Colletotrichum macroconidii sp. nov. and six new records of Colletotrichum (Glomerellaceae, Glomerellales) from southwestern China

Abstract

Colletotrichum, a genus in the phylum Ascomycota and the family Glomerellaceae, is globally recognized as a significant plant pathogen affecting various hosts. In this study, 31 Colletotrichum strains were isolated from plant hosts in southwestern China, specifically from Guizhou and Yunnan Provinces. Phylogenetic and morphological analyses revealed one novel species, Colletotrichum macroconidiisp. nov., and 11 previously known species. Six of these species represent new host and regional records: C. fioriniae, C. trichellum, C. juglandicola, C. nanhuaense, C. jiangxiense, and C. magnum, in addition to five known species: C. nymphaeae, C. metake, C. fructicola, C. siamense, and C. gloeosporioides. Multilocus phylogenetic analyses placed strains of C. jiangxiense in a well-supported clade with C. nullisetosum, C. oblongisporum, C. gracile, and C. tengchongense, which are here treated as synonyms due to their nomenclatural invalidity, morphological overlap, and phylogenetic evidence. Similarly, C. speciosum and C. simulanticitri clustered with C. nymphaeae and are synonymized with it based on insignificant genetic divergence and invalid nomenclatural status. Species identification was conducted using a comprehensive approach that combined multilocus phylogenetic analysis with detailed morphological characterization. This expanded dataset significantly contributes to our understanding of the genetic diversity and ecological distribution of Colletotrichum in southwestern China.

Introduction

The genus Colletotrichum (Glomerellaceae, Sordariomycetidae, Sordariomycetes, Ascomycota) was first described by Corda in 1831, with C. lineola designated as the type species. This species was originally isolated from an unidentified branch of an Apiaceae family host plant, collected in the Czech Republic. Colletotrichum lineola is characterized by acervuli, fusiform, curved, hyaline conidia with acute ends, and brown, opaque, subulate setae (Corda 1831). Species of Colletotrichum are known for their broad host range and their ability to colonize plant tissues through various infection strategies, including intracellular hemibiotrophy, subcuticular necrotrophy, and intramural necrotrophy (Bailey et al. 1992). Most Colletotrichum species exhibit a hemibiotrophic lifestyle, initiating infection with a symptomless biotrophic phase, which transitions into a necrotrophic phase that leads to death host cell and decay of tissue (Perfect et al. 1999; Münch et al. 2008; Crouch and Beirn 2009; Vargas et al. 2012; Damm et al. 2014; Rajarammohan 2021; Tsushima et al. 2021). In addition to their role as plant pathogens, some Colletotrichum species function as endophytes or saprotrophs, demonstrating their ecological versatility (Promputtha et al. 2007; Hyde and Soytong 2008; Yan et al. 2015). Currently, the genus has 874 named species, of which 764 are currently accepted in the Species Fungorum database (https://www.speciesfungorum.org; July 28, 2025). Colletotrichum species are globally distributed and capable of infecting a wide range of host plants, with over 1,350 documented host associations involving more than 720 plant species across diverse families, including vegetables, legumes, cereals, and both woody and herbaceous plants (Talhinhas and Baroncelli 2021). Colletotrichum species are among the most economically significant plant pathogens worldwide, responsible for anthracnose and blight diseases across a wide range of hosts, including fruits, vegetables, ornamental plants, and staple crops (Dean et al. 2012; Jayawardena et al. 2016b, 2021). Characteristic symptoms of Colletotrichum infections include sunken, dark lesions with distinct margins on leaves, stems, and fruits, leading to wilting, tissue necrosis, fruit rot, and premature fruit drop (Prusky et al. 2000; Hyde et al. 2014; Trkulja et al. 2024). These diseases contribute to significant global agricultural losses, affecting crops such as apple, mango, banana, citrus, pepper, coffee, and strawberry (Than et al. 2008; Cannon et al. 2012; Huang et al. 2013; Lima et al. 2013; Trkulja et al. 2024).

Morphologically, Colletotrichum species exhibit a variety of structures, including conidiomata, conidia, setae, and appresoria though considerable variation exists across species (Crouch et al. 2009). Accurate identification has historically been challenging due to the morphological similarities among species and the presence of cryptic lineages (Vieira et al. 2020; Bhunjun et al. 2021). This has led to the increased adoption of molecular approaches, such as multilocus sequence typing (MLST), utilizing genetic markers like the internal transcribed spacer (ITS), actin (act), beta-tubulin (β-tubulin), glyceraldehyde-3-phosphate dehydrogenase (gapdh), and chitin synthase 1 (chs-1) (White et al. 1990; O’Connell et al. 2012; Bhunjun et al. 2021). These molecular tools have proven essential in distinguishing closely related species and elucidating complex phylogenetic relationships within the genus. The current classification of Colletotrichum comprises 15 species complexes, including the C. acutatum, C. agaves, C. boninense, C. caudatum, C. dematium, C. destructivum, C. dracaenophilum, C. gigasporum, C. gloeosporioides, C. graminicola, C. magnum, C. orbiculare, C. orchidearum, C. spaethianum, and C. truncatum species complexes, along with numerous singletons (Marin-Felix et al. 2017; Damm et al. 2019; Jayawardena et al. 2020; Bhunjun et al. 2021). The C. gloeosporioides species complex is particularly notable within the genus due to its significance as a plant pathogen, with many of its members confirmed as pathogenic (Weir et al. 2012). Notably, C. gloeosporioides and C. siamense have been identified as pathogens affecting apple trees (Chen et al. 2022), rubber (Cao et al. 2019; Athinuwat et al. 2024), mango (de Souza et al. 2013), Toxicodendron radicans (Kasson et al. 2014), Parthenocissus tricuspidata (Zhao et al. 2020; Wang et al. 2025), and Kadsura coccinea (Jiang et al. 2022), among others.

Extensive surveys in tropical and subtropical regions have led to the discovery of numerous novel Colletotrichum species and newly recognized host associations, underscoring the ecological and economic importance of this genus (Armand et al. 2023b; Jiang et al. 2025; Wang et al. 2025). In China, research on Colletotrichum has gained significant attention due to the diverse agroecological landscapes and rich plant biodiversity (Tao et al. 2013; Liu et al. 2015, 2020, 2022, 2024; Zhang et al. 2020, 2024; Jiang et al. 2025; Lu et al. 2025; Usman et al. 2025). For example, Colletotrichum asianum is a pathogen causing mango anthracnose (Mangifera sp.), severe losses in both cultivation and postharvest stages, with yield reductions of 30%–60% (Kamle and Kumar 2016; Li et al. 2019; Wu et al. 2022). Additionally, Colletotrichum gloeosporioides has been identified as a pathogen causing kiwifruit anthracnose (Actinidia chinensis) in China, where approximately 20% of surveyed trees exhibited characteristic disease symptoms (Li et al. 2017). Major crops such as mango, tea, pepper, chili, rubber, and soybean are particularly vulnerable, with outbreaks often leading to substantial yield losses (Tao et al. 2013; Liu et al. 2015, 2022, 2024; Zhang et al. 2020, 2024; Jiang et al. 2025; Lu et al. 2025; Usman et al. 2025). Despite the challenges posed by complex pathogenicity, broad host ranges, and overlapping morphological features, recent advancements in molecular phylogenetics have greatly enhanced the resolution of Colletotrichum taxonomy, enabling more precise species identification and the discovery of cryptic lineages (Cai et al. 2009; Cannon et al. 2012; Marin-Felix et al. 2017; Damm et al. 2019; Jayawardena et al. 2020; Liu et al. 2022). These advancements have facilitated the development of targeted disease management strategies and underscored the importance of ongoing monitoring for effective crop protection.

This study aims to investigate Colletotrichum strains collected from southwestern China, with a focus on their phylogenetic diversity and morphological characteristics. By integrating molecular and morphological data, this research seeks to enhance the understanding of Colletotrichum diversity in the region and contribute to more accurate fungal diagnosis and management strategies in these vital ecosystems.

Materials and methods

Sampling and examination of specimens

The samples were collected from 2023 to 2024 in Guizhou and Yunnan Provinces, China. The specimens were taken to the laboratory in paper bags to be examined and described. Morphological characters such as conidiophore, conidiogenous cell, and conidia, were studied using a Zeiss microscope (Jena, Germany) and photographed with an AxioCam 208 color camera (Carl Zeiss Microscopy GmbH, Jena, Germany), while the size measurements were taken with the assistance of ZEN 3.0 (Blue Edition) software (Jena, Germany). Photoplates were made using Adobe Photoshop 2025 version 26.5 (Adobe Systems, CA, USA). Only the new species and newly recorded taxa are illustrated in this study.

The cultures were acquired by the tissue isolation technique as described by Norphanphoun and Hyde (2023). Single hyphal tips were transferred onto 2% Potato Dextrose Agar (PDA), Oatmeal agar (OA) and Water agar (WA) plates and incubated at room temperature (25 °C ± 2 °C): 12 hours dark and 12 hours light. The cultural features were observed and documented at 5, 7, and 14 days. Dried cultures were prepared as described by Sinclair and Dhingra (1995) and deposited at the Herbarium of the Department of Plant Pathology, Agricultural College, Guizhou University (HGUP). Dried and living cultures have been deposited in the culture collection at the Plant Pathology Department of the College of Agriculture, Guizhou University, China (GUCC). The enumeration for the new taxon was conducted in the MycoBank online database (https://www.mycobank.org; Robert et al. 2013).

DNA extraction, amplification via PCR, and sequencing

Genomic DNA was extracted from fresh fungal mycelia growing on PDA at room temperature (25 °C ± 2) for two weeks using the Biospin Fungal Genomic DNA Extraction Kit (BioFlux) following the manufacturer’s protocols. Polymerase chain reactions (PCR) were carried out using the following primer pairs: ITS5/ITS4 to amplify the internal transcribed spacer region (ITS), ACT512F/ACT738R for actin (act), GDF/GDR for partial glyceraldehyde-3-phosphate dehydrogenase region (gapdh), T1/ Btub4Rd or Bt2a/Bt2b for beta-tubulin (β-tubulin), CHS-79F/CHS-354R for chitin synthase (chs-1), H3F/H3R for imidazoleglycerol-phosphate dehydratase (his3), and CL1C/CL2C for calmodulin (cal) (Weir et al. 2012; Schena et al. 2014; Kim et al. 2021; Li et al. 2021).

The amplification reactions were carried out using the following protocol: 20 μL reaction volume containing 1 µl of DNA template, 1 µL (20 µM stock concentration) of each forward and reverse primers, 10µl of 2Mix (Vazyme Biotech Co., Ltd), and 7 µl of double-distilled water (ddH₂O). The PCR thermal cycling program for each locus is described in Table 1. The purification and sequencing of PCR products using the amplification primers specified above were conducted at Sangon Biotech (Shanghai, China) Co., Ltd. for Sanger sequencing. After sequencing, the sequence data were uploaded on GenBank, and the relevant information is listed in Table 2.

Table 1.

Polymerase chain reactions (PCR) thermal cycling programs for each locus.

Gene	Primers	PCR thermal cycle protocols*
ITS	ITS1/ITS4	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 52 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
actin	ACT512F/ACT738R	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 60.3 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
gapdh	GDF/GDR	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 61.9 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
β-tubulin	T1/T2	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 55.7 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
	2a/2b	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 61.3 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
chs-1	CHS-79F/ CHS-354R	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 59.4 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
his3	H3F/H3R	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 62.1 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min
cal	CL1C/CL2C	ID 95 °C for 5 min, 35 cycles of D at 95 °C for 30 s, A at 62.1 °C for 30 s, E at 72 °C for 1 min, FE at 72 °C for 10 min

*ID: initial denaturation; D = denaturation; A = annealing; E = elongation; FE = final extension

Table 2.

GenBank accession numbers of the sequences used in phylogenetic analyses in this study.

