, becoming greyish white to dark grey at the center with age, elevation flat or raised, aerial and dense, with an entire margin, producing orange conidial mass.
Specimens examined.
Thailand • Chiang Mai Province, around vicinity of Mushroom Research Center, associated with leaf spots of Schefflera sp. (Araliaceae), 7 Jul 2021, D. Gomdola Div15-L1 (
GenBank accession numbers.
Known hosts, distributions, and lifestyles
(listed chronologically). Endophytic on leaves of Pennisetum purpureum in Thailand (
Saprobic
on an unknown wild fruit in Thailand (
Pathogenic; anthracnose on leaves of Camellia sinensis in China (
Fruit lesion
of Capsicum annuum in Thailand (
Leaf spots of Acacia confusa and Bauhinia blakeana in China (
Associated with anthracnose of fruits and leaves of Mangifera indica (
Notes.
Our isolates (
Intra-species nucleotide differences between our isolates and the ex-type of C. endophyticum (
| Gene regions |
|
|
|
|
|
|
|---|---|---|---|---|---|---|
| ITS | 1/476 | 1/476 | 4/476 | 1/476 | 4/476 | 4/476 |
| GAPDH | 4/227 | 8/261 | 1/228 | 4/249 | 5/258 | 8/240 |
| CHS1 | 2/247 | 3/279 | 4/279 | 5/245 | 1/279 | 5/277 |
| ACT | 1/234 | 2/220 | 2/236 | 3/234 | 2/235 | 2/234 |
| TUB2 | 0/668 | 8/668 | 0/656 | N/A | 0/668 | 10/668 |
| H3 | N/A | 6/360 | 0/360 | N/A | 1/360 | 7/360 |
Our isolates morphologically resemble the ex-type of C. endophyticum (
Based on phylogenetic and morphological species concepts, we identify our isolates as Colletotrichum endophyticum. This study represents six new host records for C. endophyticum associated with leaf spots of Begonia luxurians and Schefflera sp., spots on dried dead leaves of Castanopsis sp. and Artocarpus heterophyllus, leaf blight of Ficus auriculata, and leaf blotches of Cassia sp. in Thailand. Additionally, this is the first report for its sexual morph.
Colletotrichum fructicola
Description.
Associated with leaf spots, blight, and blotches. Leaf spots circular or irregular, pale brown to brown, surrounded with a dark brown margin. Leaf blight brown, surrounded with a dark brown margin. Leaf blotches reddish brown to dark brown. Sexual morph on substrate: Ascomata 75–150 × 80–150 µm (x̄ = 95 × 98 µm, n = 5), solitary, semi-immersed, globose to subglobose, brown, ostiolate. Setae not observed. Sexual morph on PDA: Ascomata 100–200 × 100–190 µm (x̄ = 131 × 123 µm, n = 5), solitary or aggregated, semi-immersed or superficial, globose to subglobose, black. Setae not observed. Peridium 8–38 µm thick (x̄ = 19.1 µm, n = 10), composed of 3–4 layers of thick-walled pseudoparenchymatous cells of textura angularis. Asci 55–70 × 8–11 µm (x̄ = 63 × 9.5 µm, n = 10), operculate, unitunicate, cylindrical to clavate or cymbiform, 6–8-spored. Ascospores 13–23.5 × 4–6.5 µm (x̄ = 17.9 × 5.1 µm, n = 25; L/W ratio = 3.5), uniseriate or biseriate, hyaline, subellipsoidal or oblong, reniform to falcate, somewhat fusiform, slightly curved, smooth-walled, guttulate, aseptate, mostly with obtuse or acute ends. Asexual morph on substrate: Conidiomata 80–150 × 60–140 µm (x̄ = 100 × 90 µm, n = 5), solitary, semi-immersed, globose to subglobose, brown, producing creamy to orange conidial mass. Setae not observed. Asexual morph on PDA: Conidiomata 200–800 µm diam. (x̄ = 350 µm, n = 10), semi-immersed, scattered or aggregated, globose to subglobose, exuding creamy to orange conidial mass. Setae not observed. Conidiophores formed directly from mycelium, hyaline, cylindrical, branched, or unbranched. Conidiogenous cells 6.5–22 × 2–4 µm (x̄ = 12.1 × 3 µm, n = 10), hyaline, cylindrical, or ampulliform, straight or flexuous, tapering towards the apex. Conidia 12–18 × 4.5–5.5 µm (x̄ = 15.4 × 5 µm, n = 25; L/W ratio = 3.1), hyaline, cylindrical to ovoid, smooth-walled, guttulate, aseptate, with rounded ends. Chlamydospores 6–7 × 6.5–8 µm (x̄ = 6.3 × 7.2 µm, n = 5), globose to subglobose, pale brown. Appressoria Not observed.
