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Research Article
Three new species of Camporesiomyces (Tubeufiaceae, Tubeufiales, Dothideomycetes) associated with coffee in Yunnan, China
expand article infoMei-Yan Han‡, Samantha C. Karunarathna‡, Li Lu‡§|, De-Ge Zheng‡, Nakarin Suwannarach#, Abdallah M. Elgorban¤, Dong-Qin Dai‡, Li-Juan Zhang§|, Wan-Tong Zhao‡, Ekachai Chukeatirote§, Saowaluck Tibpromma‡
‡ College of Biology and Food Engineering, Qujing Normal University, Qujing, China
§ School of Science, Mae Fah Luang University, Chiang Rai, Thailand
| Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand
¶ Center of Excellence in Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai, Thailand
# Office of Research Administration, Chiang Mai University, Chiang Mai, Thailand
¤ King Saud University, Riyadh, Saudi Arabia

Corresponding author: Ekachai Chukeatirote ( ekachai@mfu.ac.th )

Corresponding author: Saowaluck Tibpromma ( saowaluckfai@gmail.com )

Academic editor: Ajay Kumar Gautam
© 2025 Mei-Yan Han, Samantha C. Karunarathna, Li Lu, De-Ge Zheng, Nakarin Suwannarach, Abdallah M. Elgorban, Dong-Qin Dai, Li-Juan Zhang, Wan-Tong Zhao, Ekachai Chukeatirote, Saowaluck Tibpromma.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Han M-Y, Karunarathna SC, Lu L, Zheng D-G, Suwannarach N, Elgorban AM, Dai D-Q, Zhang L-J, Zhao W-T, Chukeatirote E, Tibpromma S (2025) Three new species of Camporesiomyces (Tubeufiaceae, Tubeufiales, Dothideomycetes) associated with coffee in Yunnan, China. MycoKeys 117: 315-336. https://doi.org/10.3897/mycokeys.117.154573
Open Access

Abstract

During our surveys of microfungi associated with coffee plants in Yunnan Province, China, three saprobic fungi were isolated from dead coffee branches. Multigene phylogenetic analyses (ITS, LSU, tef1-α, and rpb2) and morphological characteristics resulted in the identification of three novel species in Camporesiomyces, namely C. bhatii, C. coffeae, and C. puerensis. Detailed morphological descriptions, illustrations, and phylogenetic analyses of these three new species are provided, along with morphological comparisons to closely related taxa. These findings have global implications for understanding the diversity of microfungi associated with coffee trees.

Key words:

Coffee-associated microfungi, morphology, multigene phylogeny, taxonomy, three novel species

Introduction

Coffee (roasted beans of Coffea) is one of the most widely consumed beverages globally and ranks as the second most valuable traded commodity after crude oil (Ochoa-Henriquez et al. 2024). Originally from the tropical forests of Ethiopia, coffee is now cultivated globally across equatorial regions, particularly in frost-free areas with adequate rainfall and well-drained soils (Davis et al. 2006). Coffee-producing countries worldwide have their own unique cultural heritage and production methods (Buzzanell 1979). To date, approximately 177 varieties of coffee are available worldwide [Global Biodiversity Information Facility database (GBIF), available at: https://www.gbif.org/species/2895315 (accessed March 4, 2025)]. However, only a few have commercial value due to their moderate caffeine content and distinctive flavor (Wintgens, 2012). Coffea arabica and Co. canephora are the most widely cultivated coffee species, with Co. arabica accounting for approximately 60–70% and Co. canephora around 30–40% of global coffee production. In comparison, Co. liberica is grown in a few countries and contributes less than 1% (Bilen et al. 2022). Coffea arabica is prized for its mild flavor and aromatic complexity, while Co. eugenioides, a progenitor of Co. arabica, contributes genetic diversity critical for breeding programs (Scalabrin et al. 2020). Coffea canephora is valued for its higher caffeine content and its resilience to pests and climatic fluctuations (Davis et al. 2006; Avelino et al. 2015). Coffea liberica, is a plant of the humid, lowland tropics (Soerianegara and Lemmens 1993), characterized by its large beans and resistance to certain diseases (Melia et al. 2025).

The history of coffee cultivation in China can be traced back to 1884, when British merchants first introduced coffee to Taiwan Province of China. The earliest coffee cultivation on mainland China began in the early 20th century when French missionaries brought coffee seedlings to Binchuan County, Yunnan Province, for cultivation (Zhang et al. 2014). Coffee production in Yunnan Province accounts for approximately 90% of China’s total coffee production (Neilson and Wang 2019). Pu’er City is one of the largest coffee production areas in China, with the largest planting area and output. It primarily grows Arabica coffee, which enjoys a high reputation in the international market for its unique flavor and high quality (Li 2014). Xishuangbanna Autonomous Prefecture is also one of China’s important coffee-producing areas. Coffee planting there is mainly concentrated in Jinghong City, Menghai County, and Mengla County (Rigal et al. 2018).

