Abstract
The genus Diaporthe comprises a functionally diverse group of fungi, inhabiting plant tissues as pathogens, endophytes, and saprobes. Many species cause significant diseases in economically and ecologically important woody plants. Despite extensive research on the taxonomy and phylogeny of Diaporthe over the past two decades, species delineation remains challenging due to the high morphological plasticity among the more than 1,000 described taxa. In practice, molecular phylogenetic divergence has become the primary criterion for species delimitation, as morphological characters often fail to resolve species boundaries. In this study, we employed an integrative taxonomic approach combining multi-locus phylogeny with morphological characterization to investigate Diaporthe isolates associated with leaf spot diseases on Fagaceae, a dominant tree family in Chinese forests. Our investigation resolved eight distinct lineages, including five novel species described herein: namely D. castanopsidicola, D. changjiangensis, D. cyclobalanopsidicola, D. liangxii, and D. lithocarpicola from Guangdong and Hainan Provinces, China. These findings significantly contribute to resolving the cryptic diversity of Diaporthe and expand our understanding of the fungal communities driving foliar diseases in Fagaceae.
Key words: Ascomycota , cryptic species, foliar pathogens, multi-locus phylogeny, Sordariomycetes , Southern China, taxonomy
Introduction
The genus Diaporthe (syn. Phomopsis), a hyper-diverse lineage within the Diaporthaceae (Diaporthales), represents one of the most ecologically successful groups of Ascomycota (Voglmayr et al. 2012; Dissanayake et al. 2017; Fan et al. 2020; Udayanga et al. 2021; Jiang et al. 2023, 2024; Zhu et al. 2023). Members of this genus are characterized by typical ordinal features, particularly in the teleomorph, such as immersed ascomata with elongated perithecial necks, and unitunicate, clavate to cylindrical asci containing hyaline, fusoid to cylindrical ascospores (Dissanayake et al. 2017; Yang et al. 2018; Jiang et al. 2021). In the anamorphic state, Diaporthe is identified by ostiolate conidiomata and cylindrical phialides producing three distinct types of hyaline, aseptate conidia (alpha, beta, and gamma) (Dissanayake et al. 2017; Jiang et al. 2021; Liu et al. 2024). While these morphological traits occur in several related taxa, Diaporthe forms a robust, monophyletic clade clearly distinguishable through molecular phylogenetic analyses (Voglmayr et al. 2017; Fan et al. 2018; Senanayake et al. 2018; Yang et al. 2020; Cai et al. 2024; Jiang et al. 2025a, 2025b).
Ecologically, Diaporthe species occupy a complex niche continuum, functioning as endophytes, saprobes, or opportunistic pathogens (Gomes et al. 2013; Huang et al. 2015; Gao et al. 2016; Cao et al. 2022). Many species are notorious pathogens of economically important woody plants, causing devastating diseases such as leaf spots, fruit rots, branch cankers, and dieback globally (Gomzhina and Gannibal 2022; Bai et al. 2023; Wan et al. 2023). Notable examples include D. alnicola and D. amygdali on Alnus in China (Li et al. 2025), and the D. ambigua complex associated with kiwifruit rot in Chile (Díaz et al. 2017). Similarly, D. foeniculina and allied species have been implicated in severe shoot blights in crops ranging from citrus to blueberry across Europe (Vakalounakis et al. 2019; Hilário et al. 2020).
Given its economic and ecological significance, the taxonomy of Diaporthe has been a focal point of mycology since the genus was established (Nitschke 1870). Historically, species identification relied heavily on host association and morphology (Mostert et al. 2001; Santos and Phillips 2009). However, modern investigations have revealed that high morphological plasticity and broad host ranges render traditional identification unreliable (Jiang et al. 2021; Zhu et al. 2024). Consequently, the field now employs integrative taxonomy. Recent studies employing genealogical concordance phylogenetic species recognition (GCPSR) and coalescence-based models, such as the General Mixed Yule-Coalescent (GMYC) and Poisson Tree Processes (PTP), have been instrumental in resolving cryptic species complexes and refining the generic boundaries (Hilário et al. 2020, 2021a, 2021b; Dissanayake et al. 2024; Pereira and Phillips 2024). These advanced approaches resulted in both the description of numerous novel lineages and the necessary synonymization of previously over-described taxa (Ren et al. 2025).
Despite these advances, knowledge of Diaporthe diversity remains uneven across host families. Fagaceae, comprising dominant tree species (e.g., Castanopsis, Lithocarpus, Quercus) in the subtropical evergreen broad-leaved forests of China, plays a critical role in forest ecosystem structure and carbon cycling. While leaf spot diseases are frequently observed in these forests, the diversity and taxonomy of the associated Diaporthe species remain poorly understood. This study aims to bridge this gap by (1) conducting a systematic survey of Diaporthe associated with leaf spots on Fagaceae in China, and (2) applying an integrative taxonomic framework to rigorously delimit species boundaries and unveil cryptic diversity.
Materials and methods
Surveys and isolations
In 2019, surveys of leaf spot diseases affecting Fagaceae were conducted in Yangshan County, Qingyuan City, Guangdong Province, and Changjiang Li Autonomous County, Hainan Province, China. Symptomatic leaves were collected from nine host species: Castanopsis chunii, C. hystrix, Cyclobalanopsis austrocochinchinensis, Cy. blakei, Cy. glauca, Cy. kerrii, Lithocarpus amygdalifolius, L. levis, and L. naiadarum. Samples were placed in sealed plastic bags and transported to the laboratory. Fungal isolation was performed following surface sterilization. Leaves were rinsed under running tap water to remove surface debris, immersed in 75% ethanol for 3 min, rinsed with sterile distilled water for 2 min, and dried with sterile absorbent cotton. Leaf segments (approx. 5 × 5 mm) containing the margin between diseased and healthy tissue were excised using a sterile scalpel and plated onto potato dextrose agar (PDA; 200 g potatoes, 20 g dextrose, and 20 g agar per L). Plates were incubated at 25 °C in the dark until mycelial growth was observed. Hyphal tips were transferred to fresh PDA plates to obtain pure cultures. Holotype specimens were deposited in the Herbarium of the Chinese Academy of Forestry (CAF; http://museum.caf.ac.cn/), and ex-type living cultures were deposited in the China Forestry Culture Collection Center (CFCC; https://cfcc.caf.ac.cn/).