Culture characteristics.
Colonies on PDA reaching approximately 80 mm diam. after 7 d of incubation at 25 °C; mycelium initially white, becoming dark grey at the center when aged, elevation flat or raised, aerial and dense, with an entire margin.
Colletotrichum fructicola. a. Leaf spots (Castanea sp.); b. Leaf blight (Hedychium sp.); c. Leaf blotches (Rhododendron sp.); d. Ascomata on substrate (Rhododendron sp.); e. Conidiomata sporulating on substrate (Hedychium sp.); f. Top and reverse of colony on PDA after 21 d; g. Clusters of ascomata formed on culture; h. Section through ascomata; i. Peridium; j, k. Asci; l–q. Ascospores; r. Chlamydospore; s. Conidiogenesis and developing conidia; t–v. Conidia. Scale bars: 500 μm (d, e, g); 100 μm (h); 10 μm (i–v).
Specimens examined.
Thailand • Chiang Mai Province, Omkoi District, Yiang Piang Subdistrict, associated with leaf spots of Castanea sp. (Fagaceae), 16 Oct 2019, D. Gomdola DG367-L2 (
GenBank accession numbers.
Known hosts, distributions, and lifestyles
(listed chronologically). Pathogenic on plants; Leaf spots of Ficus edulis in Germany and Limonium spp. in Israel (
Leaf blotch
of Aesculus chinensis in China (
Shot-hole
on leaves of Prunus sibirica in China (
Brown sunken cladode spots
of Nopalea cochenillifera in Brazil (
Anthracnose
of Dioscorea spp. in Nigeria (
Fruit rot
of Persea americana in Australia (
Ripe rot
of Vitis spp. in Brazil (
Bitter rot
of Malus domestica in China (
Associated with spathe rot, spadix rot, and leaf spots of Anthurium andraeanum in Sri Lanka (
Colletotrichum fructicola was also reported from Fragaria × ananassa in Canada and the USA (
Pathogenic on a nematode
in China; infects horsehair worms (Chordodes formosanus), a parasite of praying mantises (
Pathogenic on humans
; causes Colletotrichum keratitis, a fungal infection of human eyes (
Endophytic
on Tetragastris panamensis and Theobroma cacao in Panama (
Notes.
Our isolates (
Our isolates morphologically resemble the ex-type of C. fructicola (ICMP 18581), having hyaline, smooth-walled, guttulate, and aseptate conidia and ascospores, with the conidia being cylindrical to ovoid with rounded ends and ascospores being oblong, reniform to falcate with obtuse or acute ends (
Based on phylogenetic and morphological species concepts, we identify our isolates as Colletotrichum fructicola. This study represents three new host records for C. fructicola associated with leaf spots of Castanea sp., leaf blight of Hedychium sp., and leaf blotches of Rhododendron sp. in Thailand.
Colletotrichum jiangxiense
Description.
Associated with leaf spots of Artocarpus sp. Leaf spots circular, oval, or irregular, pale brown to brown, surrounded with a dark brown margin. Sexual morph: Not observed. Asexual morph on substrate: Conidiomata 200–500 × 150–400 µm (x̄ = 300 × 200 µm, n = 10), acervular, semi-immersed, scattered or gregarious, globose to subglobose, dark brown to black, erumpent, exuding creamy orange conidial mass. Setae not observed. Asexual morph on PDA: Conidiomata 200–1000 µm diam. (x̄ = 400 µm, n = 10), semi-immersed, scattered or gregarious, globose to subglobose, dark brown to black, exuding creamy orange conidial mass. Setae not observed. Conidiophores hyaline, sometimes branched. Conidiogenous cells 12–19 × 2–5 µm (x̄ = 14.9 × 3.2 µm, n = 10), hyaline, cylindrical to ampulliform, straight or flexuous. Conidia 14–21 × 4–6.5 µm (x̄ = 16.5 × 5.2 µm, n = 30; L/W ratio = 3.2), hyaline, cylindrical to ellipsoidal, smooth-walled, guttulate, aseptate, with rounded ends (sometimes tapering towards one end), forming conidial anastomosis tubes. Conidial anastomosis tubes 2–16 × 1–2 µm (x̄ = 8.2 × 1.5 µm, n = 10), hyaline, smooth-walled, aseptate. Appressoria not observed.