Microfungi play a crucial role in agricultural ecosystems and are key nutrient regulating organisms in coffee ecosystems. They are globally distributed and exhibit high diversity (Bahram & Netherway, 2022). In recent years, scientists have conducted in-depth research on coffee-related fungi, identifying approximately 600 species (Lu et al. 2022b; Poma-Angamarca et al. 2024). However, relatively less attention has been paid to saprophytic fungi associated with coffee, with only more than 60 species reported globally to date (Han et al. 2024; Lu et al. 2025). Coffee-associated saprophytic fungi play a crucial ecological role in ecosystems and have also attracted researchers’ interest in recent years due to their potential for biocontrol (Lu et al. 2022a). For example, Rodríguez et al. (2016) demonstrated that coffee saprobes, as decomposers, significantly reduce anthracnose incidence in Coffea arabica seedlings through pathogen suppression, indicating their potential as biocontrol agents. In addition, Rodríguez et al. (2024) showed that certain yeast strains isolated from Robusta coffee significantly inhibit the growth of Aspergillus carbonarius and the production of ochratoxin A (OTA). These strains demonstrated an 85% inhibition rate of pathogen growth and a 90% toxin degradation efficiency in vitro. Thus, investigating the diversity of coffee saprophytic fungi is crucial, as it lays the groundwork for future studies exploring their potential applications in sustainable coffee farming.

Tubeufiales was established by Boonmee et al. (2014) based on phylogenetic analyses and morphological characteristics to accommodate the monophyletic family Tubeufiaceae, which has previously been classified in Pleosporales. Later, two additional families, Bezerromycetaceae and Wiesneriomycetaceae were incorporated into Tubeufiales by Liu et al. (2017) based on divergence time estimates. The latest comprehensive revision of Tubeufiales was conducted by Lu et al. (2018b), who expanded the circumscription of the type family Tubeufiaceae and revised the taxonomy of tubeufiaceous species. This revision notably helped highlight cases of misidentified fungi and determine which morphological characters are crucial for classifying species accurately. Members of Tubeufiales (Bezerromycetaceae, Tubeufiaceae, and Wiesneriomycetaceae) exhibit a widespread distribution, ranging from temperate to tropical regions (Boonmee et al. 2014; Liu et al. 2017; Xu et al. 2022; Hyde et al. 2024), and are commonly found on decorticated or decaying woody and herbaceous substrates as saprobic fungi. They often coexist with other decaying fungi (Lu et al. 2018b).

Tubeufiaceae was established by Barr (1979) based on the type genus Tubeufia to accommodate some bitunicate ascomycetes that grow on decaying wood. Members of Tubeufiaceae are widely distributed in tropical, subtropical, and temperate regions, and are often found on woody substrates in both terrestrial and freshwater habitats (Boonmee et al. 2011, 2014; Luo et al. 2017; Lu et al. 2018b, Tibpromma et al. 2018; Ma et al. 2024). A total of 47 genera are listed in Tubeufiaceae (Hyde et al. 2024), with Tubeufia being the largest genus. In addition, some studies have demonstrated that members of the Tubeufiaceae can produce biologically active secondary metabolites with antifungal, antibacterial, antidiabetic, and anticancer properties (Itazaki et al. 1990; Hanada et al. 1996; Hu et al. 2006; Lu et al. 2018; Zhang et al. 2023; Zhang et al. 2024).

Based on integrated morphological and phylogenetic analyses of combined ITS, LSU, tef1-α, and rpb2 sequences, Camporesiomyces was proposed by Hyde et al. (2020) to accommodate a new species, C. mali (type species), and two new combinations, C. patagonicus and C. vaccinii (formerly belonging to the genera Acanthostigma and Helicoma, respectively) (Carris 1989; Sanchez et al. 2012). Morphologically, the type species, C. mali, is distinguished from other members of Tubeufiaceae by its unique multi-loculate ascomata and narrowly fusiform, hyaline, multi-septate ascospores (Hyde et al. 2020). The asexual morph of Camporesiomyces is characterized by monoblastic or polyblastic conidiogenous cells, which are denticulate, with helical and hyaline conidia that are smooth-walled (Carris 1989). Camporesiomyces patagonicus and C. vaccinii were previously classified as Acanthostigma patagonicum and Helicoma vaccinii, respectively (Carris 1989; Sanchez et al. 2012). However, the natural classification of these two taxa has been uncertain (Boonmee et al. 2014; Lu et al. 2018). Although the holotypes of A. patagonicum and H. vaccinii have not been rechecked, the DNA data derived from the ex-type strains show they belong to Camporesiomyces (Hyde et al. 2020). Currently, the genus comprises only three species (C. mali, C. patagonicus, and C. vaccinii), all of which exhibit consistent morphological autapomorphies and molecular divergence that warrant their segregation from allied genera (Hyde et al. 2020).

In this study, we aim to introduce three new species of Camporesiomyces found on dead coffee branches collected from Pu’er and Xishuangbanna, Yunnan Province, China. Morphological characteristics and multi-locus phylogeny analyses, based on combined ITS, LSU, tef1-α, and rpb2 sequence data, were used to confirm the taxonomic placement of the new species.

Materials and methods

Specimen collection, examination, and isolation

Dead branches of Arabica Coffee and Liberica Coffee with fungal fruiting bodies were collected from Pu’er City and Xishuangbanna Autonomous Prefecture, Yunnan Province, China. The specimens were placed in self-sealing bags, with sampling information recorded on the cover (Rathnayaka et al. 2025) and transported to the mycology laboratory at Qujing Normal University. The fruiting body structures were observed using a LEICA S8 APO optical microscope (Olympus, Tokyo, Japan). Micro-morphological characteristics were observed with a compound microscope (OLYMPUS BX53, Olympus, Tokyo, Japan), and photographed with an OLYMPUS DP74-CU camera fitted onto the microscope. Measurement was made using the Tarosoft (R) Image Framework v. 0.9.7 program. The photo plate was created in Adobe Photoshop CS3 Extended version 10.0 software (Adobe Systems, USA). Single spore isolation was performed following the method outlined by Senanayake et al. (2020) to obtain pure cultures on potato dextrose agar (PDA). Dried herbarium samples were preserved at the Herbarium of Guizhou Medical University, Guiyang, China (GMB-W). Living cultures were deposited in the Guizhou Medical University Culture Collection (GMBCC), Guiyang. The Facesoffungi (FoF) and Index Fungorum (IF) numbers were obtained as explained in Jayasiri et al. (2015) and Index Fungorum (2025), respectively.