Species	Strain	Host	Location	Accession numbers								References
				ITS	gapdh	chs–1	act	β-tubulin	his	cal	ApMat
Colletotrichum acutatum species complex
C. abscissum	COAD 1877 T	Citrus sinensis	Brazil	KP843126	KP843129	KP843132	KP843141	KP843135	KP843138	–	–	Crous et al. (2015)
C. acerbum	CBS 128530 T	Malus domestica	New Zealand	JQ948459	JQ948790	JQ949120	JQ949780	JQ950110	JQ949450	–	–	Damm et al. (2012)
C. acutatum	CBS 112996 T	Carica papaya	Australia	JQ005776	JQ948677	JQ005797	JQ005839	JQ005860	JQ005818	–	–	Damm et al. (2012)
C. americanum	RGM 3380 T	Drimys winteri	Chile	OR644563	OR644970	OR645022	OR645076	OR645128	OR659700	–	–	Zapata et al. (2024)
C. americanum	RGM 3129	Laurelia sempervirens	Chile	OR644556	OR644963	OR645015	OR645069	OR645121	OR659693	–	–	Zapata et al. (2024)
C. arboricola	CBS 144795 T	Fuchsia magellanica	Chile	MH817944	MH817950	–	MH817956	MH817962	OR659702	–	–	Crous et al. (2018)
C. australe	CBS 116478 T	Trachycarpus fortunei	South Africa	JQ948455	JQ948786	JQ949116	JQ949776	JQ950106	JQ949446	–	–	Damm et al. (2012)
C. bannaense	CGMCC 3.18887 T	Rubber, leaf	China	MG209637	MG242005	MG241995	MG242001	MG209659	–	–	–	Liu et al. (2021)
C. brisbanense	CBS 292.67 T	Capsicum annuum	Australia	JQ948291	JQ948621	JQ948952	JQ949612	JQ949942	JQ949282	–	–	Damm et al. (2012)
C. cairnsense	BRIP 63642 T	Capsicum annuum	Australia	KU923672	KU923704	KU923710	KU923716	KU923688	KU923722	–	–	De Silva et al. (2017)
C. carthami	SAPA100011 T	Carthamus tinctorius	Japan	AB696998	–	–	–	AB696992	–	–	–	Uematsu et al. (2012)
C. chrysalidocarpi	ZHKUCC 23-0848 T	Chrysalidocarpus lutescens	China	OR287501	OR493925	OR493908	OR493891	OR453377	–	–	–	Zhang et al. (2023)
C. chrysalidocarpi	ZHKUCC 23-0847	Chrysalidocarpus lutescens	China	OR287500	OR493924	OR493907	OR493890	OR453376	–	–	–	Zhang et al. (2023)
C. chrysanthemi	IMI 364540 T	Chrysanthemum coronarium	China	JQ948273	JQ948603	JQ948934	JQ949594	JQ949924	JQ949264	–	–	Damm et al. (2012)
C. clavatum	IMI 398854 T	Olea europaea	Italy	JN121126	–	–	–	JN121213	–	–	–	Faedda et al. (2011)
C. cosmi	CBS 853.73 T	Cosmos sp.	Netherlands	JQ948274	JQ948604	JQ948935	JQ949595	JQ949925	JQ949265	–	–	Damm et al. (2012)
C. costaricense	CBS 330.75 T	Coffea arabica	Costa Rica	JQ948180	JQ948510	JQ948841	JQ949501	JQ949831	JQ949171	–	–	Damm et al. (2012)
C. cuscutae	IMI 304802 T	Cuscuta sp.	Dominica	JQ948195	JQ948525	JQ948856	JQ949516	JQ949846	–	–	–	Damm et al. (2012)
C. eriobotryae	GLMC 1935 T	Eriobotrya japonica	China	MF772487	MF795423	MN191653	MN191648	MF795428	MN191658	–	–	Damm et al. (2020)
C. eriobotryae	GLMC 1936	Eriobotrya japonica	China	MF772488	MF795424	MN191654	MN191649	MF795429	MN191659	–	–	Damm et al. (2020)
C. filicis	CBS 101611	Fern	Costa Rica	JQ948196	JQ948526	JQ948857	JQ949517	–	–	–	–	Damm et al. (2012)
C. fioriniae	GUCC 25-0050	Parthenocissus tricuspidata	China	PV768508	PV979835	PV979804	PV979756	PV979905	PV979867	PV979784	–	In this sudy
C. fioriniae	CBS 128517 T	Fiorinia externa	USA	JQ948292	JQ948622	JQ948953	JQ949613	JQ949943	JQ949283	–	–	Damm et al. (2012)
C. fioriniae	CBS 129948	Tulipa sp.	UK	JQ948344	JQ948674	JQ949005	JQ949665	JQ949995	JQ949335	–	–	Damm et al. (2012)
C. fioriniae	CBS 119293	Vaccinium corymbosum	New Zealand	JQ948314	JQ948644	JQ948975	JQ949635	JQ949965	JQ949305	–	–	Damm et al. (2012)
C. fioriniae	IMI 363003, CPC 18928	Camellia reticulata	China	JQ948339	JQ948669	JQ949000	JQ949660	JQ949990	JQ949330	–	–	Damm et al. (2012)
C. godetiae	CBS 133.44 T	Clarkia hybrida	Denmark	JQ948402	JQ948733	JQ949063	JQ949723	JQ950053	JQ949393	–	–	Damm et al. (2012)
C. guajavae	IMI 350839 T	Psidium guajava	India	JQ948270	JQ948600	JQ948931	JQ949591	JQ949921	JQ949261	–	–	Damm et al. (2012)
C. indonesiense	CBS 127551 T	Eucalyptus sp.	Indonesia	JQ948288	JQ948618	JQ948949	JQ949609	JQ949939	JQ949279	–	–	Damm et al. (2012)
C. javanense	CBS 144963a T	Capsicum annuum	Indonesia	MH846576	MH846572	MH846573	MH846575	MH846574	MH846571	–	–	de Silva et al. (2019)
C. johnstonii	CBS 128532 T	Solanum lycopersicum	New Zealand	JQ948444	JQ948775	JQ949105	JQ949765	JQ950095	JQ949435	–	–	Damm et al. (2012)
C. kinghornii	CBS 198.35 T	Phormium sp.	UK	JQ948454	JQ948785	JQ949115	JQ949775	JQ950105	JQ949445	–	–	Damm et al. (2012)
C. kniphofiae	CBS 143496 T	Kniphofia uvaria	UK	MH107884	MH107998	MH107990	MH107975	MH108037	–	–	–	Crous et al. (2018)
C. laticiphilum	CBS 112989 T	Hevea brasiliensis	India	JQ948289	JQ948619	JQ948950	JQ949610	JQ949940	JQ949280	–	–	Damm et al. (2012)
C. limetticola	CBS 114.14 T	Citrus aurantiifolia	USA, Florida	JQ948193	JQ948523	JQ948854	JQ949514	JQ949844	JQ949184	–	–	Damm et al. (2012)
C. lupini	CBS 109225 T	Lupinus albus	Ukraine	JQ948155	JQ948485	JQ948816	JQ949476	JQ949806	–	–	–	Damm et al. (2012)
C. melonis	CBS 159.84 T	Cucumis melo	Brazil	JQ948194	JQ948524	JQ948855	JQ949515	JQ949845	JQ949185	–	–	Damm et al. (2012)
C. miaoliense	NTUCC 20-001-1 T	Fragaria × ananassa	China	MK908419	MK908470	MK908522	MK908573	MK908624	–	–	–	Chung et al. (2020)
C. nagasakiense	NBRC 116456 T	Petroselinum crispum	Japan	LC718411	LC722608	LC722651	LC722733	LC722773	LC722691	–	–	Takata et al. (2024)
C. nymphaeae	GUCC 25-0047	Rotten wood	China	PV768507	PV979834	PV979803	PV979755	PV979904	PV979866	–	–	In this sudy
C. nymphaeae	GUCC 25-0056	Rotten wood	China	PV768509	PV979836	PV979805	PV979757	PV979906	PX213640	–	–	In this sudy
C. nymphaeae	CBS 515.78 T	Nymphaea alba	Netherlands	JQ948197	JQ948527	JQ948858	JQ949518	JQ949848	JQ949188	–	–	Damm et al. (2012)
C. nymphaeae	CBS 134234	Citrus aurantiifolia	China	KC293582	KC293742	KY856139	KY855974	KC293662	KY856310	–	–	Guarnaccia et al. (2017)
C. nymphaeae	CBS 516.78	Nuphar luteum, leaf spot	Netherlands	JQ948198	JQ948528	JQ948859	JQ949519	JQ949849	JQ949189	–	–	Damm et al. (2012)
C. nymphaeae	CGMCC 3.15228	Nuphar lutea subsp. polysepala	USA, Florida	KC293581	KC293741	KY856138	KY855973	KC293661	KY856309	–	–	Liu et al. (2013)
C. nymphaeae (=C. speciosum)	YMF 1.07301 T	Ageratina adenophora	China	OK030881	–	–	–	–	–	–	–	Sui et al (2024)
C. nymphaeae (≡C. simulanticitri)	YMF 1.07302	Betula spp.	China	OK030878	OK513680	OK513577	OK513615	–	–	–	–	Yu et al. (2022)
C. paranaense	CBS 134729 T	Malus domestica	Brazil	KC204992	KC205026	KC205043	KC205077	KC205060	KC205004	–	–	Braganca et al. (2016)
C. paxtonii	IMI 165753 T	Musa sp.	Saint Lucia	JQ948285	JQ948615	JQ948946	JQ949606	JQ949936	JQ949276	–	–	Damm et al. (2012)
C. perseicola	RGM 3376 T	Persea lingue	Chile	OR644585	OR644992	OR645045	OR645098	OR645150	OR659723	–	–	Zapata et al. (2024)
C. phormii	CBS 118194 T	Phormium sp.	Germany	JQ948446	JQ948777	JQ949107	JQ949767	JQ950097	JQ949437	–	–	Damm et al. (2012)
C. pyricola	CBS 128531 T	Pyrus communis	New Zealand	JQ948445	JQ948776	JQ949106	JQ949766	JQ950096	JQ949436	–	–	Damm et al. (2012)
C. rhombiforme	CBS 129953 T	Olea europaea	Portugal	JQ948457	JQ948788	JQ949118	JQ949778	JQ950108	JQ949448	–	–	Damm et al. (2012)
C. roseum	CBS 145754 T	Lapageria rosea	Chile	MK903611	MK903603	–	MK903604	MK903607	OR659735	–	–	Crous et al. (2019)
C. salicis	CBS 607.94 T	Salix sp.	Netherlands	JQ948460	JQ948791	JQ949121	JQ949781	JQ950111	JQ949451	–	–	Damm et al. (2012)
C. schimae	LC13880 T	Schima sp.	China	MZ595885	MZ664105	MZ799347	MZ664183	MZ674003	MZ673905	–	–	Liu et al. (2022)
C. schimae	LC13881	Schima sp.	China	MZ595887	MZ664106	MZ799348	MZ664185	MZ674005	MZ673907	–	–	Liu et al. (2022)
C. schimae	PC9	Carica papaya	China	OQ642139	OQ723034	OQ723037	OQ723035	OQ723036	–	–	–	Direct Submission
C. schimae	CNUCC 324C-1-5-2	Camellia sinensis var. assamica	China	PP809328	PP832118	PP843051	PP824745	PP832196	PP832157	–	–	Sui et al (2024)
C. schimae	CNUCC 528-2-2	Ilex chinensis	China	PP809337	PP832126	PP843060	PP824754	PP832205	PP832166	–	–	Sui et al (2024)
C. scovillei	CBS 126529 T	Capsicum sp.	Indonesia	JQ948267	JQ948597	JQ948928	JQ949588	JQ949918	JQ949258	–	–	Damm et al. (2012)
C. simmondsii	CBS 122122 T	Carica papaya	Australia	JQ948276	JQ948606	JQ948937	JQ949597	JQ949927	JQ949267	–	–	Damm et al. (2012)
C. sloanei	IMI 364297 T	Theobroma cacao	Malaysia	JQ948287	JQ948617	JQ948948	JQ949608	JQ949938	JQ949278	–	–	Damm et al. (2012)
C. subsalicis	LC13863 T	Populus alba	China	MZ852849	–	MZ799346	MZ664128	MZ673953	MZ673836	–	–	Sui et al (2024)
C. tamarilloi	CBS 129814 T	Solanum betaceum	Colombia	JQ948184	JQ948514	JQ948845	JQ949505	JQ949835	JQ949175	–	–	Damm et al. (2012)
C. walleri	CBS 125472 T	Coffea sp.	Vietnam	JQ948275	JQ948605	JQ948936	JQ949596	JQ949926	JQ949266	–	–	Damm et al. (2012)
C. wanningense	CGMCC 3.18936 T	Rubber tree	China	MG830462	MG830318	MG830302	MG830270	MG830286	–	–	–	Sui et al (2024)
C. trichellum (outgroup)	CBS 217.64 T	Hedera helix	UK	GU227812	GU228204	GU228302	GU227910	GU228106	–	–	–	Damm et al. (2009)
C. rusci (outgroup)	CBS 119206 T	Ruscus sp.	Italy	GU227818	GU228210	GU228308	GU227916	GU228112	GU228014	–	–	Damm et al. (2009)
Colletotrichum bambusicola, coccodes, trichellum species complexes
C. bambusicola	CNUCC 307307 T	Phyllostachys edulis	China	MT199632	MT192844	MT192871	MT188638	MT192817	–	–	–	Wang et al. (2021)
C. coccodes	CBS 369.75 T	Solanum tuberosum	The Netherlands	HM171679	HM171673	JQ005796	HM171667	JQ005859	JX546779	–	–	Liu et al. (2011)
C. guangxiense	CNUCC 310138 T	Phyllostachys edulis	China	MT199633	MT192834	MT192861	MT188628	MT192805	–	–	–	Wang et al. (2021)
C. hsienjenchang	MAFF 243051 T	Phyllostachys bambusoides	Japan	AB738855	–	AB738846	AB738845	–	AB738847	–	–	Sato et al. (2012)
C. metake	MAFF 244029 T	Pleioblastus simonii	Japan	AB738859	–	–	–	OK236390	–	–	–	Sato et al. (2012)
C. metake	MAFF 243970	Pleioblastus simonii	Japan	OK090430	–	–	–	OK236391	–	–	–	Liu et al. (2022)
C. metake	CNUCC 311173	Chimonobambusa quadrangularis	China	MT192601	MT192858	MT188625	MT188652	MT192831	–	–	–	Wang et al. (2021)
C. metake	MAFF 241800	Pleioblastus simonii	Japan	OK090431	–	OK236386	OK236388	OK236392	–	–	–	Liu et al. (2022)
C. metake	GUCC 25-0038	Bamboo	China	PV791792	–	PV979808	PV979758	PV979909	PV979871	PV979785	–	In this study
C. nigrum	CBS 169.49 T	Capsicum sp.	Argentina	JX546838	JX546742	JX546693	JX546646	JX546885	JX546791	–	–	Liu et al. (2013)
C. nigrum (=C. dianense)	CGMCC 3.18943	Alternanthera philoxeroides	China	–	PP482514	PP482513	PP482512	PP482516	PP482515	–	–	Chang et al. (2024)
C. nigrum (=C. dianense)	YMF 1.04943	Alternanthera philoxeroides	China	OL842189	OL981284	OL981310	OL981258	–	–	–	–	Zheng et al. (2022)
C. obovoides	LC6085 T	unidentified plant, leaf	China	MZ595838	–	MZ799345	MZ664136	MZ673959	MZ673857	–	–	Liu et al. (2022)
C. parabambusicola	LC13884 T	Bamboo, dead culm	China	MZ595904	MZ664098	MZ799338	MZ664202	MZ674022	MZ673924	–	–	Liu et al. (2022)
C. rusci	CBS 119206 T	Ruscus sp.	Italy	GU227818	GU228210	GU228308	GU227916	GU228112	GU228014	–	–	Liu et al. (2013)
C. trichellum	CBS 217.64 T	Hedera helix	Germany	GU227812	GU228204	GU228302	GU227910	GU228106	–	–	–	Liu et al. (2013)
C. trichellum	CBS 118198	Hedera sp.	UK	GU227813	GU228205	GU228303	GU227911	GU228107	–	–	–	Liu et al. (2013)
C. trichellum	GUCC 25-0039	Hedera sp.	China	PV791793	PV979841	PV979809	PV979759	PV979910	PV979872	–	–	In this study
C. acutatum (outgroup)	CBS 112996 T	Carica sp.	Australia	JQ005776	JQ948677	JQ005797	JQ005839	JQ005860	JQ005818	–	–	Liu et al. (2013)
C. fioriniae (outgroup)	CBS 128517 T	Fiorinia externa	USA	JQ948292	JQ948622	JQ948953	JQ949613	JQ949943	JQ949283	–	–	Damm et al. (2012)
Colletotrichum gloeosporioides species complex
C. aenigma	ICMP 18608 T	Persea americana	Israel	JX010244	JX010044	JX009774	JX009443	JX010389	–	JX009683	KM360143	Weir et al. (2012)
C. aeschynomenes	ICMP 17673 T	Aeschynomene virginica	USA	JX010176	JX009930	JX009799	JX009483	JX010392	–	JX009721	KM360145	Weir et al. (2012)
C. alatae	CBS 304.67 T	Dioscorea alata	India	JX010190	JX009990	JX009837	JX009471	JX010383	–	JX009738	KC888932	Weir et al. (2012)
C. alienum	ICMP 12071 T	Malus domestica	New Zealand	JX010251	JX010028	JX009882	JX009572	JX010411	–	JX009654	KM360144	Weir et al. (2012)
C. aotearoa	ICMP 18537 T	Coprosma sp.	New Zealand	JX010205	JX010005	JX009853	JX009564	JX010420	–	JX009611	KC888930	Weir et al. (2012)
C. arecicola	CGMCC 3.19667 T	Areca catechu	China	MK914635	MK935455	MK935541	MK935374	MK935498	–	–	MK935413	Cao et al. (2020)
C. artocarpicola	MFLUCC 18-1167 T	Artocarpus	Thailand	MN415991	MN435568	MN435569	MN435570	MN435567	–	–	–	Bhunjun et al. (2019)
C. asianum	ICMP 18580 T	Coffea arabica	Thailand	FJ972612	JX010053	JX009867	JX009584	JX010406	–	FJ917506	FR718814	Zhang et al. (2023)
C. australianum	VPRI 43075 T	Citrus sinensis	Australia, Vic	MG572138	MG572127	MW091987	MN442109	MG572149	–	–	MG572171	Wang et a. (2021)
C. camelliae	CGMCC 3.14925 T	Camellia sinensis	China	KJ955081	KJ954782		KJ954363	KJ955230	–	KJ954634	KJ954497	Wang et a. (2021)
C. camelliae (≡ C. analogum)	YMF1.06943 T	Ageratina adenophora	China	OK030860	OK513663	OK513559	OK513599	OK513629	–	–	PP498774	Yu et al. (2022)
C. cangyuanense	YMF1.05001T	Ageratina adenophora	China	OK030864	OK513667	OK513563	OK513603	OK513633	–	–	–	Yu et al. (2022)
C. castaneae	GUCC 21268.4 T	Castanea mollissima	China	OP722991	OP737973	OP715778	OP715812	OP720868	–	–	–	Zhang et al. (2023)
C. changpingense	MFLUCC 15-0022 T	Fragaria × ananassa	China	KP683152	KP852469	KP852449	KP683093	KP852490	–	–	–	Wang et a. (2021)
C. chiangmaiense	MFLUCC 18-0945 T	Magnolia garrettii	Thailand	MW346499	MW548592	MW623653	MW655578		–	–	–	Zhang et al. (2023)
C. chrysophilum	CMM4268 T	Musa sp.	Brazil	KX094252	KX094183	KX094083	KX093982	KX094285	–	KX094063	KX094325	Wang et a. (2021)
C. cigarro	ICMP 18539 T	Olea europaea	Australia	JX010230	JX009966	JX009800	JX009523	JX010434	–	JX009635	–	Weir et al. (2012)
C. clidemiae	ICMP 18658 T	Clidemia hirta	USA, Hawaii	JX010265	JX009989	JX009877	JX009537	JX010438	–	JX009645	KC888929	Weir et al. (2012)
C. cobbittiense	BRIP 66219a T	Magnolia garrettii	Thailand	MH087016	MH094133	MH094135	MH094134	MH094137	–	–	–	Zhang et al. (2023)
C. conoides	CAUG17 T	Capsicum annuum	China	KP890168	KP890162	KP890156	KP890144	KP890174	–	KP890150	–	Wang et a. (2021)
C. cordylinicola	ICMP 18579 T	Cordyline fruticosa	Thailand	JX010226	JX009975	JX009864	HM470235	JX010440	–	HM470238	JQ899274	Weir et al. (2012)
C. cycadis	BRIP 71326a T	Cycas revoluta	China	MT439915	MT439919	MT439917		MT439921	–	–		Zhang et al. (2023)
C. dimorphum	YMF1.07309 T	Ageratina adenophora	China	OK030867	OK513670	OK513566	OK513606	OK513636	–	–	PP498776	Yu et al. (2022)
C. dracaenigenum	MFLUCC 19-0430 T	Dracaena sp.	Thailand	MN921250	MT215577	MT215575	MT313686		–	–	–	Zhang et al. (2023)
C. endophyticum	MFLUCC 13-0418 T	Pennisetum purpureum	Thailand	KC633854	KC832854	MZ799261	KF306258	MZ673954	–	KC810018	–	Zhang et al. (2023)
C. fici-septicae	MFLUCC 20-0166 T	Ficus septica	China	MW114367	MW183774	MW177701	MW151585		–	–	–	Zhang et al. (2023)
C. fructicola	MFLUCC 22-0181	Pineapple	Thailand	OQ048649	OQ067350	OQ067349	OQ067348	OQ067351	–	–	–	Armand et al. (2023a)
C. fructicola	MFLUCC 22-0182	Pineapple	Thailand	OQ048650	OQ067354	OQ067353	OQ067352	OQ067355	–	–	–	Armand et al. (2023a)
C. fructicola	MFLUCC 17-1752	Rhizophora apiculata	Thailand	OR828931	OR840868	OR840856	OR840845	OR840862	–	OR840851	–	Norphanphoun and Hyde (2023)
C. fructicola	MFLUCC 17-1753	Rhizophora apiculata	Thailand	OR828932	OR840869	OR840857	OR840846	OR840863	–	OR840852	–	Norphanphoun and Hyde (2023)
C. fructicola	ICMP 18581 T	Coffea arabica	Thailand	JX010165	JX010033	JX009866	FJ907426	JX010405	–	FJ917508	JQ807838	Weir et al. (2012)
C. fructicola	ICMP 18613	Limonium sinuatum	Israel	JX010167	JX009998	JX009772	JX009491	JX010388	–	JX009675	–	Weir et al. (2012)
C. fructicola	ICMP 18727	Fragaria × ananassa	USA	JX010179	JX010035	JX009812	JX009565	JX010394	–	JX009682	–	Weir et al. (2012)
C. fructicola	GUCC 25-0021	Juglans regia	China	PV791817	PV979865	PV979833	PV979783	PV979934	PV979896	PV979786	PV979935	In this study
C. fructicola	GUCC 25-0022	Juglans regia	China	PV791816	PV979864	PV979832	PV979782	PV979933	PV979895	PV979787	PV979936	In this study
C. fructicola	GUCC 25-0023	Juglans regia	China	PV791815	PV979863	PV979831	PV979781	PV979932	PV979894	PV979788	PV979937	In this study
C. fructicola	GUCC 25-0024	Juglans regia	China	PV791814	PV979862	PV979830	PV979780	PV979931	PV979893	PV979789	PV979938	In this study
C. fructicola	GUCC 25-0025	Juglans regia	China	PV791813	PV979861	PV979829	PV979779	PV979930	PV979892	PV979790	PV979939	In this study