Culture characteristics.
Colonies on PDA reaching approximately 55 mm diam. after 7 d of incubation at 25 °C; mycelium white to grey, elevation flat, cottony, with an entire margin.
Colletotrichum jiangxiense. a. Leaf spots (Artocarpus sp.); b. Conidiomata on substrate; c. Top and reverse of colony on PDA after 7 d; d, e. Conidiogenesis and developing conidia; f, g. Conidia; h. Conidia forming conidial anastomosis tubes. Scale bars: 200 μm (b); 50 μm (d); 20 μm (e); 10 μm (f–h).
Specimen examined.
Thailand • Chiang Mai: Omkoi District, Yiang Piang Subdistrict, associated with leaf spots of Artocarpus sp. (Moraceae), 16 Oct 2019, D. Gomdola DG360 (
Known hosts, distributions, and lifestyles
(listed chronologically). Endophytic on Camellia sinensis in China (
Pathogenic; brown lesions on leaves of Camellia sinensis in China (
Anthracnose
on fruits of Persea americana in Mexico (Ayvar-Serna et al. 2020;
Leaf spots
of Fraxinus americana in China (
Associated with leaf spots of Artocarpus sp. in Thailand (this study).
Notes.
Our isolate (
Based on phylogenetic and morphological species concepts, we identify our isolate as Colletotrichum jiangxiense. This study represents the first report of C. jiangxiense associated with leaf spots on Artocarpus sp. and establishes a new geographical record in Thailand.
Colletotrichum spaethianum species complex
Colletotrichum dendrobii , sp. nov.
Holotype.
Etymology.
The epithet refers to the host genus, Dendrobium, from which the species was isolated.
Description.
Associated with pod blight of Dendrobium sp. Pod blight elongated, pale brown to brown, surrounded with a dark brown margin. Sexual morph: Not observed. Asexual morph on substrate: Conidiomata 200–300 × 180–200 µm (x̄ = 240 × 190 µm, n = 5), acervular, semi-immersed, scattered or gregarious, dark brown to black. Setae 30–200 µm long (x̄ = 103 µm, n = 30), scattered or aggregated, straight or flexuous, smooth-walled, 1–5-septate, brown to dark brown, base darker and apex paler, base 3.5–12.5 µm wide (x̄ = 7.1 µm, n = 30), cylindrical or ampulliform, tapering to 1.5–5.5 µm (x̄ = 3.1 µm, n = 30) at the apex. Asexual morph on PDA: Conidiomata 200–1000 µm diam. (x̄ = 450 µm, n = 10), semi-immersed, scattered or aggregated, globose to subglobose, black, exuding orange conidial mass. Setae not observed. Conidiophores 13–30 µm long (x̄ = 19.7 µm, n = 10), hyaline, branched or unbranched, smooth-walled. Conidiogenous cells 6–14(–20) × 2.5–4.5 µm (x̄ = 8.3 × 3.5 µm, n = 10), hyaline, cylindrical to ampulliform, sometimes elongated, straight or flexuous. Conidia 11–19 × 4–6 µm (x̄ = 16.7 × 4.8 µm, n = 30; L/W ratio = 3.5), hyaline, falcate or fusiform, sometimes cylindrical or irregular in shape, smooth-walled, guttulate, aseptate, tapering towards both ends, base conical or rounded, apex acute. Appressoria 15–17 × 17–19 µm (x̄ = 15.6 × 17.8 µm, n = 5), hyaline, single-celled, globose to subglobose, smooth-walled or verruculose.
Colletotrichum dendrobii (
Culture characteristics.
Colonies on PDA reaching approximately 70 mm diam. after 7 d of incubation at 25 °C; mycelium initially greyish white, becoming dark grey to olivaceous brown with age; elevation flat, with an entire or undulate margin.
Specimen examined.
Thailand • Chiang Mai Province, Omkoi District, Yiang Piang Subdistrict, associated with pod blight of Dendrobium sp. (Orchidaceae), 16 Oct 2019, D. Gomdola DG386 (
Additional specimen examined.