DNA extraction, PCR amplification, and sequencing

Fresh mycelia grown on PDA (15–30 days) were scraped and transferred to 1.5 ml microcentrifuge tubes for DNA extraction. The Biospin Fungus Genomic DNA Extraction Kit BSC14S1 (BioFlux, China) was used to obtain DNA products from the above mycelia, according to the manufacturer’s protocol. The extracted DNA was then preserved at -20 °C for future use. Polymerase chain reaction (PCR) was performed for four loci; the details of the different loci, primers, and PCR thermal cycle programs used for amplification are presented in Table 1. PCR amplification was performed in a reaction volume of 25 µL, consisting of 12.5 µL of 2× Bench Top™ Taq Master Mix, 8.5 µL of ddH2O, 2 µL of each forward and reverse primer, and 2 µL of the DNA template. The PCR products were purified and sequenced by Sangon Biotechnology Co., Ltd. (Shanghai, China). All newly generated sequences in this study were deposited in GenBank (http://www.ncbi.nlm.nih.gov/genbank/) and are listed in Table 2.

Table 1.
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Loci, primers, and amplification procedure used in this study.

Loci Primers PCR conditions References
ITS ITS5/ITS4 94 °C: 3 mins, (94 °C: 45 s, 55 °C: 50 s, 72 °C: 1 min) × 35 cycles, 72 °C: 10 mins White et al. (1990)
LSU LR5/LR0R Vilgalys and Hester (1990)
tef1-α 983F/2218R Rehner and Samuels (1994)
rpb2 5F/7cR 95 °C: 5 mins, (95 °C: 1 min, 55 °C: 2 min, 72 °C: 90 s) × 40 cycles, 72 °C: 10 mins Liu et al. (1999)
Table 2.
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Names, strain numbers, and corresponding GenBank accession numbers of the taxa used in the phylogenetic analyses. The newly generated sequences are in bold black. After the strain number, “T” indicates the type strains. “NA” indicates sequence unavailability.

Fungal species Strain numbers ITS LSU tef1-α rpb2
Acanthohelicospora aurea GZCC 16-0059 KY321322 KY321325 KY792599 MF589910
Acanthohelicospora aurea GZCC 16-0060 KY321323 KY321326 KY792600 MF589911
Acanthostigma chiangmaiense MFLUCC 10-0125T JN865209 JN865197 KF301560 NA
Acanthostigma multiseptatum ANM 475T GQ856145 GQ850492 NA NA
Acanthostigma multiseptatum ANM 665 GQ856144 GQ850493 NA NA
Acanthostigma perpusillum UAMH 7237T AY916492 AY856892 NA NA
Acanthostigma scopulum ANM 95 GQ856142 GQ850490 NA NA
Acanthostigma scopulum ANM 386 GQ856141 GQ850489 NA NA
Berkleasmium fusiforme MFLUCC 17-1978T MH558693 MH558820 MH550884 MH551007
Berkleasmium fusiforme MFLUCC 17-1979 MH558694 MH558821 MH550885 MH551008
Bezerromyces pernambucoensis URM7414 KX470393 KX518626 KX518634 NA
Bezerromyces pernambucoensis URM7412 KX470391 KX518624 KX518632 NA
Camporesiomyces bhatii GMBCC 1120T PQ763360 PQ842543 PV388888 PV388894
Camporesiomyces bhatii GMBCC 1125 PQ763361 PQ842544 PV388889 PV388895
Camporesiomyces coffeae GMBCC 1130T PQ763358 PQ842545 PV388890 PV388896
Camporesiomyces coffeae GMBCC 1131 PQ763359 PQ842546 PV388891 PV388897
Camporesiomyces mali KUMCC 19-0216T NR_169709 NG_075312 MN794018 NA
Camporesiomyces patagoniensis BBB MVB 573 JN127358 JN127359 NA NA
Camporesiomyces puerensis GMBCC 1113T PQ763356 PQ842541 PV388886 PV388892
Camporesiomyces puerensis GMBCC 1114 PQ763357 PQ842542 PV388887 PV388893
Camporesiomyces vaccinii CBS 216.90 MH862204 MH873889 NA NA
Chlamydotubeufia cylindrica MFLUCC 16-1130T MH558702 MH558830 MH550893 MH551018
Chlamydotubeufia huaikangplaensis MFLUCC 16-0023 KY678766 KY678758 KY792597 MF535259
Dematiohelicosporum guttulatum MFLUCC 17-2011T MH558705 MH558833 MH550896 MH551021
Dictyospora thailandica MFLUCC 16-0001T KY873627 KY873622 KY873286 MH551023
Dictyospora thailandica MFLUCC 18-0641 MH558706 MH558834 MH550897 MH551022
Helicoarctatus aquaticus MFLUCC 17-1996T MH558707 MH558835 MH550898 MH551024
Helicoma muelleri CBS 964.69T AY916453 MH871278 NA NA
Helicoma nematosporum MFLUCC 16-0011 MH558722 MH558848 MH550913 MH551039
Helicoma rubriappendiculatum MFLUCC 18-0491T MH558723 NG_069593 MH550914 MH551040
Helicomyces hyalosporus GZCC 16 0070 MH558728 MH558854 MH550919 MH551044
Helicomyces torauatus MELUCC16-0217 MH558732 MH558858 MH550923 MH551048
Helicosporium nanningense GZCC 22-2175T OR066418 OR066425 OR058864 OR058857
Helicosporium nanningense GZCC 23-0588 OR066419 OR066426 OR058865 OR058858
Helicosporium setiferum MFLUCC 17-1994T MH558735 MH558861 MH550926 MH551051
Helicosporium setiferum MFLUCC 17-2007 MH558737 MH558863 MH550928 MH551053
Helicosporium sexuale GZCC 22-2007 OP508731 OP508771 OP698082 OP698071
Helicosporium sexuale MFLUCC 16-1244 MZ538503 MZ538537 MZ567082 MZ567111
Helicosporium vegetum BCC 8125 AY916491 MH871277 NA NA
Helicosporium vegetum CBS 254.75 NA DQ470982 DQ471105 NA
Helicosporium vegetum CBS 941.72T AY916488 AY856883 NA NA
Helicosporium vesicarium MFLUCC 17-1795T MH558739 MH558864 MH550930 MH551055
Helicosporium viridiflavum MFLUCC 17-2336T MH558738 NA MH550929 MH551054
Helicotubeufia guangxiensis MFLUCC 17-0040T MH290018 MH290024 MH290029 MH290034
Helicotubeufia hydei MFLUCC 17-1980T MH290021 MH290026 MH290031 MH290036
Kamalomyces thailandicus MFLUCC 13-0233T MF506884 MF506882 MF506886 NA
Kamalomyces thailandicus MFLUCC 11-0158 MF506883 MF506881 MF506885 MF506887
Neoacanthostigma fusiforme MFLUCC 11-0510T KF301529 KF301537 NA NA
Neohelicomyces aquaticus KUMCC 15-0463 KY320529 KY320546 KY320562 MH551065
Neotubeufia krabiensis MFUCC 16-1125T MG012031 MG012024 MG012010 MG012017
Thaxteriella inthanonensis MFLUCC 11-0003T JN865211 JN865199 NA NA
Thaxteriellopsis lignicola MFLUCC 16-0026 MH558768 MH558893 MH550960 MH551094
Thaxteriellopsis lignicola MELUCC 16-0024 MH558767 MH558892 MH550959 MH551093
Tubeufia bambusicola MFLUCC 17-1803T MH558771 MH558896 MH550963 MH551097
Tubeufia javanica MFLUCC 12-0545T KJ880034 KJ880036 KJ880037 NA