C. fructicola	GUCC 25-0045	Actinidia chinensis var. deliciosa	China	PV791799	PV979847	PV979815	PV979765	PV979916	PV979878	PV979800	PV979947	In this study
C. fructicola	GUCC 25-0051	Rosa chinensis	China	PV791796	PV979844	PV979812	PV979762	PV979913	PV979875	PV979801	–	In this study
C. fructivorum	CBS 124.22	n/a	USA	MH854714	–	–	–	JX145176	–	–	–	Vu et al. (2019)
C. fructivorum	CBS 133125 T	Vaccinium macrocarpon	USA	JX145145	MZ664047	MZ799259	MZ664126	JX145196	–	–	JX145300	Wang et a. (2021)
C. gardeniae	GUCC 12049 T	Gardenia jasminoides	China	OP722995	OP737963	OP715766	OP715801	OP720858	–	–	–	Zhang et al. (2023)
C. gardeniae	GUCC 12048	Gardenia jasminoides	China	OP722989	OP737962	OP715765	OP715800	OP720857	–	–	–	Zhang et al. (2023)
C. gardeniae	GUCC 12047	Gardenia jasminoides	China	OP722964	OP737961	OP715764	OP715799	OP720856	–	–	–	Zhang et al. (2023)
C. gloeosporioides	IMI 356878 T	Citrus sinensis	Italy	JQ005152	JQ005239	JQ005326	JQ005500	JQ005587	–	–	–	Damm et al. (2012)
C. gloeosporioides	LGMF800	Commelina benghalensis	Rincão	KM278577	KM257052	KJ579936	KJ569194	KJ579891	–	–	–	Waculicz-Andrade et al. (2017)
C. gloeosporioides	LGMF748	Sida rhombifolia	Rincão	KM257023	KM257047	KJ579931	KJ569189	KJ579839	–	–	–	Waculicz-Andrade et al. (2017)
C. gloeosporioides	LGMF524	Citrus	Mogi Guaçú	JQ580610	JQ580791	KJ579927	KJ569185	KJ579739	–	–	–	Waculicz-Andrade et al. (2017)
C. gloeosporioides	ICMP 18730	Citrus sp.	New Zealand	JX010157	JX009981	JX009861	JX009548		–	–	–	Weir et al. (2012)
C. gloeosporioides	CBS 112999 T	Citrus sinensis	Italy	JX010152	JX010056	JX009818	JX009531	JX010445	–	JX009731	JQ807843	Weir et al. (2012)
C. gloeosporioides	ICMP 19121	Citrus limon	Italy	JX010148	JX010054	JX009903	JX009558		–	–	–	Weir et al. (2012)
C. gloeosporioides	GUCC 25-0041	Camellia Japonica	China	PV791803	PV979851	PV979819	PV979769	PV979920	PV979882	–	–	In this study
C. gloeosporioides	GUCC 25-0042	Camellia Japonica	China	PV791802	PV979850	PV979818	PV979768	PV979919	PV979881	–	–	In this study
C. grevilleae	CBS 132879 T	Grevillea sp.	Italy	KC297078	KC297010	KC296987	KC296941	KC297102	–	KC296963	–	Wang et a. (2021)
C. grossi	CAUG7 T	Capsicum sp.	China	KP890165	KP890159	KP890153	KP890141	KP890171	–	KP890147	–	Wang et a. (2021)
C. hebeiense	MFLUCC 13-0726 T	Vitis vinifera cv. Cabernet Sauvignon	China	KF156863	KF377495	KF289008	KF377532	KF288975	–	–	KF377562	Wang et a. (2021)
C. hederiicola	MFLU 15-0689 T	Hedera helix	Italy	MN631384	–	MN635794	MN635795	–	–	–	–	Zhang et al. (2023)
C. helleniense	CBS 142418 T	Poncirus trifoliata	Greece	KY856446	KY856270	KY856186	KY856019	KY856528	–	KY856099	–	Wang et a. (2021)
C. henanense	CGMCC 3.17354 T	Camellia sinensis	China	KJ955109	KJ954810	–	KM023257	KJ955257	–	KJ954662	KJ954524	Wang et a. (2021)
C. horii	ICMP 10492 NT	Diospyros kaki	Japan	GQ329690	GQ329681	JX009752	JX009438	JX010450	–	JX009604	JQ807840	Wang et a. (2021)
C. hystricis	CBS 142411 T	Citrus hystrix	Italy	KY856450	KY856274	KY856190	KY856023	KY856532	–	KY856103	–	Wang et a. (2021)
C. jiangxiense	22N642	n/a	South Korea	OR805434	OR826297	OR826295	–	OR826299	–	–	–	Direct Submission
C. jiangxiense	SYD-9	n/a	China	OR467495	OR472539	OR472537	OR472535	OR472541	–	–	–	Direct Submission
C. jiangxiense	SYD-4	n/a	China	OR467494	OR472538	OR472536	OR472534	OR472540	–	–	–	Direct Submission
C. jiangxiense	CGMCC 3.17363 T	Camellia sinensis	China	KJ955201	KJ954902	–	KJ954471	KJ955348	–	KJ954752	KJ954607	Liu et al. (2015)
C. jiangxiense	GUCC 25-0027	Juglans regia	China	PV791811	PV979859	PV979827	PV979777	PV979928	PV979890	PV979792	PV979940	In this study
C. jiangxiense	GUCC 25-0031	Juglans regia	China	PV791808	PV979856	PV979824	PV979774	PV979925	PV979887	PV979795	PV979943	In this study
C. jiangxiense	GUCC 25-0052	Pteris henryi	China	PV791795	PV979843	PV979811	PV979761	PV979912	PV979874	–	–	In this study
C. jiangxiense	GUCC 25-0053	Parthenocissus tricuspidata	China	PV791794	PV979842	PV979810	PV979760	PV979911	PV979873	PV979802	PV979948	In this study
C. jiangxiense (=C. gracile)	YMF 1.07329	Ageratina adenophora	China	OK030869	OK513672	OK513568	OK513608	OK513638	–	–	–	Yu et al. (2022)
C. jiangxiense (=C. gracile)	YMF1.06939 T	Ageratina adenophora	China	OK030868	OK513671	OK513567	OK513607	OK513637	–	–	PP498777	Yu et al. (2022)
C. jiangxiense (=C. nullisetosum)	YMF1.06946 T	Mango	China	OK030872	OK513675	OK513571	OK513611	OK513641	–	–	PP498779	Yu et al. (2022)
C. jiangxiense (=C. nullisetosum)	YMF1.07328	Mango	China	OK030873	OK513676	OK513572	OK513612	OK513642	–	–	–	Yu et al. (2022)
C. jiangxiense (=C. oblongisporum)	YMF1.06938 T	Ageratina adenophora	China	OK030874	OK513677	OK513573	–	OK513643	–	–	PP498780	Yu et al. (2022)
C. jiangxiense (=C. oblongisporum)	YMF 1.07326	Ageratina adenophora	China	OK030875	OK513678	OK513574	–	OK513644	–	–	–	Yu et al. (2022)
C. juglandicola	CGMCC3.24312 T	Juglans regia	China	OQ263015	OQ282973	OR004793	OQ282966	OQ282980	–	–	–	Zhang et al. (2023)
C. juglandicola	CGMCC3.24313	Juglans regia	China	OQ263018	OQ282977	OR004797	OQ282970	OQ282984	–	–	–	Zhang et al. (2023)
C. juglandicola	GUCC 25-0043	Camellia Japonica	China	PV791801	PV979849	PV979817	PV979767	PV979918	PV979880	PV979798	PV979945	In this study
C. juglandicola	GUCC 25-0044	Ilex sp.	China	PV791800	PV979848	PV979816	PV979766	PV979917	PV979879	PV979799	PV979946	In this study
C. kahawae	CIFC Uga7	Coffea arabica	Uganda: Kapchorwa	HE655531	–	–	–	HE655592	–	–	HE657410	Silva et al. (2012)
C. kahawae	ICMP 17816 T	Coffea arabica	Kenya	JX010231	JX010012	JX009813	JX009452	JX010444	–	JX009642	JQ894579	Weir et al. (2012)
C. kahawae	ICMP:17811	Coffea arabica	Malawi	JX010233	JX009970	JX009817	JX009555	JX010430	–	JX009641	OQ871550	Weir et al. (2012)
C. kunmingense	GUCC 12053 T	Ophiopogon japonicus	China	OP722975	OP737965	OP715769	OP715804	OP720861	–	–	–	Zhang et al. (2023)
C. ledongense	CGMCC3.18888 T	Hevea brasiliensis	China	MG242009	MG242017	MG242019	MG242015	MG242011	–	MG242013	–	Liu et al. (2018)
C. ligustri	GUCC 12111 T	Ilex chinensis	China	OP722988	OP737968	OP715773	OP740216	OP720864	–	–	–	Zhang et al. (2023)
C. macroconidii	GUCC 25-0028 T	Juglans regia	China	PV791810	PV979858	PV979826	PV979776	PV979927	PV979888	PV979793	PV979941	In this study
C. macroconidii	GUCC 25-0063	Juglans regia	China	PV791809	PV979857	PV979825	PV979775	PV979926	PV979889	PV979794	PV979942	In this study
C. makassarense	CBS 143664 T	Capsicum annuum	Indonesia	MH728812	MH728820	MH805850	MH781480	MH846563	–	–	MH728831	Zhang et al. (2023)
C. menglaense	YMF1.04960 T	Air	China	MH023505	MH023507	MH023508	MH023506		–	–	–	Zhang et al. (2023)
C. musae	CBS 116870 T	Musa sp.	USA	JX010146	JX010050	JX009896	JX009433	HQ596280	–	JX009742	KC888926	Weir et al. (2012)
C. nanhuaense	YMF1.04993 T	Ageratina adenophora	China	OK030870	OK513673	OK513569	OK513609	OK513639	–	–	PP498778	Yu et al. (2022)
C. nanhuaense	GUCC 25-0026	Juglans regia	China	PV791812	PV979860	PV979828	PV979778	PV979929	PV979891	PV979791	–	In this study
C. nupharicola	CBS 470.96 T	Nuphar lutea subsp. polysepala	USA	JX010187	JX009972	JX009835	JX009437	JX010398	–	JX009663	JX145319	Weir et al. (2012)
C. perseae	CBS 141365 T	Persea americana	Israel	KX620308	KX620242	–	KX620145	KX620341	–	KX620206	KX620177	Sharma et al. (2017)
C. proteae	CBS 132882 T	Protea sp.	South Africa	KC297079	KC297009	KC296986	KC296940	KC297101	–	KC296960	–	Wang et a. (2021)
C. pseudotheobromicola	MFLUCC 18-1602 T	Avocado	Israel	MH817395	MH853675	MH853678	MH853681	MH853684	–	–	–	Zhang et al. (2023)
C. psidii	CBS 145.29 T	Psidium sp.	Italy	JX010219	JX009967	JX009901	JX009515	JX010443	–	JX009743	KC888931	Weir et al. (2012)
C. queenslandicum	ICMP 1778 T	Carica papaya	Australia	JX010276	JX009934	JX009899	JX009447	JX010414	–	JX009691	KC888928	Weir et al. (2012)
C. rhexiae	CBS 133134 T	Rhexia virginica	USA	JX145128	MZ664046	MZ799258	MZ664127	JX145179	–	–	JX145290	Zhang et al. (2023)
C. rhizophorae	MFLUCC 17-1927 T	Rhizophora apiculata	Thailand	OR828933	OR840870	OR840858	OR840847	OR840864	–	OR840853	–	Norphanphoun and Hyde (2023)
C. salsolae	ICMP 19051 T	Salsola tragus	Hungary	JX010242	JX009916	JX009863	JX009562	JX010403	–	JX009696	KC888925	Weir et al. (2012)
C. siamense	MFLUCC 18-1162	Rosa sp.	Thailand	MN788676	MN995328	MN995335	MN995334	MN995329	–	–	–	Hyde et al. (2020)
C. siamense	MFLUCC 22-0138	Cyclosorus sp.	Thailand	OP802366	OP801723	OP801705	OP801688	OP801742	–	–	–	Seifollahi et al. (2023)
C. siamense	ICMP 18578 T	Coffea arabica	Thailand	JX010171	JX009924	JX009865	FJ907423	JX010404	–	FJ917505	JQ899289	Weir et al. (2012)
C. siamense	ICMP 18572	Vitis vinifera	USA	JX010160	JX010061	JX009783	JX009487	–	–	–	–	Weir et al. (2012)
C. siamense	ICMP 18739	Carica papaya	South Africa	JX010161	JX009921	JX009794	JX009484	–	–	–	–	Weir et al. (2012)
C. siamense	ICMP 18571	Fragaria × ananassa	USA	JX010159	JX009922	JX009782	JX009482	–	–	–	–	Weir et al. (2012)
C. siamense	MFLUCC 17-0571	Pandanaceae	Thailand	MG646967	MG646934	MG646931	MG646938	MG646926	–	–	–	Wang et a. (2021)
C. siamense (= C. parvisporum)	SAUCC 201152	n/a	China	MW786746	MW876478	MW883693	MW883702	MW888977	–	–	–	Direct Submission
C. siamense (= C. parvisporum)	SAUCC 200204	n/a	China	MW786641	MW846239	MW883685	MW883694	MW888969	–	–	–	Direct Submission
C. siamense (= C. parvisporum)	MFLUCC 22-0151	Cyclosorus sp., leaf	Thailand	OP802371	OP801726	OP801708	OP801691	OP801746	–	–	–	Seifollahi et al. (2023)
C. siamense (= C. parvisporum)	MFLUCC 22-0164	Cyclosorus sp., leaf	Thailand	OP802369	OP801724	OP801706	OP801689	OP801744	–	–	–	Seifollahi et al. (2023)
C. siamense (= C. parvisporum)	MFLUCC 22-0159	Nephrolepis cordifolia	Thailand	OP802373	OP801727	OP801709	OP801692	OP801747	–	–	–	Seifollahi et al. (2023)
C. siamense (= C. parvisporum)	YMF 1.06942	Ageratina adenophora	China	OK030876	OK513679	OK513575	OK513613	OK513645	–	–	–	Yu et al. (2022)
C. siamense	GUCC 25-0032	Juglans regia	China	PV791807	PV979855	PV979823	PV979773	PV979924	PV979886	PV979796	PV979944	In this study
C. siamense	GUCC 25-0033	Pteridophyta	China	PV791806	PV979854	PV979822	PV979772	PV979923	PV979885	PV979797	–	In this study
C. siamense	GUCC 25-0034	Pteridophyta	China	PV791805	PV979853	PV979821	PV979771	PV979922	PV979884	–	–	In this study
C. siamense	GUCC 25-0035	Pteridophyta	China	PV791804	PV979852	PV979820	PV979770	PV979921	PV979883	–	–	In this study
C. siamense	GUCC 25-0048	Parthenocissus quinquefolia	China	PV791798	PV979846	PV979814	PV979764	PV979915	PV979877	–	–	In this study
C. siamense	GUCC 25-0049	Parthenocissus tricuspidata	China	PV791797	PV979845	PV979813	PV979763	PV979914	PV979876	–	–	In this study
C. subhenanense	YMF1.06865 T	Ageratina adenophora	China	OK030883	OK513684	OK513581	OK513618	OK513647	–	–	PP498782	Yu et al. (2022)
C. syzygiicola	MFLUCC 10-0624 T	Syzygium samarangense	Thailand	KF242094	KF242156	–	KF157801	KF254880	–	KF254859	–	Wang et a. (2021)
C. tainanense	CBS 143666 T	Capsicum annuum	China	MH728818	MH728823	MH805845	MH781475	MH846558	–	–	MH728836	Zhang et al. (2023)
C. temperatum	CBS 133122 T	Vaccinium macrocarpon	USA	JX145159	MZ664045	MZ799254	MZ664125	JX145211	–	–	JX145298	Wang et a. (2021)
C. tengchongense	YMF 1.04950 T	Isoetes sinensis	China	OL842169	OL981264	OL981290	PP498771	PP498785	–	PP498784	PP498773	Zhang et al. (2023)
C. thailandica	MFLUCC 17-1924 T	Rhizophora apiculata	Thailand	OR828935	OR840872	OR840860	OR840849	OR840866	–	OR840855		Norphanphoun and Hyde (2023)
C. theobromicola	CBS 124945 T	Theobroma cacao	Panama	JX010294	JX010006	JX009869	JX009444	JX010447	–	JX009591	KC790726	Weir et al. (2012)
C. ti	ICMP 4832 T	Cordyline sp.	New Zealand	JX010269	JX009952	JX009898	JX009520	JX010442	–	JX009649	KM360146	Weir et al. (2012)
C. tropicale	CBS 124949 T	Theobroma cacao	Panama	JX010264	JX010007	JX009870	JX009489	JX010407	–	JX009719	KC790728	Weir et al. (2012)
C. viniferum	GZAAS5.08601 T	Vitis vinifera, cv. ‘Shuijing’	China	JN412804	JN412798	–	JN412795	JN412813	–	JQ309639		Wang et a. (2021)
C. vulgaris	YMF 1.04940 T	Hippuris vulgaris	China	OL842170	OL981265	OL981291	OL981239	–	–			Zhang et al. (2023)
C. wuxiense	CGMCC 3.17894 T	Camellia sinensis	China	KU251591	KU252045	KU251939	KU251672	KU252200	–	KU251833	KU251722	Wang et a. (2021)
C. xanthorrhoeae	ICMP 17903 T	Xanthorrhoea preissii	Australia	JX010261	JX009927	JX009823	JX009478	JX010448	–	JX009653	KC790689	Weir et al. (2012)
C. xishuangbannaense	MFLUCC 19-0107 T	Magnolia liliifera	China	MW346469	MW537586	MW660832	MW652294		–	–	–	Zhang et al. (2023)
C. yuanjiangense	YMF1.04996 T	Ageratina adenophora	China	OK030885	OK513686	OK513583	OK513620	OK513649	–	–	–	Yu et al. (2022)
C. yulongense	CFCC 50818 T	Vaccinium dunalianum var. urophyllum	China	MH751507	MK108986	MH793605	MH777394	MK108987	–	MH793604	PP861147	Zhang et al. (2023)
C. boninense (outgroup)	CBS 123755 T	Crinum asiaticum var. sinicum	Japan	JQ005153	JQ005240	JQ005327	JQ005501	JQ005588	–	JQ005674	–	Damm et al. (2012)
C. brasiliense (outgroup)	CBS 128501 T	Passiflora edulis	Brazil	JQ005235	JQ005322	JQ005409	JQ005583	JQ005669	–	JQ005756	–	Damm et al. (2012)
Colletotrichum magnum species complex
C. brevisporum	BCC 38876 T	Neoregelia sp.	Thailand	JN050238	JN050227	–	JN050216	JN050244	–	–	–	Noireung et al. (2012)
C. cacao	CBS 119297 T	Theobroma cacao	Costa Rica	MG600772	MG600832	MG600878	MG600976	MG601039	MG600916	–	–	Damm et al. (2019)
C. guangdongense	ZHKUCC 21-0105 T	Citrus maxima	China	OL708415	OL855854	OL855864	OL855875	OL855885	ON315370	–	–	Liu et al. (2023)
C. guangdongense	ZHKUCC 21-0106	Citrus maxima	China	OL708420	OL855855	OL855865	OL855876	OL855886	–	–	–	Liu et al. (2023)
C. kaifengense	CAASZK33 T	Citrullus lanatus	China	MZ475245	OL456715	OL901183	OL449313	OL456674	OM057724	–	–	Guo et al. (2022)
C. kaifengense	CAASZK32	Citrullus lanatus	China	MZ475244	OL456714	OL901182	OL449312	OL456673	OM057723	–	–	Guo et al. (2022)
C. lobatum	IMI 79736 T	Piper catalpaefolium	Trinidad and Tobago	MG600768	MG600828	MG600874	MG600972	MG601035	MG600912	–	–	Damm et al. (2019)
C. magnum	CBS 519.97 T	Citrullus lanatus	USA	MG600769	MG600829	MG600875	MG600973	MG601036	MG600913	–	–	Damm et al. (2019)
C. magnum	IMI391662	Citrullus lanatus	USA	MG600771	MG600831	MG600877	MG600975	MG601038	MG600915	–	–	Damm et al. (2019)
C. magnum	CBS575.97	Citrullus lanatus	USA	MG600770	MG600830	MG600876	MG600974	MG601037	MG600914	–	–	Damm et al. (2019)
C. magnum	CAASZK9	Citrullus lanatus	China	MZ475213	OL899046	OL901142	OL986316	OL988321	OM057685	–	–	Gou et al. (2022)
C. magnum	CAASZK34	Citrullus lanatus	China	MZ475246	OL899044	OL986134	OL986314	OL988319	OL986209	–	–	Gou et al. (2022)
C. magnum	CAASZKF13	Citrullus lanatus	China	MZ475166	OL456677	OL901145	OL449275	OL456636	OM057688	–	–	Gou et al. (2022)
C. magnum	CAASZK2	Citrullus lanatus, fruit	China	MZ475206	OL456679	OL901147	OL449277	OL456638	OM057690	–	–	Gou et al. (2022)
C. magnum (=C. liaoningense)	CGMCC3.17616, CAUOS2	Chili pepper	China	KP890104	KP890135	KP890127	KP890097	KP890111	–	–	–	Diao et al. (2016)
C. magnum (=C. liaoningense)	CAUOS6	Capsicum annuum var. conoides	China	–	–	KP890131	–	KP890115	–	–	–	Diao et al. (2017)
C. magnum (=C. liaoningense)	CAUOS3	Capsicum annuum var. conoides	China	KP890105	KP890136	KP890128	–	KP890112	–	–	–	Diao et al. (2017)
C. magnum (=C. liaoningense)	CAUOS4	Capsicum annuum var. conoides	China	KP890106	KP890137	KP890129	–	KP890113	–	–	–	Diao et al. (2017)
C. magnum	GUCC 25-0029	Juglans regia	China	PV791818	PV979839	PV979806	PV979837	PV979907	PV979869	–	–	In this study
C. magnum	GUCC 25-0030	Juglans regia	China	PV791819	PV979840	PV979807	PV979838	PV979908	PV979870	–	–	In this study
C. merremiae	CBS 124955 T	Merremia umbellata	Panama	MG600765	MG600825	MG600872	MG600969	MG601032	MG600910	–	–	Damm et al. (2019)
C. okinawense	MAFF 240517 T	Carica papaya, petiole	Japan	MG600767	MG600827	–	MG600971	MG601034	–	–	–	Damm et al. (2019)
C. panamense	CBS 125386 T	Merremia umbellata	Panama	MG600766	MG600826	MG600873	MG600970	MG601033	MG600911	–	–	Damm et al. (2019)
C. qilinense	CAASZK13 T	Citrullus lanatus, fruit	China	MZ475217	OL456694	OL901162	OL449292	OL456653	OM057703	–	–	Guo et al. (2022)
C. qilinense	CAASZK15	Citrullus lanatus	China	MZ475219	OL456696	OL901164	OL449294	OL456655	OM057705	–	–	Guo et al. (2022)
C. dracaenophilum (outgroup)	CBS 118199 T	Dracaena sanderana	China	JX519222	JX546707	JX519230	JX519238	JX519247	JX546756	–	–	Damm et al. (2019)
C. yunnanense (outgroup)	CBS 13213 T	Buxus sp.	China	JX546804	JX546706	JX519231	JX519239	JX519248	JX546755	–	–	Damm et al. (2019)