Thailand • Chiang Mai Province, Omkoi District, Yiang Piang Subdistrict, associated with pod blight of Dendrobium sp. (Orchidaceae), 17 Oct 2019, D. Gomdola DG387 (
GenBank accession numbers.
Known hosts, distributions, and lifestyles.
Associated with pod blight of Dendrobium sp. in Thailand (this study).
Notes.
Our isolates (
Morphologically, the conidiophores and conidiogenous cells of our isolates are hyaline, while those of C. verruculosum (IMI 45525) are pale brown. Additionally, conidiophores of our isolates are shorter than those of C. verruculosum (up to 30 µm vs. 110 µm long). Conidiogenous cells of our isolates are also smaller compared to those of C. verruculosum (6–14(–20) × 2.5–4.5 µm vs. 10–25 × 3–5 µm). Setae of our isolates are 1–5-septate and 30–200 µm long, while those of C. verruculosum are 2–4-septate and 70–160 µm long. The conidial L/W ratio of our isolates differs from that of C. verruculosum (L/W ratio = 3.5 vs. 4.6) (
Based on phylogenetic analyses and morphological data following recommendations proposed by
The primary feature distinguishing Colletotrichum dendrobii from C. verruculosum (IMI 45525) is its variable and irregular conidial shape, along with their distinct phylogenetic lineages.
Colletotrichum singleton species
Colletotrichum musichiangmaiense , sp. nov.
Holotype.
Etymology.
The compound epithet refers to the host genus, Musa, from which the species was isolated, and the location, Chiang Mai, where the fungus was collected.
Description.
Associated with leaf blight of Musa sp. Leaf blight elongated, pale brown to brown, surrounded with a dark brown margin. Sexual morph: Not observed. Asexual morph on PDA: Conidiomata 200–1000 µm diam. (x̄ = 450 µm, n = 10), semi-immersed, scattered or gregarious, globose to subglobose, dark brown to black, exuding creamy orange conidial mass. Setae not observed. Conidiophores formed directly from mycelium, hyaline, aseptate, smooth-walled. Conidiogenous cells 9–16 × 2–6 µm (x̄ = 13.2 × 4.2 µm, n = 10), hyaline, cylindrical to ampulliform or clavate, straight or flexuous, tapering towards the apex. Conidia 12–19 × 4.5–6 µm (x̄ = 15.7 × 5.4 µm, n = 30; L/W ratio = 2.9), hyaline, cylindrical to clavate, straight or slightly flexuous, smooth-walled, guttulate, aseptate, mostly with obtusely rounded ends (sometimes tapering towards one end). Appressoria not observed.
Culture characteristics.
Colonies on MEA reaching approximately 75 mm diam. after 7 d of incubation at 25 °C; mycelium initially greyish white, becoming olivaceous brown with age, elevation flat, cottony, and fluffy, aerial and filamentous with an entire margin.
Specimen examined.
Thailand • Chiang Mai Province, Omkoi District, Yiang Piang Subdistrict, associated with leaf blight of Musa sp. (Musaceae), 16 Oct 2019, D. Gomdola DG385-T2 (
Known hosts, distributions, and lifestyles.
Associated with leaf blight of Musa sp. in Thailand (this study).
Notes.