Phylogenetic analyses

The assembly of the forward and reverse primers for the recently obtained sequence was accomplished using BioEdit version 7.0.5.3 and SeqMan version 7.0.0 software packages (DNASTAR, Madison, WI) (Plasterer 1997; Hall 1999). For phylogenetic analysis, relevant sequences were sourced from GenBank based on BLASTn search results and in accordance with the most current literature (Tanaka et al. 2017; Hyde et al. 2018). The sequence data were aligned using the online multiple alignment program MAFFT version 7 (https://mafft.cbrc.jp/alignment/server/) (Katoh and Standley 2013) to ensure that all sequences were in the correct orientation. The alignments were automatically adjusted using trimAL v1.2 (Capella-Gutiérrez et al. 2009) and manually optimized with BioEdit v7.0.5.3 (Hall 1999). The multiple gene sequences were combined using the SequenceMatrix-Windows-1.7.8 software. The aligned multigene sequences were transformed into the PHYLIP format for Maximum Likelihood (ML) analysis and the NEXUS format for Bayesian Inference (BI) using the online utility ALTER (ALignment Transformation EnviRonment) (Glez-Peña et al. 2010). Maximum likelihood (ML) and Bayesian inference (BI) analyses were performed using the online CIPRES Science Gateway platform (https://www.phylo.org/portal2/home.action) (Dissanayake et al. 2020; Ma et al. 2023). The ML tree was constructed using the RAxML-HPC v.8 tool on XSEDE (version 8.2.12) (Stamatakis 2006) with the GTRGAMMA model and 1,000 bootstrap pseudoreplicates. The BI was carried out using the tool MrBayes on XSEDE (3.2.7a) (Huelsenbeck and Ronquist 2001; Stamatakis et al. 2008; Ronquist et al. 2012), using the Markov Chain Monte Carlo (MCMC) method, six simultaneous Markov chains were run for 2,000,000 generations, and trees were sampled at every 100th generation. Prior to conducting BI, the model of evolution for each gene region was estimated using MrModelTest version 2 (Ma 2016), The phylogenetic trees were edited by FigTree v. 1.4.0 (http://tree.bio.ed.ac.uk/software/figtree/).

Results

Phylogenetic analyses

Three new species (Camporesiomyces bhatii, C. coffeae, and C. puerensis) formed a distinct clade within Camporesiomyces, with strong statistical support (ML = 100%, PP = 1.00). Camporesiomyces clade was sister to Helicosporium within the family Tubeufiaceae, and all genera in Tubeufiaceae are distinctly separated (Fig. 1). The RAxML tree was based on a combined dataset of ITS, LSU, tef1-α, and rpb2 gene sequence data, which comprised 3,255 characters (ITS:1– 437, LSU: 438–1257, rpb2: 1258–2357, tef1-α: 2358–3255), including gaps. Bezerromyces pernambucoensis (URM7414) and B. pernambucoensis (URM7412) were used as outgroup taxa. The topology of the phylogenetic tree constructed using the maximum likelihood (ML) method exhibited a high degree of similarity to that generated by Bayesian inference (BI). Consequently, the ML tree was selected to illustrate the evolutionary history of Camporesiomyces.