Ex-type/ex-epitype/ex-neotype/ex-lectotype strains are marked with ^T; BRIP Queensland Plant Pathology Herbarium; CBSCBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands; CFCC China Forestry Culture Collection Center; CGMCC China General Microbiological Culture Collection Center; CMM Culture Collection of Phytopathogenic Fungi Prof. Maria Menezes, Federal Rural University of Pernambuco, Brazil; CNUCC Capital Normal University Culture Collection Center; CPCCulture collection of Pedro Crous, housed at CBS; GUCC the Plant Pathology Department of the College of Agriculture, Guizhou University, China; GZAAS Guizhou Academy of Agricultural Sciences, Guiyang, China; ICMP International Collection of Microorganisms from Plants; IMI International Mycological Institute; LC the LC Culture Collection (a personal culture collection of Lei Cai, housed in the Institute of Microbiology, Chinese Academy of Sciences); LGMF Culture Collection of Laboratory of Genetics of Microorganisms, Federal University of Parana, Curitiba, Brazil; MAFF the Genetic Resources Center, National Agriculture and Food Research Organization, Tsukuba, Ibaraki, Japan; MFLUMae Fah Luang University Herbarium Collection, Chiang Rai, Thailand; MFLUCCMae Fah Luang University Culture Collection, Chiang Rai, Thailand; NBRCNITE Biological Resource Center; NTUCCthe Department of Plant Pathology and Microbiology, National Taiwan University Culture Collection; RGM the Chilean Collection of Microbial Genetic Resources; SAUCCCulture collection of the Department of Plant Pathology, College of Plant Protection, Shenyang Agricultural University, China; VPRI the Victorian Plant Pathology Herbarium; YMF the Herbarium of the Laboratory for Conservation and Utilization of Bio-Resources, Yunnan University, Kunming, Yunnan, China; ZHKUCC the Culture Collection of Zhongkai University of Agriculture and Engineering. The strains generated in this study are in bold.