Our isolate (
Discussion
The main aim of this study was to identify 20 isolates of Colletotrichum obtained from various diseased plants in Thailand based on morphological and phylogenetic species concepts. These isolates represent eight distinct species, encompassing two newly described taxa and six species that were reported from different hosts (accounting for 13 new host records). The Colletotrichum isolates obtained herein are distributed across three species complexes, except for one isolate that is a singleton species. The number of species and species complexes within this genus has been a topic of ongoing taxonomic debate (
Based on our phylogenies, the species complexes are listed below in descending order of species richness, with the number of species in each complex indicated in brackets: C. gloeosporioides (84), C. acutatum (50), C. boninense (41), C. graminicola-caudatum (37), C. destructivum (25), C. dematium (19), C. gigasporum (15), C. dracaenophilum (14), C. orchidearum (13), C. spaethianum (12), C. magnum (10), C. orbiculare (9), C. agaves (7), C. truncatum (6), and C. bambusicola (5). Among these, the C. gloeosporioides species complex is the most speciose (
Colletotrichum species exhibit life modes that allow them to manifest as saprobes, endophytes, and pathogens, depending on environmental conditions (
In addition to our two newly described species, we establish 13 new host records, representing six spp. isolated from various sites in Thailand, as listed below: C. castaneae (Jasminum grandiflorum), C. chrysophilum (Terminalia sp.), C. endophyticum (leaf spots of Artocarpus heterophyllus, Begonia luxurians, Castanopsis sp.; and Schefflera sp.; leaf blight of Ficus auriculata; and leaf blotches of Cassia sp.), C. fructicola (leaf spots of Castanea sp., leaf blight of Hedychium sp., and leaf blotches of Rhododendron sp.), C. jiangxiense, and C. schimae (leaf spots of Jasminum sp.). These species have also been documented from other hosts in previous studies (
Colletotrichum castaneae was previously found associated with Castanea mollissima in China (
Interestingly, this study presents the first detailed description and illustration of the sexual morph of Colletotrichum endophyticum, offering new insights into its reproductive biology and lifecycle. So far, this species was solely reported in its asexual state as saprobes, endophytes, and pathogens. Therefore, this finding emphasizes the importance of examining both sexual and asexual morphs for a comprehensive understanding of Colletotrichum taxonomy, ecology, and reproductive strategies.
Over the past few years, extensive research has been conducted to identify and classify Colletotrichum taxa (
In this study, we observed a similar pattern where species could potentially be synonymized. The type strains of both Colletotrichum chrysophilum [URM 7368; W.A.S. Vieira, W.G. Lima, M.P.S. Câmara, and V.P. Doyle] and C. noveboracense [CBS 146410; F. Khodadadi, P.L. Martin, V.P. Doyle, J.B. Gonzalez, and S.G. Aćimović] are positioned within the same clade, with strong statistical support (100% ML and 1.00 PP) (Fig.
To accurately identify and classify Colletotrichum species, a polyphasic approach, incorporating morphological traits with multigene phylogenetic analyses and geographical and/or ecological data, is recommended (
Acknowledgments
We thank Dr. Shaun Pennycook for suggesting and validating the nomenclature of the new species. Deecksha Gomdola thanks Mae Fah Luang University for providing the MFU student scholarship 2020 (reference: GR-ST-FS-63-06) to pursue a PhD program and for allocating the thesis writing grant [reference: 7702(6)/201 (No.0275)] and the research presentation or publication grant [reference No.0285]. Deecksha Gomdola also expresses her gratitude to the Mushroom Research Foundation (MRF) and the National Science and Technology Development Agency (NSTDA: Project No. P-19-52624), under the Doi Inthanon National Park Permission No. 0907.4/8218 and No. 0907.4/19647. Rajesh Jeewon thanks the University of Mauritius. Kevin D. Hyde and Fatimah Al-Otibi extend their appreciation to the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
This research was funded by the Research Innovation Fund Project of Shandong Institute of Pomology (GSS2022QN02) and the Technology Program of Tai’an, China (2021NS092), Guizhou University, King Saud University, Mae Fah Luang University, Mushroom Research Foundation, National Science and Technology Development Agency.
Author contributions
Conceptualization: DG, RJ. Data curation: DG, XT. Formal analysis: DG. Funding acquisition: DG, YW, KDH. Investigation: DG. Methodology: DG. Resources: KDH, YW, LF. Software: DG. Supervision: RJ, RSJ, EHC, KDH, YW. Validation: RJ, RSJ, EHC, KDH, YW, LF, FA, XT. Visualization: DG. Writing - original draft: DG. Writing - review and editing: DG, RJ, RSJ, EHC, KDH, YW, LF, FA, XT.
Author ORCIDs
Deecksha Gomdola https://orcid.org/0000-0002-0817-1555
Rajesh Jeewon https://orcid.org/0000-0002-8563-957X
Eric H.C. McKenzie https://orcid.org/0000-0001-9400-7112
Ruvishika S. Jayawardena https://orcid.org/0000-0001-7702-4885
Fatimah Al-Otibi https://orcid.org/0000-0003-3629-5755
Xia Tang https://orcid.org/0000-0003-2705-604X
Kevin D. Hyde https://orcid.org/0000-0002-2191-0762
Data availability
All data generated or analyzed during this study are included in this published article.
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Supplementary material
Isolates and their corresponding GenBank accession numbers
Data type: xlsx
Explanation note: Type, ex-type, and reference spe-cies are denoted with T. Our isolates are in bold font.