Figure 1. 

The ML analysis and Bayesian inference (BI) analyses yielded nearly identical tree topologies, with bootstrap support values for ML equal to or greater than 50%, and BI analysis values equal to or greater than 0.90 PP are provided at each node. Newly generated sequences are in red, while the ex-type strains are in bold.

The best-scoring RAxML tree, with a final ML optimization likelihood value of -26167.982465 is presented. The matrix contained 1,266 distinct alignment patterns, with 21.89% of the characters being undetermined or gaps. Estimated base frequencies were as follows: A = 0.243800, C = 0.255403, G = 0.265360, T = 0.235437; substitution rates: AC = 0.936178, AG = 5.168669, AT = 2.168333, CG = 0.811291, CT = 10.165688, GT = 1.000000; gamma distribution shape parameter α = 0.816594. The phylogenetic tree resulting from RAxML analysis is shown in Fig. 1.

According to the phylogenetic tree, our three new species (Camporesiomyces bhatii, C. coffeae, and C. puerensis) stand out as distinct entities within the genus Camporesiomyces. Among these, C. bhatii (GMBCC 1120 (ex-type) and GMBCC 1125) formed a sister lineage to C. vaccinii (CBS 216.90) with 96% ML/0.99 PP support. The other two new species, C. puerensis (GMBCC 1113 (ex-type) and GMBCC 1114) and C. coffeae (GMBCC 1130 (ex-type) and GMBCC 1131), clustered together but formed well-separated branches (98% ML/1.00 PP support), further emphasizing their distinctiveness. These two new species create a distinct branch that diverges from C. patagoniensis (BBB MVB 573) with 70% ML/0.99 PP support.

Taxonomy

Camporesiomyces bhatii M.Y. Han & Tibpromma, sp. nov.

Index Fungorum: IF903588
Facesoffungi Number: FoF17675
Fig. 2

Etymology.

The species epithet “bhatii” honors Prof. Jayarama Darbhe Bhat for his immense contributions to mycology.

Figure 2. 

Camporesiomyces bhatii (GMB-W1176, holotype) a, b colonies on the natural substrate c, d conidiophore with conidia and conidiogenous cells e, f conidiogenous cells g–m conidia n a germinated conidium o, p Culture on PDA from above and reverse. Scale bars: 50 μm (c); 20 μm (d); 10 μm (e, g–n); 5 μm (f).

Holotype.

GMB-W1176

Description.

Saprobic on dead branches of Coffea liberica. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate superficial, solitary or in clusters, hairy, yellow at apex, velvety. Mycelium exposed on the surface of substrate, except on the roots. Conidiophores 23–87.4 × 2.4–5.7 μm (x̄ = 63 × 3.7 μm, n = 30) macronematous, mononematous, either solitary or forming small clusters, smooth or occasionally verruculose, cylindrical, brown, slightly flexuous, simple, unbranched, 2–8-septate, sometimes slightly constricted at the septa. Conidiogenous cells 8.8–21.7 × 3–4.7 μm (x̄ = 17 × 3.5 μm, n = 20), polyblastic, terminal, cylindrical, slightly tapering, conspicuously denticulate on conidial secession, pale brown. Conidia 16–30 × 3.3–6.3 μm (x̄ = 24 × 6.5 μm, n = 30), solitary, acrogenous, cylindrical, obclavate or fusiform, 3–8-septate, subhyaline to pale brown.

Culture characteristics.

Conidia germinate on PDA within 24 h at 28 °C, colony on PDA reaching 3 cm diam. after two weeks, circular or irregular, umbonate, with the entire margin dark brown to black. The reverse side displays predominantly black colonies with brown peripheral edges.

Material examined.

China • Yunnan Province, Xishuangbanna Autonomous Prefecture (21°41'N, 101°25'E 570 m), on dead branches of Coffea liberica, 27 August 2024, M.Y. Han & Tibpromma, (DL4 = GMB-W1176, holotype), ex-type GMBCC 1120, other living culture GMBCC 1125.

GenBank number.

GMBCC 1120 = ITS: PQ763360, LSU: PQ842543, tef1-α: PV388888, rpb2: PV388894 and GMBCC 1125 = ITS: PQ763361, LSU: PQ842544, tef1-α: PV388889, rpb2: PV388895.

Notes.

In the phylogenetic tree, our collection Camporesiomyces bhatii [GMBCC 1120 (ex-type) and GMBCC 1125] formed a well-separated lineage, sister to C. vaccinii [CBS 216.90, ex-type] with 96% ML/0.99 PP support (Fig. 1). ITS and LSU gene sequences blast results showed that our strain showed 94.14% and 99.23% similarities to C. vaccinii (MH862204) and (MH873889), respectively (Carris 1989). In the morphological comparison between C. bhatii and C. vaccinii, significant differences are observed in both conidial and conidiophore dimensions, as well as conidial shape (Carris 1989). The conidia of C. bhatii are notably longer than those of C. vaccinii (16–30 × 3.3–6.3 μm) compared to (8.0–13.0 × 2.0–4.0 μm). Conversely, the conidiophores of C. bhatii are shorter than those of C. vaccinii (23–87.4 × 2.4–5.7 μm) compared to (64–145 × 4.2–5.0 μm) (Carris 1989). Additionally, the conidia of C. bhatii exhibit complex shapes, such as obclavate or fusiform, and are subhyaline to pale brown. In contrast, the conidia of C. vaccinii are spirally coiled and hyaline or pale brown (Carris 1989). These distinct morphological characteristics provide a clear basis for differentiating the two species. Therefore, based on molecular data and morphological comparison, C. bhatii was identified as a new species.