Phylogenetic analyses

The raw readings were processed and organized into contigs using Geneious Prime 2025.0.3 Java Version 11.0.24+8 (64-bit) software (http://www.geneious.com). The newly generated sequences were used as queries to conduct a BLASTn search against the non-redundant (nr) database in GenBank. The retrieval of similar sequences was conducted, followed by the construction of numerous alignments. The GenBank taxonomy browser was utilized to verify all sequences classified as Colletotrichum in the database. BioEdit version 7.2.5 (Hall 1999) was used to assign open reading frames of the protein-coding sequences of actin, gapdh, β-tubulin, chs-1, and cal according to reference sequences in the GenBank database. The combined sequence data of all loci were used to perform maximum likelihood (ML), maximum parsimony (MP), and Bayesian posterior probability analysis (BI).

The dataset of each gene region was independently aligned with the ‘auto’ strategy (based on data size) in MAFFT (Katoh et al. 2019) and trimmed with the ‘gappyout’ method (based on gaps’ distribution) in TrimAl (Capella-Gutiérrez et al. 2009). BioEdit v. 7.0.9.0 (Hall 1999) was utilised for manual editing, where needed. The best-fit nucleotide substitution models for each dataset were selected based on the Bayesian information criterion (BIC) with rate heterogeneity by ModelFinder (Kalyaanamoorthy et al. 2017). Afterwards, all datasets were concatenated with partition information for the subsequent phylogenetic analyses.

Maximum likelihood (ML), maximum parsimony (MP) and Bayesian posterior probability (BI) analysis were performed using the CIPRES Science Gateway (https://www.phylo.org/portal2) (Miller et al. 2010). The maximum likelihood tree was constructed using RAxMLHPC2 on XSEDE with bootstrapping of 1000 replicates. The ML analysis utilized the GTR + GAMMA model. The maximum parsimony phylogenetic tree was performed using PAUP XSEDE (Swofford 2002). The Bayesian posterior probability (BI) analysis employed a Markov Chain Monte Carlo (MCMC) algorithm with MrBayes on XSEDE, involving four MCMC chains running for 1,000,000 generations and sampling at intervals of 100 generations. The first 25% of constructed trees were eliminated as burn-in, and the remaining trees were used to compute posterior probabilities (Ronquist et al. 2012). The resultant phylograms were visualised with FigTree v. 1.4.4 (Rambaut 2018) and formatted using Adobe Illustrator CC 22.0.0 (Adobe Systems, USA).

The Genealogical Concordance Phylogenetic Species Recognition (GCPSR) model with a pairwise homoplasy index (PHI) test was used to analyze the newly generated taxon and its most phylogenetically close neighbors (Quaedvlieg et al. 2014). The PHI test was performed in SplitsTree v. 4.14.6 (Huson 1998; Huson and Bryant 2006) with a five-locus concatenated dataset (ITS, act, gapdh, β-tubulin, and chs-1) to determine the recombination level among phylogenetically closely related species. A pairwise homoplasy index below a 0.05 threshold (Φw < 0.05) indicated the presence of significant recombination in the dataset. The relationship between closely related species was visualized by constructing a split graph.

Results

Single-locus phylogenetic trees were initially generated for each gene region to assess individual tree topologies and clade support prior to constructing a combined gene tree. In this study, we introduce a novel Colletotrichum species and report 11 previously described species, with six representing new geographical and/or host records. Phylogenetic analyses were conducted using a comprehensive multilocus dataset, incorporating multi-gene regions depending on the species complex. Maximum Likelihood (ML), Maximum Parsimony (MP), and Bayesian posterior probability analysis (BI) methods were applied to infer phylogenetic relationships.

The topologies resulting from the ML, MP, and BI analyses were congruent, with no significant conflicts observed among the trees generated by the three methods. The combined analyses resolved the newly obtained strains within five Colletotrichum species complexes: C. acutatum, C. bambusicola, C. trichellum, C. gloeosporioides, and C. magnum. In each case, the isolates clustered with previously accepted members of the respective complexes, and the resulting trees exhibited strong nodal support, as shown in Figs 1, 3, 5, 12.

Figure 1.

Phylogenetic tree constructed using a maximum likelihood (ML) analysis based on a combined ITS, gapdh, chs-1, act, β-tubulin, and his3 sequences, representing Colletotrichum acutatum species complex. The tree topology of the ML analysis was identical to the Maximum Parsimony (MP) and Bayesian posterior probability (BI) analyses. The final RAxML tree with a likelihood value of -11021.830194 is presented here. The evolutionary model GTR+GAMMA was applied to all the genes. The analysis included sixty-nine (69) taxa with a total of 2183 characters, with 803 distinct alignment patterns, and 7.96% were gaps and undetermined characters. Estimated base frequencies were as follows: A = 0.224158, C = 0.310418, G = 0.240273, T = 0.225151; substitution rates AC = 1.652392, AG = 4.067151, AT = 1.193894, CG = 0.729619, CT = 7.723186, GT = 1.0; gamma distribution shape parameter α = 0.285275; tree length = 0.845774. Bootstrap support values for ML and MP ≥ 70% and Bayesian Posterior Probabilities (BI) ≥ 0.90 are indicated at the nodes as ML/MP/BI. The tree is rooted with C. rusci (CBS 119206) and C. trichellum (CBS 217.64). Type strains are denoted in bold and T and sequences generated in this study are in yellow. Bar = 0.04 represents the estimated number of nucleotide substitutions of site per branch.

Figure 3.

Phylogenetic tree constructed using a maximum likelihood (ML) analysis based on a combined ITS, gapdh, chs-1, act, β-tubulin, and his3 sequences, representing Colletotrichum bambusicola, coccodes, and trichellum species complexes. The tree topology of the ML analysis was identical to the Maximum Parsimony (MP) and Bayesian posterior probability (BI) analyses. The final RAxML tree with a likelihood value of -9417.584651 is presented here. The evolutionary model GTR+GAMMA was applied to all the genes. The analysis included twenty (20) taxa with a total of 2341 characters, with 764 distinct alignment patterns, and 21.79% were gaps and undetermined characters. Estimated base frequencies were as follows: A = 0.217665, C = 0.311394, G = 0.244784, T = 0.226158; substitution rates AC = 1.493066, AG = 4.079410, AT = 1.470090, CG = 1.247573, CT = 7.473294, GT = 1.0; gamma distribution shape parameter α = 0.267361; tree length = 0.838159. Bootstrap support values for ML and MP ≥ 70% and Bayesian Posterior Probabilities (BI) ≥ 0.90 are indicated at the nodes as ML/MP/PP. The tree is rooted with C. acutatum (CBS 112996) and C. fioriniae (CBS 128517). Type strains are denoted in bold and T and sequences generated in this study are in yellow. Bar = 0.04 represents the estimated number of nucleotide substitutions of site per branch.

Figure 5.

Phylogenetic tree constructed using a Bayesian posterior probability (BI) analysis based on a combined ITS, gapdh, act, chs-1, β-tubulin, and cal sequences, representing Colletotrichum gloeosporioides species complex. The tree topology of the ML analysis was identical to the Maximum Parsimony (MP) and Bayesian posterior probability (BI) analyses. The final RAxML tree with a likelihood value of -11975.336 is presented here. The evolutionary model GTR+GAMMA was applied to all the genes. The analysis included one hundred ninety-five (195) taxa with a total of 1689 characters, with 852 distinct alignment patterns, 495 parsimony-informative, 192 singleton sites, and 1002 constant sites. Estimated base frequencies were as follows: A = 0.229358, C = 0.298987, G = 0.241845, T = 0.226158; substitution rates AC = 1.085133, AG = 3.378568, AT = 1.335103, CG = 0.972042, CT = 5.776668, GT = 1.0; gamma distribution shape parameter α = 0.371036; tree length = 1.239. Bootstrap support values for ML and MP ≥ 50% and Bayesian Posterior Probabilities (BI) ≥ 0.90 are indicated at the nodes as ML/MP/BI. The tree is rooted with C. acidae (MFLUCC 17-2659) and C. truncatum (CBS 151.35). Type strains are denoted in bold and T and sequences generated in this study are in yellow. Bar = 0.03 represents the estimated number of nucleotide substitutions of site per branch.

Figure 12.

Colletotrichum jiangxiense (new host record). a–c. Pteris henryi, host habitat; d, e. Culture on OA (d-above, e-reverse); f. Conidiomata on OA; g, h. Conidiophores, conidiogenous cells giving rise to conidia; i, j. Appressoria; k–o. Conidia. Scale bars: 10 µm (i, j); 5 µm (g, h, k–o).

One novel species, Colletotrichum macroconidii (strain GUCC 25-0028 and GUCC 25-0063), was recovered as a distinct lineage within the gloeosporioides species complex. This taxon is supported by bootstrap values and Bayesian posterior probabilities, confirming its phylogenetic distinctiveness. In addition, 11 known species were identified from our collections. Among them, six species represent new records for China or for specific host associations: C. fioriniae from Parthenocissus tricuspidata, C. trichellum from Hedera spp., C. juglandicola from Camellia japonica and Ilex spp., C. nanhuaense from Juglans regia, C. jiangxiense from Juglans regia, and C. magnum from Juglans regia. The remaining five species: C. nymphaeae, C. metake, C. fructicola, C. siamense, and C. gloeosporioides are recognized taxa that were previously reported in similar or nearby regions. Detailed phylogenetic placements and morphological characteristics of all taxa are provided in the respective species descriptions and notes.

To further validate the taxonomic independence of the novel taxon, the Genealogical Concordance Phylogenetic Species Recognition (GCPSR) criterion was applied. The Pairwise Homoplasy Index (PHI) test was employed to detect recombination events among closely related taxa. A Φw value below 0.05 indicates significant recombination, while values above this threshold support genetic separation (Figs 2, 6). For the new species C. macroconidii, the PHI test produced non-significant values (Φw = 1.0), indicating no detectable recombination with the closely related species, C. gardeniae (Fig. 6). These results support the recognition of C. macroconidii as a phylogenetically and evolutionarily distinct species.

Figure 2.

The results of the pairwise homoplasy index (PHI) test for closely related species of Colletotrichum nymphaeae in this study using both LogDet transformation and splits decomposition. PHI test results (Φw) > 0.05 indicate no significant recombination within the dataset.

Figure 6.

The results of the pairwise homoplasy index (PHI) test for closely related species of Colletotrichum macroconidii sp. nov. in this study using both LogDet transformation and splits decomposition. PHI test results (Φw) > 0.05 indicate no significant recombination within the dataset.

. Colletotrichum fioriniae

(Marcelino & Gouli) Pennycook, Mycotaxon 132(1): 150 (2017) [2016]

7ED6EA67-6CC3-5AF3-A21D-300A598AF318

Description.

See Damm et al. (2012a).

Material examined.

China • Guizhou Province, Tongren, Parthenocissus tricuspidata (Siebold & Zucc.) Planch., 2024.07.11, coll. Gou Shiqi, D94/GSQ/GZ38 (dried culture, HGUP 25-0036), living culture GUCC 25-0050.

Notes.

Parthenocissus tricuspidata (Vitaceae, Vitales) is a vigorous liana widely cultivated for ornamental and ecological purposes due to its exceptional adaptability to various environmental conditions, including low temperatures, drought, and nutrient-poor soils (Steinbrecher et al. 2011). Its strong adhesive and climbing abilities make it a preferred species for covering walls, pavilions, and rocks (He et al. 2011). However, with its increasing use in landscape greening, the plant has become more susceptible to a growing number of pathogenic threats. Recent studies have documented several fungal pathogens associated with P. tricuspidata, including Phyllosticta partricusidatae (Zhou et al. 2015), Colletotrichum siamense (Zhao et al. 2020), Septoria tormentillae (Wang et al. 2020), Diaporthe tulliensis (Huang et al. 2021a), Coniella vitis (Yin et al. 2023), Neophysopella vitis (Zhou et al. 2024), C. gloeosporioides, and C. siamensethe (Wang et al. 2025), highlighting the plant vulnerability to fungal diseases.

In the present study, the fungal strain GUCC 25-0050 was isolated from symptomatic leaves of P. tricuspidata. Based on multilocus phylogenetic analyses of ITS, gapdh, chs-1, act, β-tubulin, and his3 gene regions (Fig. 1), this strain was identified as Colletotrichum fioriniae. This identification is supported by its phylogenetic placement. This is the first report of C. fioriniae on P. tricuspidata in China. The identification of C. fioriniae as a pathogen of P. tricuspidata underscores the need for further investigation into its epidemiology, pathogenicity, and potential impact on ornamental and ecological plantings.

. Colletotrichum nymphaeae

(Pass.) Aa, Netherlands Journal of Plant Pathology, Supplement 1 84(3): 110 (1978)

AFAAD7F4-9DE0-5748-87FF-65CD75D06835

• = Colletotrichum speciosum Z.F. Yu, in Yu et al, J. Fungi 8(2, no. 185): 24 (2022).

• = Colletotrichum simulanticitri Z.F. Yu, in Yu et al, J. Fungi 8(2, no. 185): 24 (2022).

Description.

See more details in Damm et al. (2012a).

Material examined.

China • Guizhou Province, Tongren, on unidentified branch, 2024.07.11, coll. Gou Shiqi, D79/FDS28/GZ35 (dried culture, HGUP 25-0033), living culture GUCC 25-0047; ibid., on rotten branch, 2024.07.11, coll. Gou Shiqi, E43/FDS8/GZ45 (dried culture, HGUP 25-0040), living culture GUCC 25-0056.

Notes.

This species was originally described from Nymphaea alba leaves in Kortenhoef by Van der Aa (1978). According to Damm et al. (2012a), C. nymphaeae is reliably distinguished from related taxa by the beta tubulin (β-tubulin) gene, although other loci show high intraspecific variability. Based on morphological characteristics and multilocus phylogenetic analyses, strains GUCC 25-0047 and GUCC 25-0056 were identified as Colletotrichum nymphaeae. Morphologically, the new strain in this study produces an asexual morph on PDA with hyaline to pale brown, septate, branched, and smooth-walled conidiophores. The conidiogenous cells were cylindrical, measuring 9.7–15 × 2.8–4.3 µm, sharing similar coloration and wall texture, and were characterized by a distinct collarette and evident periclinal thickening. Conidia were hyaline, aseptate, smooth-walled, and straight, varying from cylindrical to cylindric-clavate, measuring 9.9–15.5 × 3.5–5 µm, consistent with the characteristics of the genus Colletotrichum. Phylogenetically, both strains clustered with C. nymphaeae, showing high sequence similarity and revealed low genetic differences for each gene with the type strain (CBS 515.78): GUCC 25-0047 (ITS = 99% [2/541], gapdh = 98% [5/248], chs-1 = 99% [1/282], act = 99% [1/246], β-tubulin = 99% [1/490], his3 = 99% [2/382]) and GUCC 25-0056 (ITS =99% [3/541], gapdh = 99% [3/248], chs-1 = 99% [1/282], act = 100% [0/246], β-tubulin = 100% [0/490], his3 = 99% [2/382]). These results indicate only minor genetic variation from C. nymphaeae. The species formed a well-supported clade, clearly distinct from other taxa within the C. acutatum species complex (Fig. 1). This placement was further supported by the PHI test, which indicated significant recombination among the two collections (Fig. 2).