Camporesiomyces coffeae M.Y. Han &Tibpromma, sp. nov.

Index Fungorum: IF903589
Facesoffungi Number: FoF17676
Fig. 3

Etymology.

The species epithet “coffeae” refers to the host genus Coffea.

Figure 3. 

Camporesiomyces coffeae (GMB-W1181, holotype) a, b colonies on the natural substrate c, d conidiophore and conidiogenous cells e conidiogenous cells f–j conidia k a germinated conidium l, m culture on PDA from above and reverse. Scale bars: 50 μm (c); 20 μm (d); 5 μm (e); 10 μm (f–k).

Holotype.

GMB-W1181

Description.

Saprobic on dead branches of Coffea arabica. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate superficial, solitary or clusters, hairy, subhyaline at apex, bunch of flowers-like, shiny. Mycelium exposed on surface of substrate, except on the roots. Conidiophores 43–97 × 2.8–4.5 μm (x̄ = 68 × 3.7 μm, n = 30) macronematous, mononematous, either solitary or forming small clusters, smooth or occasionally verruculose, cylindrical, dark brown, slightly flexuous, simple, unbranched, with longitudinal striations in the upper part, 2–9-septate, sometimes slightly constricted at the septa. Conidiogenous cells 8–21 × 2.3–4.6 μm (x̄ = 15.4 × 3.4 μm, n = 30), polyblastic, terminal, cylindrical, with longitudinal striations, with several conspicuous denticles at apex, brown. Conidia 20–50 × 3.3–6.5 μm (x̄ = 26.7 × 4.5 μm, n = 30), solitary, acrogenous, guttules, cylindrical, fusiform, 3–7-septate, subhyaline or hyaline, obtuse or conical at both ends.

Culture characteristics.

Conidia germinating on PDA within 24 h at 28 °C, colony on PDA reaching 2 cm diam. after two weeks, circular or irregular, umbonate, with entire margin dark grey to black, irregular, the reverse side displays predominantly black colonies with brown peripheral edges.

Material examined.

China • Yunnan Province, Pu’er City, Simao District (22°35'53"N, 100°59'17"E, 1186.4 m), on dead branches of Coffea arabica, 8 August 2024, M.Y. Han & Tibpromma, (YYT15 = GMB-W1181, holotype), ex-type GMBCC 1130; other living culture GMBCC 1131.

GenBank number.

GMBCC 1130 = ITS: PQ763358 LSU: PQ842545 tef1-α: PV388890, rpb2: PV388896 and GMBCC 1131 = ITS: PQ763359 LSU: PQ842546 tef1-α: PV388891, rpb2: PV388897.

Notes.

Based on multi-gene phylogenetic analysis, Camporesiomyces coffeae [GMBCC 1130 (ex-type) and GMBCC 1131] is a distinct species, sister to C. puerensis [GMBCC 1113 (ex-type) and GMBCC 1114] with 98% ML/1.00 PP support (Fig. 1). Comparative genomic analysis demonstrated that the sequence similarity between C. coffeae and C. puerensis was 93.00% in the ITS locus, 98.33% in the LSU locus, 93.34% in the tef1-α locus, and 89.03% in the rpb2 locus (Table. 3). Morphologically, C. coffeae can be distinguished from C. puerensis by the dimensions and septation of conidiophores and conidia, in that the conidia and conidiophores of C. coffeae are smaller than C. puerensis (20–50 × 3.3–6.5 μm vs. 21.7–83 × 4–9.4 μm) and (43–97 × 2.8–4.5 μm vs. 52–176.5 × 2.8–5.6 μm), respectively (Figs 3, 4). In addition, the conidia of C. coffeae are subhyaline or hyaline, whereas those of C. puerensis are hyaline at their ends but yellow in the middle. Moreover, the conidiophores of C. coffeae are distinguished by longitudinal striations in their apical sections, a feature that is absent in C. puerensis. Camporesiomyces coffeae is similar to C. puerensis in having polyblastic, terminal, conidiogenous cells, which are brown (Figs 3, 4). Based on the differences between the two species, we describe C. coffeae as a new species.

Table 3.
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Nucleotide comparisons of C. bhatii (GMBCC 1120, ex-type), C. coffeae (GMBCC 1130, ex-type), and C. puerensis (GMBCC 1113, ex-type) based on ITS, LSU, tef1-α, and rpb2; all of them were compared, excluding gaps.

Species ITS (%) LSU (%) tef1-α (%) rpb2 (%)
C. puerensis vs. C. coffeae 44/628(7.0%) 15/900(1.67%) 61/914 (6.65%) 118/1076 (10.97%)
C. puerensis vs. C. bhatii 42/484 (8.68%) 17/895 (1.9%) 70/924 (7.58%) 165/1076 (15.76%)
C. coffeae vs. C. bhatii 37/515 (7.18%) 12/901 (1.33%) 76/969 (7.84%) 172/1070 (16.07%)

Camporesiomyces puerensis M.Y. Han &Tibpromma, sp. nov.

Index Fungorum: IF903590
Facesoffungi Number: FoF17677
Fig. 4

Etymology.

The name reflects the type location, “Pu’er” City, China.

Figure 4. 

Camporesiomyces puerensis (GMB-W1121, holotype) a, b colonies on the natural substrate c, d conidiophores and conidiogenous cells e, f conidiogenous cells g–l conidia m a germinated conidium n, o culture on PDA from above and reverse. Scale bars: 50 μm (c); 30 μm (d); 10 μm (e, f); 20 μm (g–m).