Additionally, Colletotrichum speciosum (YMF 1.07301) and Colletotrichum simulanticitri (YMF 1.07302) clustered with the type strain of C. nymphaeae (CBS 515.78), along with five other strains, including GUCC 25-0047 and GUCC 25-0056 (Fig. 1). Phylogenetic analyses based on single loci showed that chs-1, β-tubulin, and his3 provided the highest resolution for this species complex, producing a topology most consistent with the combined multilocus tree and supported by significant bootstrap values. Colletotrichum speciosum was described by Yu et al. (2022) based only on ITS sequence data, which shows 99% similarity (538/541 bp) with C. nymphaeae, suggesting a close relationship. However, since only ITS is not sufficient for species delimitation in Colletotrichum, and only a single strain is known, we tentatively consider C. speciosum a synonym of C. nymphaeae. Further studies using multilocus genes and more strains are needed to confirm this synonymy. In comparison, C. simulanticitri was described using four loci (ITS, gapdh, chs-1, and act), providing stronger molecular support (Yu et al. 2022). Both the combined analysis and single-locus trees placed C. simulanticitri together with C. nymphaeae strains. Pairwise comparisons between C. simulanticitri (YMF 1.07302) and the type strain of C. nymphaeae (CBS 515.78) revealed low genetic differences: ITS = 0.3% (2/542 bp), gapdh = 0.8% (2/238 bp), chs-1 = 0% (0/226 bp), and act = 0% (0/246 bp). Morphologically, C. simulanticitri shares similarities with C. nymphaeae, including hyaline, cylindrical to oblong conidia (10–13.5 × 4–5 µm for C. simulanticitri), as well as dark brown, septate, clavate, or oval appressoria (Damm et al. 2012a; Yu et al. 2022). Notably, the morphological characters of C. simulanticitri were observed on CMA, whereas C. nymphaeae was observed on PDA in this study. Taken together, the close phylogenetic position, minimal genetic divergence, and morphological similarity strongly indicate that C. simulanticitri and C. nymphaeae represent the same species. This is further confirmed by the PHI test, which indicated significant recombination between the taxa (Fig. 2). It is also important to note that both C. speciosum and C. simulanticitri are currently listed as nomen invalidum (nom. inval.) in MycoBank. The name C. simulanticitri has already been synonymized with C. nymphaeae, and C. speciosum is invalid due to a violation of Article 40.8 of the International Code of Nomenclature. Based on morphological features, multilocus phylogeny, recombination analysis, and taxonomic status, we recognize C. simulanticitri as a synonym of C. nymphaeae, and provisionally treat C. speciosum as its synonym as well.

. Colletotrichum metake

Sacc., Annls mycol. 6(6): 557 (1908)

34378B8C-48A3-5C43-9203-00EA7751A640

Description.

See Sato et al. (2012).

Material examined.

CHINA • Guizhou Province, Guiyang, symptomatic leaves of Poaceae spp. (bamboo), 2024.01.13, coll. Wang Xingchang, BB-1-19-1-4-A/GZ26 (dried culture, HGUP 25-0026), living culture GUCC 25-0038.

Notes.

In China, research on Colletotrichum species infecting bamboo remains scarce. For instance, Ren et al. (2008) identified C. coccodes as the causal pathogen of anthracnose on Bambusa pervariabilis. More recently, Wang et al. (2021a) reported C. metake as a seed endophyte from Chimonobambusa quadrangularis. Despite the high diversity of bamboo hosts, Colletotrichum species associated with bamboo are relatively understudied. To date, only five Colletotrichum species have been reported from members of the Bambusoideae subfamily. Among these, C. coccodes (Ren et al. 2008), C. graminicola (Parris 1959), and C. trichellum (Crouch et al. 2009) were identified based on morphological characteristics. In contrast, C. hsienjenchang and C. metake were described using a combination of morphological and phylogenetic analyses (Sato et al. 2012; Wang et al. 2021a). Additionally, Zhou (2018) investigated the endophytic fungal communities in bamboo seeds and found Colletotrichum to be one of the dominant genera, although the isolates were not identified to species level.

In the present study, strain GUCC 25-0038, isolated from bamboo in Guizhou Province, was identified as Colletotrichum metake based on multilocus phylogenetic analyses (Fig. 3). This represents a novel strain of C. metake, which was previously reported from bamboo in China by Wang et al. (2021a).

. Colletotrichum trichellum

(Fr.) Pat., Cat. Rais. Pl. Cellul. Tunisie (Paris): 127 (1897)

AFD619EA-5280-5BFD-BD81-022E6AE3EC64

Fig. 4

Figure 4.

Colletotrichum trichellum (GUCC 25-0039, new record). a, b. Culture on OA (a-above, b-reverse); c. Conidiomata; d, e. Conidiogenous cells giving rise to conidia; f, g. Setae; h–l. Conidia. Scale bars: 10 µm (d, e, i–l); 25 µm (f–h).

Description.

See Sutton (1962).

Material examined.

CHINA • Guizhou Province, Guiyang, Guizhou University, symptomatic leaves of Hedera L. (Ivy), 2024.01.20, coll. Wang Xingchang, 1.28cct1-2/GZ27 (dried culture, HGUP 25-0027), living culture GUCC 25-0039.

Notes.

Colletotrichum trichellum is known to cause leaf and stem spots on English ivy (Hedera helix) and has been reported as a pathogen in several countries, including Canada, Germany, Guatemala, the Netherlands, New Zealand, and the United Kingdom (Damm et al. 2009; Jayawardena et al. 2021). This species is particularly notable for its ability to infect Hedera species, producing distinct symptoms such as necrotic lesions and chlorosis on leaves and stems. In the present study, strain GUCC 25-0039, isolated from Hedera spp. (ivy), was identified as Colletotrichum trichellum based on both morphological characteristics and multilocus phylogenetic analyses. Morphologically, the strain in this study produced an asexual morph on oatmeal agar (OA) with acervular conidiomata that appeared dark and were surrounded by mycelium, bearing white to cream conidial masses; conidiophores were hyaline, smooth-walled, septate, and branched, while setae were medium to dark brown, smooth to finely verruculose, with a knobbed and rounded tip, 1–3-septate, and 25–50 µm long. Conidiogenous cells were subcylindrical, straight to curved, and measured 16.4–26.4 × 2.6–4.4 µm. Conidia were hyaline, aseptate, falcate, slightly curved, fusiform, and abruptly tapered at both ends, measuring 20.4–25.7 × 3.8–5.5 µm. The overall morphology is consistent with that of C. trichellum as previously reported (Sutton 1962; Damm et al. 2009). Phylogenetic analyses of combined gene regions placed strain GUCC 25-0039 within the well-supported C. trichellum clade (Fig. 3). The multilocus phylogenetic tree showed a clear monophyletic grouping of GUCC 25-0039 with reference strains of C. trichellum, supported by 100% ML, 100% MP, 1.00 BI, confirming its taxonomic identity within the Colletotrichum trichellum species complex. Morphologically, the strain exhibited characteristics consistent with those described for C. trichellum, including simple to branched conidiophores and hyaline, fusiform, aseptate conidia with slight curvature, which are features considered diagnostic for this species (Sutton 1962; Damm et al. 2009). Additionally, the presence of elongated, acicular conidia further supported its placement within the genus Colletotrichum. Given the pathogenic potential of C. trichellum on ornamental Hedera species and its widespread distribution, its detection poses a potential concern for the horticultural industry. This study represents the first geographical record of C. trichellum associated with Hedera spp. in China.

. Colletotrichum macroconidii

Norph. & M.T. Zou sp. nov.

99741D26-19A0-5F80-9596-59580E10F947

903940

Figs 7, 8

Figure 7.

Colletotrichum macroconidii (GUCC 25-0028, ex-type). a, b. Culture on PDA 10-days (a-above, b-reverse); c, d. Conidiomata on PDA; e. Conidioma immersed in PDA. Scale bars: 10 µm (e).

Figure 8.

Colletotrichum macroconidii (GUCC 25-0028, ex-type). a, b. Culture on WA with sterile toothpick 10-days (a-above, b-reverse); c. Conidiomata on WA; d. Setose conidioma; e. Conidiophores, conidiogenous cells, giving rise to conidia; f, g. Setae; h–l. Conidia. Scale bars: 50 µm (d); 10 µm (e–l).

Etymology.

The epithet ‘macroconidii’ refers to the distinctly large conidia produced by this species, which serve as a key morphological feature distinguishing it from closely related taxa in the Colletotrichum gloeosporioides species complex. Derived from Greek makros (large) and Latin conidium (spore), meaning having large conidia.

Holotype.

HGUP 25-0018.

Ex-type.

GUCC 25-0028.

Description.

Associated with symptomatic leaves of Juglans regia L. Sexual morph: Not observed. Asexual morph: Conidiomata pycnidial, globose, dark brown, superficial on WA, releasing conidia in a yellow mass, slimy from which setae and conidiophores were produced. Setae dark brown, concoloured, smooth-walled to finely verruculose, 1–3-septate, (80–)100–150(–180) µm long, base cylindrical to conical, 4–6(–9) µm diam., tip acute to roundish. Chlamydospores in cultures observed, in branched chains, brown, verrucose, 11–25 × 5–8 µm. Conidiophores pale brown, septate, strongly branched, smooth-walled or verrusculose, up to 90 µm long. Conidiogenous cells enteroblastic, pale brown, smooth-walled or verruculose, cylindrical to elongate ampulliform, 10–25 × 3–5 µm, opening 1.5–2 µm diam, collarette distinct, 1–2 µm long, periclinal thickening visible. Conidia (14.4–)15–17(–18.2) × (4.5–)5–6(–6.8) μm (mean ± SD = 16 ± 0.4 × 5 ± 0.9 μm), n = 80, L/W ratio = 2.8, hyaline, aseptate, straight, smooth-walled, cylindrical, the apex and base rounded, guttules contents with 1–2 large drop.

Culture characteristics.

Colonies on PDA reaching 7–8 cm diam after 7 d at room temperature (±25 °C), under light 12 hr/dark 12 hr, colonies rhizoid to filamentous, dense, flat or raised surface, with filiform margin, white from above and white to pale-yellow reverse, with producing grouped-pycnidia. Colonies on WA with/without sterilized toothpick, reaching 5 cm in diameter after 7 d at room temperature (±25 °C), under light 12 hr/dark 12 hr, colonies rhizoid to filamentous, dense, with flat surfaces and filiform margins, appearing white to pale gray-green from both the surface and reverse. Pycnidia developed both on the agar surface and as immersed structures within the medium.

Material examined.

China • Yunnan Province, Dali, Symptomatic leaves of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ5-7/GZ12_1, (dried culture, HGUP 25-0018, holotype), ex-type living culture GUCC 25-0028; ibid., GZ12_2, living culture GUCC 25-0063.

Notes.

Within the Colletotrichum gloeosporioides species complex (Fig. 5), two strains (GUCC 25-0028 and GUCC 25-0063) formed a distinct sister clade to C. gardeniae with 74% ML, 0.69 BI support (Fig. 5). Morphologically, C. macroconidii exhibits typical features of the complex, including hyaline, smooth-walled, aseptate, straight, cylindrical conidia with rounded apices and bases. However, it can be distinguished from its closest relatives by subtle yet consistent differences in conidial morphology. In particular, this species produces narrower conidia compared to C. gardeniae, associated with leaf spots on Gardenia jasminoides, which has conidia measuring 16.2 ± 0.8 × 6.1 ± 0.4, with a length/width (L/W) ratio of 2.7. In addition, C. macroconidii produces setae, which were not observed in C. gardeniae (Zhang et al. 2023b). It is important to note that the micromorphological characteristics of C. gardeniae were obtained from cultures grown on PDA, whereas those of C. macroconidii were observed on WA, as the latter species did not sporulate on PDA. On PDA, C. gardeniae formed flat colonies with an entire margin, white to vinaceous buff in color, turning orange in the center due to sporulation, and partially covered by short white aerial mycelium, with a growth rate of 72 mm in 7 days (Zhang et al. 2023b). In contrast, C. macroconidii produced rhizoid to filamentous, dense colonies with flat to raised surfaces and filiform margins. Colonies were white from above and white to pale-yellow in reverse, forming grouped pycnidia; however, no sporulation was observed within 7–10 days (Fig. 7). In addition to morphological differentiation, phylogenetic analysis based on multilocus sequence data supports the distinctiveness of C. macroconidii. Sequence similarity between these two species revealed differences of 3/547 base pairs in the ITS region, 4/231 base pairs in the gapdh region, 2/249 base pairs in the chs-1 region, no differences in the act region, and 5/689 base pairs in the β-tubulin region. The C. gardeniae was missing his3, Apmat, and cal gene sequences. The pairwise homoplasy index (PHI) test showed no significant evidence of recombination between C. macroconidii and C. gardeniae (Φw = 1.0000; Fig. 6), further confirming its genetic separation. Taken together, the combination of morphological and the absence of recombination supports the recognition of Colletotrichum macroconidii as a novel species within the C. gloeosporioides species complex.

. Colletotrichum fructicola

Prihast., L. Cai & K.D. Hyde, Fungal Diversity 39: 96 (2009)

B6C5FE05-0C61-5C79-868B-0B8F21CF0F9B

Description.

See Wang et al. (2017).

Material examined.

China • Yunnan Province, Dali, on symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ11-9/GZ05 (dried culture, HGUP 25-0011), living culture GUCC 25-0021; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ1-6/GZ06 (dried culture, HGUP 25-0012), living culture GUCC 25-0022; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ18-4/GZ07 (dried culture, HGUP 25-0013), living culture GUCC 25-0023; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ21-6/GZ08 (dried culture, HGUP 25-0014), living culture GUCC 25-0024; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ5-5/GZ09 (dried culture, HGUP 25-0015), living culture GUCC 25-0025; CHINA, Guizhou Province, Tongren, on symptomatic leaves of Actinidia chinensis var. deliciosa (Kiwi Fruit Leaf), 2024.07.11, coll. Gou Shiqi, D20/FDS1/GZ33 (dried culture, HGUP 25-0032), living culture GUCC 25-0045; • ibid. on symptomatic leaves of Rosa chinensis Jacq., 2024.07.11, coll. Gou Shiqi, D97/FDS23ROSE/GZ39 (dried culture, HGUP 25-0037), living culture GUCC 25-0051.

Notes.

Colletotrichum fructicola has been increasingly recognized as a significant plant pathogen with a broad host range in China. In 2017 (Wang et al. 2017), it was identified as the pathogen of walnut anthracnose on Juglans regia in Shandong Province, where infected fruits exhibited subcircular to irregular sunken lesions with pink conidial masses. The identification was confirmed through morphological characteristics and molecular analyses (Wang et al. 2017). Colletotrichum fructicola was also reported as the causal agent of leaf spot disease on Actinidia chinensis, based on morphological characteristics and multilocus phylogenetic analyses (Huang et al. 2022). Recently, C. fructicola was associated with anthracnose on Rosa chinensis in Henan Province, presenting as irregular brown specks that expanded into large necrotic lesions. Pathogenicity assays and molecular data supported its identification (Du et al. 2023). Peng et al. (2023) reported that among Colletotrichum species in China, C. fructicola is the dominant taxon causing Camellia anthracnose, making it the most prevalent species on Camellia spp. These findings emphasize the expanding distribution and host range of C. fructicola in China.

In the present study, strains GUCC 25-0021, GUCC 25-0022, GUCC 25-0023, GUCC 25-0024, and GUCC 25-0025, isolated from Camellia japonica, as well as GUCC 25-0045 and GUCC 25-0051, obtained from Actinidia chinensis and Rosa chinensis, respectively, were identified as C. fructicola based on morphological features and multilocus phylogenetic analyses. The morphological characteristics observed in this study revealed the production of an asexual morph on PDA, with hyphae that were hyaline to light brown, branched, and septate. Conidiophores, which were hyaline and occasionally branched, formed singly or in clusters on the hyphae and measured 15–25.5 × 3.5–6.5 µm. Conidia were single-celled, hyaline, elliptical to oblong, measured 9–14.5 × 4.5–5 µm, with smooth walls and no septa. These morphological features are consistent with those of C. fructicola (Wang et al. 2017). Phylogenetic analyses of the combined genes placed the seven new strains within the C. fructicola clade, supported by 100% ML, 95% MP, and 1.00 BI, confirming their taxonomic identity within the Colletotrichum gloeosporioides species complex (Fig. 5). These findings are consistent with previous records and suggest that this species may be widespread and potentially associated with diverse plant hosts across China.