Holotype.

GMB-W1121

Description.

Saprobic on dead branches of Coffea arabica. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on the natural substrate are superficial, solitary or clusters, hairy, yellow at apex velvety. Mycelium exposed on the surface of the substrate, with the exception of the roots. Conidiophores 52–176.5 × 2.8–5.6 μm (x̄ = 88.4 × 4 μm, n = 30), macronematous, mononematous, solitary or forming small clusters, smooth or occasionally verruculose, cylindrical, brown, flexuous, simple, unbranched, 3–13-septate, sometimes slightly constricted at the septa. Conidiogenous cells 6.8–26 × 2.3–4.3 μm (x̄ = 15 × 3.6 μm, n = 20), polyblastic, terminal, cylindrical to slightly tapering, denticulate, smooth, slightly curved, pale brown. Conidia 21.7–83 × 4–9.4 μm (x̄ = 43.3 × 6.3 μm, n = 30), solitary, acrogenous, cylindrical, or fusiform, sometimes slightly curved, 4–9-septate, subhyaline to yellow, hyaline at both rostrate ends.

Culture characteristics.

Conidia germinating on PDA within 24 h at 28 °C. Colony on PDA reaching 2 cm diam. after two weeks, circular or irregular, umbonate, with entire margin dark brown to black, reverse side displays predominantly black colonies with brown peripheral.

Material examined.

China • Yunnan Province, Pu’er City, Simao District (22°36'36"N, 101°0'14"E, 1189 m), on dead branches of Coffea arabica, 8 August 2024, M.Y. Han & Tibpromma, (BG25 = GMB-W1121, holotype), ex-type GMBCC 1113, other living culture GMBCC 1114.

GenBank number.

GMBCC 1113 = ITS: PQ763356, LSU: PQ842541, tef1-α: PV388886, rpb2: PV388892 and GMBCC 1114 = ITS: PQ763357, LSU: PQ842542, tef1-α: PV388887 rpb2: PV388893.

Notes.

In the concatenated phylogenetic analysis, our strain Camporesiomyces puerensis [GMBCC 1113 (ex-type), and GMBCC 1114] formed a sister branch with C. coffeae [GMBCC 1130 (ex-type), and GMBCC 1131] with 98% ML/1.00 PP bootstrap support (Fig. 1), both taxa formed an independent branch under C. patagoniensis (BBB MVB 573) with 70% ML/0.99 PP statistical support. Nucleotide comparisons between C. puerensis (GMBCC 1113) and C. coffeae (GMBCC 1130) are shown in Table 3. Morphologically, C. patagoniensis exhibits the sexual morph, so the morphological comparisons with our strain were unavailable. Camporesiomyces puerensis shares a similar conidial shape with C. coffeae, but it can be distinguished by its conidial pigmentation and dimensions. The conidia of C. puerensis are subhyaline to pale green, contrasting with the subhyaline to hyaline conidia of C. coffeae (Figs 3, 4). In addition, C. puerensis produces longer and broader conidia than C. coffeae (21.7–83 × 4–9.4 μm vs. 20–50 × 3.3–6.5 μm). Camporesiomyces puerensis also produces longer and slightly wider conidiophores than C. coffeae (52–176.5 × 2.8–5.6 μm vs. 43–97 × 2.8–4.5 μm) (Figs 3, 4). Hence, we describe C. puerensis as a new species based on both morphology and phylogeny.

Discussion

In recent years, research on the diversity of microfungi on coffee plants has made significant progress, revealing their complexity and importance within the coffee ecosystem (dos Santos Gomes et al. 2024; Lu et al. 2025). These studies not only covered various ecological types, such as pathogens, endophytes, and saprophytes, but also emphasized the potential role of microfungi in coffee health and ecosystem function (Hyde et al. 2014; Bahram and Netherway 2022; Lu et al. 2022a; dos Santos Gomes et al. 2024). Pathogenic fungi are among the most widely studied microfungi in coffee ecosystems, while saprobic and endophytic fungi have received relatively less attention (Lu et al. 2022a; Poma-Angamarca et al. 2024; Lu et al. 2025). Saprophytic fungi play a crucial role in the coffee ecosystem functioning by decomposing organic matter and facilitating nutrient cycling (Dequn and Hyde 2001; Hyde et al. 2007). Some studies have suggested that the saprotrophic fungus Phialomyces macrosporus, isolated from leaf litter, could potentially be used in the management of coffee halo blight in seedlings (Botrel et al. 2018). Therefore, studying the diversity of coffee saprophytic fungi is essential. By characterizing these fungi, we can identify key species that efficiently decompose organic matter, improve soil structure, suppress pathogens, and ultimately support more resilient and productive coffee ecosystems.

In this study, we introduce three new asexual species of Camporesiomyces, viz., C. bhatii, C. coffeae, and C. puerensis, which were isolated from decaying coffee branches based on morphological and phylogenetic analyses. Camporesiomyces currently comprises three recognized species: C. mali, C. patagonicus (sexual morph), and C. vaccinii (asexual morph) (Carris 1989; Sanchez et al. 2012; Hyde et al. 2020). The three novel asexual species (C. bhatii, C. coffeae, C. puerensis) belonging to Camporesiomyces exhibit distinct morphological differences from the currently recognized taxa. Our three new Camporesiomyces species share similar morphological characteristics, including fusiform conidia and analogous conidiophores and conidiogenous cells, but differ primarily in their dimensional measurements. However, our three new species exhibit significant morphological differences from the asexual species C. vaccinii. Camporesiomyces vaccinii displays substantial morphological divergence, with spirally convolute conidia and conidiophores, characterized by multiple (1–3) percurrent proliferations (Carris 1989) (the detailed morphological distinctions are provided in Table 4). This discrepancy may be explained by geographical isolation (C. vaccinii occurring on Vaccinium elliotii stems in Georgia, USA) and host specificity (Carris 1989), suggesting environmental factors potentially drive morphological differentiation within the genus Camporesiomyces. The morphological similarity among our three species may, therefore, be attributed to their shared host and geographically proximate distribution.

Table 4.
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Morphological comparisons of asexual morphs in Camporesiomyces.

Species Conidiophores Conidiogenous cells Conidia Host/location References
C. bhatii (GMB-W1176) 23–87.4 × 2.4–5.7 μm, brown, 2–8-septate, sometimes slightly constricted at the septa. 8.8–21.7 × 3–4.7 μm, cylindrical, slightly tapering, conspicuously denticulate on conidial secession, pale brown. 16–30 × 3.3–6.3 μm, obclavate or fusiform, 3–8-septate, subhyaline to pale brown. Dead branches of Coffea liberica/China This study
C. coffeae (GMB-W1181) 43–97 × 2.8–4.5 μm, dark brown, with longitudinal striations in the upper part, 2–9-septate, sometimes slightly constricted at the septa. 8–21 × 2.3–4.6 μm, cylindrical, with longitudinal striations, with several conspicuous denticles at apex, brown. 20–50 × 3.3–6.5 μm, guttules, cylindrical, fusiform, 3–7-septate, subhyaline or hyaline, obtuse or conical at both ends. Dead branches of Coffea arabica/China This study
C. puerensis (GMB-W1121) 52–176.5 × 2.8–5.6 μm, brown, 3–13-septate, sometimes slightly constricted at the septa. 6.8–26 × 2.3–4.3 μm, cylindrical to slightly tapering, denticulate, smooth, slightly curved, pale brown. 21.7–83 × 4–9.4 μm, or fusiform, sometimes slightly curved, 4–9-septate, subhyaline to yellow, hyaline at both rostrate ends. Dead branches of Coffea arabica/China This study
C. vaccinii (LMC 0043) 64–145 μm long, 4.2–5.0 μm wide at base, 2.5–3.3 μm wide at apex, simple or rarely branched, dark brown with paleapical cells, occasionally roughened, 4–10 septate, usually with l–2 percurrent proliferations. 1.3–2.7 × 1.0–1.3 μm, mono-or polyblastic, denticulate, denticles. 8.0–13.0 μm, conidial filament coiled l.5–l.75 times, 4–8-septate, rounded at apex and tapering to a truncate base, 2.0–4.0 μm, hyaline to pale brown, smooth-walled. On stems of Vaccinium elliotii/America Carris (1989)

We present descriptions, illustrations, and phylogenetic analysis results to validate and confirm the novelty of the three species: C. bhatii, C. coffeae, and C. puerensis. This is the first report of Camporesiomyces fungi associated with coffee, contributing three new asexual species to the genus Camporesiomyces. Our findings significantly enrich the fungal diversity of Camporesiomyces and hold substantial scientific value for enhancing our understanding of the fungal community inhabiting coffee hosts.

Acknowledgements

Samantha C. Karunarathan and Saowaluck Tibpromma thank the High-Level Talent Recruitment Plan of Yunnan Province (High-End Foreign Experts program) and the Key Laboratory of Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River for their support. Nakarin Suwannarach thanks Chiang Mai University for the support. The authors extend their appreciation to the Researchers Supporting Project Number (RSP2025R56), King Saud University, Riyadh, Saudi Arabia. The authors thank Dr. Shaun Pennycook for his advice on new fungal names.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was supported by the Yunnan Provincial Department of Science and Technology “Zhihui Yunnan” plan (202403AM140023), the National Natural Science Foundation of China (Number 32260004) and the Researchers Supporting Project Number (RSP2025R56), King Saud University, Riyadh, Saudi Arabia.

Author contributions

Conceptualization: SCK, ST, and LL. Data curation: MYH, LL, WTZ, and LJZ. Formal analysis: SCK, ST, and NS. Funding acquisition: SCK, ST, and AME. Investigation: SCK and MYH. Methodology: MYH, ST, EC. Project administration: ST, SCK. Resources: MYH. Software: MYH, LL, DGZ, WTZ and LJZ. Supervision: ST and SCK. Validation: ST and SCK. Visualization: ST and MYH. Writing - original draft: MYH. Review, and editing: SCK, ST, EC, LL, DGZ, NS, AME, DQD, LJZ, and WTZ.

Author ORCIDs

Mei-Yan Han https://orcid.org/0009-0004-3705-5408

Samantha C. Karunarathna https://orcid.org/0000-0001-7080-0781

Li Lu https://orcid.org/0000-0003-0977-6414

De-Ge Zheng https://orcid.org/0009-0001-9199-6784

Nakarin Suwannarach https://orcid.org/0000-0002-2653-1913

Abdallah M. Elgorban https://orcid.org/0000-0003-3664-7853

Dong-Qin Dai https://orcid.org/0000-0001-8935-8807

Li-Juan Zhang https://orcid.org/0000-0002-3234-6757

Wan-Tong Zhao https://orcid.org/0009-0006-7298-0150

Ekachai Chukeatirote https://orcid.org/0000-0002-9968-5841

Saowaluck Tibpromma https://orcid.org/0000-0002-4706-6547

Data availability

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

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