. Colletotrichum siamense

Prihast., L. Cai & K.D. Hyde, Fungal Diversity 39: 98 (2009)

93CCC68F-4ECD-5FBD-9F4D-D037906800C3

Description.

See Prihastuti et al. (2009), Zhang et al. (2023b), Khuna et al. (2024).

Material examined.

China • Yunnan Province, Dali, on Juglans regia L., 2023.12.26, Zou Mengting, CXD2-1/GZ18 (dried culture, HGUP 25-0022), living culture GUCC 25-0032; China, • Guizhou Province, Zunyi, Xishui, National Nature Reserve In Xi’an, Pteridophyta, 2024.05.20, coll. Gou Shiqi/Wang Jiaping, Xsj1.1/GZ21 (dried culture, HGUP 25-0023), living culture GUCC 25-0033; • ibid. on Pteridophyta, 2024.05.20, coll. Gou Shiqi/Wang Jiaping, xsj1.2/GZ22 (dried culture, HGUP 25-0024), living culture GUCC 25-0034; • ibid. on Pteridophyta, 2024.05.20, coll. Gou Shiqi/Wang Jiaping, Xsj1/GZ23 (dried culture, HGUP 25-0025), living culture GUCC 25-0035; China, • Guizhou Province, Tongren, on Parthenocissus quinquefolia (L.) Planch., 2024.07.11, coll. Gou Shiqi, D86/FDS40/GZ36 (dried culture, HGUP 25-0034), living culture GUCC 25-0048; • ibid. on Parthenocissus tricuspidata (Siebold & Zucc.) Planch., 2024.07.11, coll. Gou Shiqi, D93/FDS42/GZ37 (dried culture, HGUP 25-0035), living culture GUCC 25-0049.

Notes.

Colletotrichum siamense, a member of the Colletotrichum gloeosporioides species complex, was described by Weir et al. (2012) and is now recognized as a widespread phytopathogen with a broad host range across multiple countries (Talhinhas and Baroncelli 2021, 2023). It has been reported to cause anthracnose and leaf spot diseases on a variety of economically important crops and ornamental plants (Liu et al. 2017; de Silva et al. 2019; Zhang et al. 2023c). In China, C. siamense has been documented on several ornamental and fruit crops, including Alocasia macrorrhiza (Huang et al. 2021b), Viburnum odoratissimum (Li et al. 2023), Syzygium samarangense (Yao et al. 2023), and Artocarpus heterophyllus (Khuna et al. 2024). Identification of this species typically relies on morphological characteristics and multilocus phylogenetic analyses, while its pathogenicity is confirmed through inoculation assays following Koch’s postulates. The widespread distribution and extensive host range of C. siamense highlight its significance as a potential threat to plant health and agricultural sustainability.

In the present study, strain GUCC 25-0032 isolated from Camellia japonica; strains GUCC 25-0033, GUCC 25-0034, and GUCC 25-0035 from unidentified pteridophytes; and strains GUCC 25-0048 and GUCC 25-0049 from Parthenocissus quinquefolia and P. tricuspidata, respectively, were identified as C. siamense based on morphological characteristics, and phylogenetic analyses (Fig. 5). Morphologically, the strains in this study produced an asexual morph on PDA, with hyaline to pale brown, septate, and branched conidiophores. Conidiogenous cells were hyaline to pale brown, cylindrical to ampulliform in shape, and measured 13.4–22.5 × 2.6–4.7 µm. Conidia were single-celled, hyaline, smooth-walled, cylindrical with rounded ends, guttulate, and measured 12.8–19.1 × 4.9–7 µm. Phylogenetic analyses of the combined gene sequences placed the six new strains within the C. siamense clade. The species formed a monophyletic clade, supported by a 0.92 BI, with a distinct clade separating it from other species, thereby confirming its taxonomic identity within the Colletotrichum gloeosporioides species complex (Fig. 5). This species continues to be reported from diverse host plants in China (He et al. 2011; Zhao et al. 2020; Peng et al. 2022, 2023), and the current study contributes six novel strains to the expanding records of C. siamense.

. Colletotrichum gloeosporioides

(Penz.) Penz. & Sacc., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 2(5): 670 (1884)

9225DE64-9122-58B3-9B7A-4A70CAF5923F

Description.

See Peng et al. (2023).

Material examined.

China • Guizhou Province, Guiyang, Guiyang Botanical Garden Of Medical Plants, symptomatic leaves of Camellia japonica L., 2024.4.4, coll. Wang Xingchang, Sch1-1-1/GZ29 (dried culture, HGUP 25-0028), living culture GUCC 25-0041; • ibid. on symptomatic leaves of Camellia japonica L., 2024.4.4, coll. Wang Xingchang, Sch1-1-4/GZ30 (dried culture, HGUP 25-0029), living culture GUCC 25-0042.

Notes.

Colletotrichum gloeosporioides is a well-known plant pathogenic species within Colletotrichum, responsible for anthracnose diseases across a wide range of host plants (Zhang et al. 2023c). Typical symptoms include dark, sunken lesions on leaves, stems, and fruits, which can significantly impact plant health and yield. This species is of considerable economic importance due to its broad host range, affecting major fruit crops such as papaya, mango, and citrus, as well as ornamental plants like Camellia japonica (Peng et al. 2023). Accurate identification of C. gloeosporioides relies on a combination of morphological traits, such as the shape and size of conidia and conidiophores, as well as molecular tools, particularly multilocus phylogenetic analyses. Peng et al. (2023) reported the occurrence of C. gloeosporioides on C. japonica, extending the known host range of the species and underscoring its potential impact on ornamental horticulture.

In the present study, strains GUCC 25-0041 and GUCC 25-0042, isolated from symptomatic tissues of Camellia japonica, were identified as Colletotrichum gloeosporioides based on both morphological characteristics and multilocus phylogenetic analyses (Fig. 5), consistent with previous findings (Peng et al. 2023). Morphologically, the strains in this study produced an asexual morph on PDA, characterized by hyaline, septate, and branched conidiophores. Conidiogenous cells were cylindrical to ampulliform, measured 12.9–30 × 2.3–3.9 µm, and hyaline. Conidia were single-celled, hyaline, smooth-walled, elliptical to oval with rounded ends, measuring 12.3–16 × 4.9–6.0 µm. Phylogenetic analyses of the combined genes placed the two new strains within the C. gloeosporioides species clade. The species clade formed a distinct clade, separated from other species, with support of 75% ML, confirming its taxonomic identity within the Colletotrichum gloeosporioides species complex (Fig. 5).

. Colletotrichum juglandicola

Y. Zhang ter & Lin Zhang bis, MycoKeys 99: 139 (2023)

12AC2A2E-DBE2-5A58-B06D-DBCFACC556D2

Fig. 9

Figure 9.

Colletotrichum juglandicola (GUCC 25-0043, new host record). a–c. Camellia japonica, host habitat; d, e. Culture on OA (d-above, e-reverse; f. Conidiomata on OA; g, h. Conidiophore, conidiogenous cells giving rise to conidia; i, j. Setae; k–o. Conidia. Scale bars: 10 µm (g–k); 5 µm (l–o).

Description.

See Zhang et al. (2023a).

Material examined.

China • Guizhou Province, Guiyang, Guiyang Botanical Garden Of Medical Plants, Camellia japonica L., 2024.4.4, coll. Wang Xingchang, Sch6/GZ31 (dried culture, HGUP 25-0030), living culture GUCC 25-0043; China, • Shandong Province, Taian, Tianwaicun Street, Mountain Tai, Ilex L., 2024.02, coll. Wang Xingchang, 2.26-DQ-2/GZ32 (dried culture, HGUP 25-0031), living culture GUCC 25-0044.

Notes.

Colletotrichum juglandicola was originally described by Zhang et al. (2023a) as the causal agent of anthracnose-like symptoms on Juglans regia, characterized by dark, sunken lesions on leaves, stems, and fruit. Zhang et al. (2023a) emphasized the economic impact of C. juglandicola in walnut orchards, citing its potential to significantly reduce both yield and nut quality. Like other members of the genus, C. juglandicola can infect multiple plant tissues and exhibits a broad host range, making it an important pathogen for integrated disease management. The identification was based on a combination of morphological characteristics, such as the features of conidia and conidiophores and multilocus phylogenetic analyses. In this study, strains GUCC 25-0043 (from Camellia japonica) and GUCC 25-0044 (from Ilex sp.) were identified as C. juglandicola based on both morphological characteristics and multilocus phylogenetic analyses (Figs 5, 9). Morphologically, the strains in this study produced an asexual morph on OA with acervular conidiomata that generated yellow to orange conidial masses; conidiophores were hyaline, smooth-walled, septate, and branched, while setae were medium to dark brown, smooth to finely verruculose near the tip, rounded, 1–3-septate, and 60–110 μm long; conidiogenous cells measured 11–29.5 × 2.7–5 μm, subcylindrical, and either straight or curved. Conidia were hyaline, smooth-walled, subcylindrical, with both ends rounded, measuring 15.8–20.0 × 4.8–6.6 μm, containing 1–3 guttules with granular contents. Phylogenetically, the two new strains grouped with C. juglandicola (strain GCMCC 3.2431 and GCMCC 3.24313) with support values of 91% ML and 0.93 PP (Fig. 5). This represents the first report of C. juglandicola infecting C. japonica and Ilex sp. in China, thereby expanding the known host range of the species and highlighting its potential as a pathogen across diverse plant families.

. Colletotrichum nanhuaense

Z.F. Yu, J. Fungi 8(2, no. 185): 17 (2022)

6FB9C313-240F-56A6-BF0E-B6CA37ABCD62

Fig. 10

Figure 10.

Colletotrichum nanhuaense (new host record). a, b. Culture on PDA (a-above, b-reverse); c Conidiomata on PDA; d–f. Conidiophore, conidiogenous cells giving rise to conidia; g–k. Conidia. Scale bars: 10 µm (d–k).

Description.

See more details in Yu et al. (2022).

Material examined.

China • Yunnan Province, Dali, symtomatic fruit of Juglans regia L., 2023.12.26, coll. Zou Mengting, CXD7-9/GZ10 (dried culture, HGUP 25-0016), living culture GUCC 25-0026.

Notes.

Colletotrichum nanhuaense was originally described by Yu et al. (2022) as the causal agent of leaf spot disease on Ageratina adenophora in Yunnan Province, China. In the present study, strain GUCC 25-0026, isolated from Juglans regia, was identified as C. nanhuaense based on both morphological characteristics (Fig. 10) and multilocus phylogenetic analyses (Fig. 5). Morphologically, the strain produced an asexual morph on PDA, with hyaline to dark brown, branched, and septate hyphae. Conidiophores measured 9–29.5 × 3–6.5 µm, were solitary or clustered, hyaline, and unbranched, and were formed on the hyphae. Conidia were single-celled, elliptical to ovoid-shaped, measuring 10.5–15 × 5.5–6 µm, hyaline, with smooth walls and no septa. Phylogenetic analyses placed the new strain within the C. nanhuaense clade with strong support (100% ML/1.0 PP; Fig. 5). The detection of C. nanhuaense on walnut expands its host range and raises concerns about its impact on other crops. Further research on its biology and spread, especially in Juglans, is needed, along with ongoing monitoring. Pathogenicity tests are required to confirm its role in J. regia. This is the first report of C. nanhuaense on J. regia and in China.

. Colletotrichum jiangxiense

F. Liu & L. Cai, Persoonia 35: 82 (2015)

97BA1826-F2A2-5A53-8466-C13BA2CE40E1

Fig. 12

Description.

See Liu et al. (2015), Chang et al. (2024), Zhang et al. (2023b).

Material examined.

China, • Yunnan Province, Dali, symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ4-6/GZ11 (dried culture, HGUP 25-0017), living culture GUCC 25-0027; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ11-10/GZ16 (dried culture, HGUP 25-0021), living culture GUCC 25-0031; China, • Guizhou Province, Tongren, on symptomatic leaves of Pteris henryi Christ., 2024.07.11, coll. Gou Shiqi, E14/FDS34/GZ40 (dried culture, HGUP 25-0038), living culture GUCC 25-0052; • ibid. on symptomatic leaves of Parthenocissus tricuspidata (Siebold & Zucc.) Planch., 2024.07.11, coll. Gou Shiqi, E17/FDS41/GZ41 (dried culture, HGUP 25-0039), living culture GUCC 25-0053.

Notes.

Colletotrichum jiangxiense was initially described as an endophytic fungus associated with Camellia sinensis in China (Liu et al. 2015). Since its discovery, the species has been increasingly recognized as a plant pathogen, causing anthracnose and other disease symptoms on various fruit and ornamental hosts. Recent studies have reported its involvement in anthracnose outbreaks in crops such as avocado (Ma et al. 2018; Ayvar Serna et al. 2021; Guo et al. 2022; Zhang et al. 2023b). Additionally, C. jiangxiense has been identified as a pathogen in Mexico, where it causes anthracnose on avocado (Ayvar Serna et al. 2021), although it has not yet been reported as a plant pathogen in North America. The international trade of live plants between East Asia and North America raises concerns regarding the potential introduction of C. jiangxiense into new environments. Under favorable conditions, this species may shift from an endophytic or low-pathogenic state to a more aggressive pathogenic form, posing a threat to both ornamental and economically important crops. This highlights the importance of phytosanitary surveillance and monitoring for emerging pathogen-host interactions (Liebhold et al. 2012).

Colletotrichum jiangxiense is identified based on both its morphological characteristics and phylogenetic placement, supported by multilocus gene sequence data (Liu et al. 2015). In this study, we report the first identification of C. jiangxiense on Juglans regia, Pteris henryi, and Parthenocissus tricuspidata, thus expanding its known host range. Strains GUCC 25-0027 and GUCC 25-0031, isolated from J. regia, and strains GUCC 25-0052 and GUCC 25-0053, isolated from P. henryi and P. tricuspidata, respectively, were identified as C. jiangxiense through morphological and multilocus phylogenetic analyses (Figs 5, 11). Phylogenetically, four strains clustered with C. jiangxiense in both combined gene tree and single loci tree, showing high sequence similarity and only minor genetic differences from the type strain (CBS 515.78): GUCC 25-0027 (ITS = 100% [0/514], gapdh = 100% [0/252], chs-1 = 100% [0/287], act = 100% [0/273], β-tubulin = 100% [0/689], cal = 100% [0/733], ApMat = 99% [2/949]); GUCC 25-0031 (ITS = 100% [0/514], gapdh = 100% [0/252], chs-1 = 100% [0/287], act = 100% [0/273], β-tubulin = 100% [0/700], cal = 100% [0/757], ApMat = 100% [0/950]); GUCC 25-0052 (ITS = 100% [0/514], gapdh = 100% [0/252], chs-1 = 100% [0/287], act = 100% [0/273], β-tubulin = 100% [0/732], cal = n/a, ApMat = n/a); and GUCC 25-0053 (ITS = 100% [0/514], gapdh = 100% [0/252], chs-1 = 100% [0/287], act = 100% [0/273], β-tubulin = 100% [0/732], cal = 100% [0/751], ApMat = 99% [1/944]). These results indicate only minor genetic variation from C. jiangxiense. This placement was further supported by the PHI test, which indicated significant recombination among the two collections (Fig. 11). Morphologically, the strains exhibit hyaline, smooth-walled, aseptate, and unbranched conidiophores that grow directly from the hyphae. Conidiogenous cells were hyaline, cylindrical to clavate, measuring 12.6–27.5 × 3.2–4.6 µm, smooth-walled, and lacked a collarette. Conidia were cylindrical, with both ends bluntly rounded, or one end bluntly rounded and the other acutely rounded, measuring 14.6–18 × 4.9–6.5 µm, aseptate, and smooth-walled. Appressoria were brown to pale-brown, circular, subglobose to ellipsoidal or irregular, with lobed margins, measuring 5–9 × 5–8 µm. These morphological characteristics are consistent with those of C. jiangxiense as described by Liu et al. (2015).

Figure 11.

The results of the pairwise homoplasy index (PHI) test for closely related species of Colletotrichum macroconidii sp. nov. in this study using both LogDet transformation and splits decomposition. PHI test results (Φw) > 0.05 indicate no significant recombination within the dataset.

Phylogenetic analysis placed the four new strains within a distinct clade, which also includes C. nullisetosum, C. oblongisporum, C. gracile, and C. tengchongense. This clade is strongly supported by 98% ML and 0.99 PP values (Fig. 5). However, C. nullisetosum, C. oblongisporum, and C. gracile are currently considered as nomenclaturally invalid (nom. inval.) in MycoBank, due to violations of Article 40.8 of the International Code of Nomenclature. Based on their morphological characteristics (Yu et al. 2022) and taxonomic status, we treat these three species as synonyms of C. jiangxiense. Colletotrichum tengchongense is retained as a valid species name within the clade. Morphologically, it differs from C. jiangxiense by the structure of its conidiophores and conidiogenous cells, with C. tengchongense producing lageniform or ampulliform conidiogenous cells (Liu et al. 2015; Zheng et al. 2022). Phylogenetically, C. tengchongense forms a relatively long branch within the C. jiangxiense species clade. However, its sequence similarity with the type strain of C. jiangxiense indicates only minor genetic differences: ITS = 0% [0/515], gapdh = 0% [0/274], chs-1 = 0.3% [1/287], act = 0.4% [1/246], β-tubulin = 0% [0/700], cal = 0.1% [1/715], ApMat = 0.2% [2/870]. These results suggest very limited genetic variation from C. jiangxiense, and we therefore treat C. tengchongense as conspecific with C. jiangxiense. This conclusion was further supported by the PHI test, which indicated significant recombination within the C. jiangxiense species group (Fig. 11). Nevertheless, further studies incorporating additional loci are needed to confirm the phylogenetic placement of this taxon.

. Colletotrichum magnum

(S.F. Jenkins & Winstead) Rossman & W.C. Allen, IMA Fungus 7(1): 4 (2016)

97416470-0664-5FDC-9B8E-2D786B4811D2

Fig. 14

Figure 14.

Colletotrichum magnum (GUCC 25-0029, new record). a, b. Culture on OA (a-above, b-reverse); c. Conidiomata on OA; d, e. Conidiophore, conidiogenous cells giving rise to conidia; f. Chlamydospores; g–k. Conidia. Scale bars: 10 µm (d–k).

Description.

See Rossman et al. (2016) and Damm et al. (2019).

Material examined.

China • Yunnan Province, Dali, symptomatic fruit of Juglans regia L., 2023.11.5, coll. Zou Mengting, RH1-5/GZ13 (dried culture, HGUP 25-0019), living culture GUCC 25-0029; • ibid. symptomatic fruit of Juglans regia L., 2023.11.12, coll. Zou Mengting, DJ5-8/GZ15 (dried culture, HGUP 25-0020), living culture GUCC 25-0030.

Notes.

Colletotrichum magnum is a well-recognized pathogen responsible for anthracnose disease across a wide range of plant hosts, including both agricultural crops and ornamental species, causing considerable economic losses (Rossman et al. 2016; Damm et al. 2019). The species was first described as distinct based on its unique morphological characteristics and phylogenetic placement (Rossman et al. 2016). Key diagnostic features for identifying C. magnum include its cylindrical conidia and the structure of its cylindrical to ellipsoidal conidiogenous cells, supported by molecular phylogenetic analyses using multiple genes (Rossman et al. 2016; Damm et al. 2019).

In China, Colletotrichum species have been commonly associated with Juglans regia (English walnut), causing leaf spots and lesions that reduce photosynthetic capacity and ultimately impact yield (Zhu et al. 2014; Wang et al. 2017; He et al. 2019; Li et al. 2023, 2024; Zhang et al. 2023b). Although the full pathogenic potential of C. magnum remains under investigation, it is increasingly regarded as an emerging threat to walnut production, especially in regions where walnuts are economically important. In the present study, strains GUCC 25-0029 and GUCC 25-0030, isolated from symptomatic fruit of J. regia, were identified as Colletotrichum magnum based on morphological characteristics (Fig. 14) and multilocus phylogenetic analyses (Fig. 13). Morphologically, the strain produced an asexual morph on OA, with transparent to dark brown, branched, and septate hyphae. Conidiogenous cells were cylindrical to ellipsoidal, measured 25.5–30 × 3.5–4.5 µm, solitary or clustered, hyaline, and unbranched, formed on the hyphae. Conidia were hyaline, cylindrical, straight, the apex and base rounded, measured 7.5–17.5 × 2.5–4.5 µm, hyaline, with smooth walls and no septa (Fig. 14). Phylogenetic analyses placed the new strain within the C. magnum clade with support, 97% ML, 96% MP, and 1.0 PP (Fig. 13). This represents the first report of C. magnum infecting Juglans regia in China. However, to confirm its pathogenic role on J. regia, future pathogenicity tests are required.

Figure 13.

Phylogenetic tree constructed using a maximum likelihood (ML) analysis based on a combined ITS, gapdh, chs-1, act, β-tubulin, and his3 sequences, representing Colletotrichum magnum species complex. The tree topology of the ML analysis was identical to the Maximum Parsimony (MP) and Bayesian posterior probability (BI) analysis. The final RAxML tree with a likelihood value of -6752.888091is presented here. The evolutionary model GTR+GAMMA was applied to all the genes. The analysis included twenty-three (23) taxa with a total of 2449 characters, with 434 distinct alignment patterns, and 16.59% were gaps and undetermined characters. Estimated base frequencies were as follows: A = 0.221259, C = 0.309082, G = 0.253858, T = 0.215801; substitution rates AC = 0.914360, AG = 2.159987, AT = 0.604764, CG = 0.664359, CT = 4.006750, GT = 1.0; gamma distribution shape parameter α = 0.280788; tree length = 0.381194. Bootstrap support values for ML and MP ≥ 70% and Bayesian Posterior Probabilities (BI) ≥ 0.90 are indicated at the nodes as ML/MP/BI. The tree is rooted with C. dracaenophilum (CBS 118199) and C. yunnanense (CBS 13213). Type strains are denoted in bold and T and sequences generated in this study are in yellow. Bar = 0.02 represents the estimated number of nucleotide substitutions of site per branch.

Discussion

Identification of Colletotrichum species is vital for understanding their biology, host range, pathogenic potential, and for developing effective disease management strategies. Although morphological traits such as conidial shape, size, and colony appearance have traditionally been used for species delimitation, these features often overlap among taxa, especially within species complexes, making morphology-based identification unreliable (Damm et al. 2012a, b; Cannon et al. 2012). Molecular data have become essential for resolving taxonomic ambiguities in Colletotrichum (Marin-Felix et al. 2017; Bhunjun et al. 2021; Jayawardena et al. 2021; Liu et al. 2022). Multilocus phylogenetic analyses using markers such as ITS, gapdh, act, chs-1, β-tubulin, cal, his3, and ApMat have significantly improved species resolution, especially within the C. gloeosporioides, C. acutatum, and C. boninense complexes (Weir et al. 2012; Jayawardena et al. 2021; Liu et al. 2022). These methods are crucial for distinguishing cryptic species, correcting misidentifications, and resolving invalid taxa, particularly in a genus exhibiting high morphological plasticity.

In the present study, our phylogenetic analyses confirmed that four newly collected strains clustered within a robust clade that includes Colletotrichum jiangxiense, C. nullisetosum, C. oblongisporum, C. gracile, and C. tengchongense. These findings align with earlier reports by Liu et al. (2015) and Yu et al. (2022), who highlighted morphological similarity and close phylogenetic placement among these taxa. However, C. nullisetosum, C. oblongisporum and C. gracile are now considered nomenclaturally invalid (nom. inval.) due to violations of the International Code of Nomenclature, and based on our morphological and molecular data, we support their synonymy under C. jiangxiense. This clarification builds on previous taxonomic suggestions and provides new evidence from fresh collections in Guizhou and Yunnan. Colletotrichum tengchongense, described by Zheng et al. (2022), was initially recognized as a distinct taxon due to its lageniform to ampulliform conidiogenous cells and subtle colony differences. In our analyses, however, C. tengchongense shows only minimal nucleotide differences when compared with C. jiangxiense (ITS = 0% [0/515], gapdh = 0% [0/274], chs-1 = 0.3% [1/287], act = 0.4% [1/246], β-tubulin = 0% [0/700], cal = 0.1% [1/715], ApMat = 0.2% [2/870]). These differences fall well within the range of intraspecific variation commonly reported for Colletotrichum species complexes (Cannon et al. 2012; Hyde et al. 2014; Jayawardena et al. 2021). Moreover, phylogenetic analyses consistently recover C. tengchongense and C. jiangxiense as a monophyletic lineage, without evidence of independent sub-clade formation or unique lineage divergence. Similar cases of low genetic divergence but close phylogenetic clustering have previously been used to synonymize taxa in Colletotrichum (Damm et al. 2012a, b; Liu et al. 2022). Taken together, both sequence identity and phylogenetic topology provide strong evidence that C. tengchongense does not represent a distinct species and should be treated as a later synonym of C. jiangxiense. Our data also support the synonymization of C. speciosum and C. simulanticitri with C. nymphaeae, corroborating the conclusions of Yu et al. (2022). Both taxa clustered tightly with C. nymphaeae, showing negligible sequence divergence. Colletotrichum simulanticitri, although based on four loci, exhibited low genetic distances and evidence of recombination, justifying its synonymy. Meanwhile, C. speciosum, described from ITS data only, lacked the molecular support necessary for species distinction. Both names are also currently listed as nom. inval. due to violations of the International Code of Nomenclature. They are considered synonyms here based on molecular and taxonomic criteria. This finding strengthens earlier assumptions based on partial gene data and provides a more comprehensive resolution using a broader phylogenetic framework. Altogether, the integration of our current results with previous taxonomic and phylogenetic studies highlights the crucial role of molecular data in resolving species boundaries in Colletotrichum. Given the high morphological plasticity and broad host associations across the genus, molecular tools remain essential for accurate species identification and understanding species complexes (Liu et al. 2017; de Silva et al. 2019; Peng et al. 2023; Zhang et al. 2023a, b, c; Trkulja et al. 2024; Lu et al. 2025).

This study significantly contributes to our understanding of Colletotrichum diversity in southwestern China by presenting a novel species, Colletotrichum macroconidii, alongside 11 previously known species, including six new host and regional records. The integration of detailed morphological analyses with multilocus phylogenetic data allowed us to clarify the taxonomic relationships of these species, offering valuable insights into their ecological and pathogenic roles in southwestern China (Hyde et al. 2024; Damm et al. 2009).

Our phylogenetic analyses revealed support for the distinctiveness of the newly described species, C. macroconidii, which was consistently placed as a sister clade to C. gardeniae within the C. gloeosporioides species complex. This novel taxon is characterized by its large, aseptate conidia, a feature that differentiates it from closely related species within the same complex. The phylogenetic distinctiveness of C. macroconidii was further supported by significant genetic variations, along with the absence of recombination between C. macroconidii and its closest relative, as demonstrated by the Genealogical Concordance Phylogenetic Species Recognition (GCPSR) model and the pairwise homoplasy index (Φw) test (Quaedvlieg et al. 2014; Damm et al. 2019). These results provide robust molecular evidence for recognizing C. macroconidii as a separate species within the C. gloeosporioides complex.

Additionally, we report six new host or geographic records for China: C. fioriniae, C. trichellum, C. juglandicola, C. nanhuaense, C. jiangxiense, and C. magnum. Notably, C. trichellum is reported for the first time from Hedera spp. in China, marking an important expansion of its known host range. Colletotrichum trichellum has been previously reported from various countries, including Canada, Germany, and the United Kingdom, where it causes leaf and stem spots on Hedera helix (English ivy) (Jayawardena et al. 2021). Our finding extends the host range of C. trichellum and highlights its potential to impact ornamental plants in China. This is particularly concerning as Hedera helix is widely used in landscaping, and the pathogen could have significant economic and ecological effects (Jayawardena et al. 2021). Similarly, C. juglandicola, first described from Juglans regia (walnut) in China, was detected in new host plants, including Camellia japonica and Ilex spp. These findings underscore the broad host range of C. juglandicola and its emerging importance as a pathogen of ornamental species in China (Zhang et al. 2023a). Additionally, C. magnum, a known anthracnose pathogen (Rossman et al. 2016; Damm et al. 2019), was identified from Juglans regia for the first time in China. Given the importance of walnut cultivation in the region, this represents a significant addition to the list of Colletotrichum species affecting economically valuable crops (Rossman et al. 2016).

New host associations were also confirmed for C. fioriniae, C. nanhuaense, and C. jiangxiense, which were identified on new hosts, Parthenocissus tricuspidata and Actinidia chinensis (kiwifruit). These findings further emphasize the ecological adaptability of Colletotrichum species and their ability to infect diverse plant species across different regions. In particular, C. fioriniae, traditionally associated with fruits like pear, has now been recorded infecting P. tricuspidata, highlighting the need for continued surveillance of its host range (Yu et al. 2022; Wang et al. 2025).

In conclusion, our integrative approach, combining multilocus phylogenetic analyses with morphological evidence, not only confirms species identities but also clarifies synonymy and reveals new host-pathogen associations (Dean et al. 2012; Jayawardena et al. 2020). This study reinforces the essential role of molecular phylogenetics in modern fungal taxonomy and expands the known diversity of Colletotrichum in China, with implications for agriculture, forestry, and biodiversity conservation.

Citation

Norphanphoun C, Zou M-T, Liu F-Q, Wang Y (2025) Colletotrichum macroconidii sp. nov. and six new records of Colletotrichum (Glomerellaceae, Glomerellales) from southwestern China. MycoKeys 122: 321–373. https://doi.org/10.3897/mycokeys.122.161122

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This work was supported by the Natural Science Special Research Fund of Guizhou University, Special Post (Gui Da Ling Jun He Zi [2024] 07), National Natural Science Foundation of China (No. 31972222), Program of Introducing Talents of Discipline to Universities of China (111 Program, D20023), Guizhou Science, Technology Department of International Cooperation Base project ([2018]5806), Guizhou University Research and Innovation Team Project [2024]05 and Guizhou Science and Technology Innovation Talent Team Project ([2020]5001).

Author contributions

Data curation: Chada Norphanphoun. Sample collection: Meng-Ting Zou. Formal analysis: Chada Norphanphoun. Funding acquisition: Yong Wang, Feng-Quan Liu. Supervision: Yong Wang. Writing – original draft: Chada Norphanphoun. Writing – review and editing: Chada Norphanphoun, Yong Wang.

Author ORCIDs

Chada Norphanphoun https://orcid.org/0000-0002-5756-7206

Meng-Ting Zou https://orcid.org/0009-0005-5564-4131

Yong Wang https://orcid.org/0000-0003-3831-2117

Data availability

All of the data that support the findings of this study are available in the main text.