Morphological analyses
Cultural characteristics were assessed on PDA, malt peptone agar (MPA; 30 g malt extract, 5 g mycological peptone, and 15 g agar per L), and synthetic nutrient agar (SNA; containing 0.2 g glucose, 0.2 g sucrose, 1 g KH2PO4, 1 g KNO3, 0.25 g MgSO4, 0.5 g KCl, and 14 g agar per L). To induce sporulation, sterile pine needles were placed on the surface of SNA plates. Colony morphology and growth rates were recorded after 14–20 d incubation at 25 °C. Colony colors were described according to Rayner (1970). Micromorphological structures were examined using a Zeiss Discovery V8 stereomicroscope (Jena, Germany) for conidiomata and an Olympus BX51 compound microscope (Tokyo, Japan) for microscopic features. Structures (conidiophores, conidiogenous cells, and alpha, beta, and gamma conidia) were mounted in distilled water or lactic acid for observation and photography. For each isolate, the dimensions of 50 randomly selected conidia were measured.
Molecular analyses
Genomic DNA was extracted from fungal mycelium harvested from colonies grown on PDA for 20 d using the Wizard® Genomic DNA Purification Kit (Promega, Madison, WI, USA), following the manufacturer’s protocol. Five partial gene regions were amplified: the internal transcribed spacer (ITS) regions, calmodulin (cal), histone H3 (his3), translation elongation factor 1-alpha (tef1), and beta-tubulin (tub2). The primer pairs used were ITS1/ITS4 for ITS (White et al. 1990), CAL-228F/CAL-737R for cal (Carbone and Kohn 1999), CYLH3F/H3-1b for his3 (Glass and Donaldson 1995; Crous et al. 2004), EF1-728F/EF1-986R for tef1 (Carbone and Kohn 1999), and T1/Bt2b for tub2 (Glass and Donaldson 1995; O’Donnell and Cigelnik 1997). PCR amplification was performed in a 25 μL reaction volume with the following thermal cycling conditions: initial denaturation at 94 °C for 5 min; followed by 35 cycles of denaturation at 94 °C for 30 s, annealing for 50 s at temperatures specific to each locus (48 °C for ITS; 54 °C for cal, tef1, and tub2; 57 °C for his3), and extension at 72 °C for 1 min; and a final elongation at 72 °C for 7 min. PCR products were sequenced bidirectionally by Beijing Ruibo Xingke Biotechnology Co., Ltd. (Beijing, China). Consensus sequences were assembled using SeqMan v. 7.1.0 (DNASTAR Inc., Madison, WI, USA). New sequences generated in this study were deposited in GenBank (Table 1).
Table 1.
Species of Diaporthe and their Genbank accession numbers.
Note. Ex-type strains are marked with *, and NA means not available.
Sequence alignments for individual loci were generated using MAFFT v. 7 (Katoh and Standley 2013) and manually adjusted in MEGA v. 7.0.21 where necessary. Based on BLASTn searches of ITS sequences, isolates were assigned to their respective species complexes (D. arecae, D. gardeniae, D. siamensis, and D. virgiliae SCs) for further detailed analyses. Phylogenetic analyses were performed on concatenated five-locus datasets (ITS-cal-his3-tef1-tub2) using both Maximum Likelihood (ML) and Bayesian Inference (BI) approaches implemented in the One-click Fungal Phylogenetic Tool (OFPT) v. 1.9.0 (Zeng et al. 2023). For ML analysis, the best-fit substitution models were determined using ModelFinder (Kalyaanamoorthy et al. 2017). ML trees were inferred using IQ-TREE with ultrafast bootstrap approximation (1000 replicates) (Minh et al. 2020; Hoang et al. 2018). Bayesian Inference was performed using MrBayes v. 3.2.7 (Ronquist et al. 2012), with best-fit models selected by ModelFinder under the Akaike Information Criterion (AIC). Two parallel runs of four Markov chains (one cold, three heated) were executed for 10 million generations, sampling every 100 generations. The first 25% of sampled trees were discarded as burn-in, and the remaining trees were used to calculate posterior probabilities (PP). Convergence was confirmed when the average standard deviation of split frequencies fell below 0.01. Phylogenetic trees were visualized in FigTree v. 1.4.4 (Rambaut 2018).
To evaluate the genealogical exclusivity of the novel lineages, the Pairwise Homoplasy Index (PHI) test was conducted in SplitsTree4 v. 4.14.6 (Huson and Bryant 2006) to detect recombination events (Φw). A p-value > 0.05 indicates a lack of significant recombination. Additionally, species boundaries were assessed using two coalescence-based methods: the Poisson Tree Processes (PTP) model and the Multi-rate PTP (mPTP) model. Analyses were conducted on the respective web servers (http://species.h-its.org/ptp/ and http://mptp.h-its.org/) using the ML tree as input, with 1,000,000 MCMC generations, a thinning of 100, and a burn-in of 10% (Zhang et al. 2013; Kapli et al. 2017).
Results
Phylogeny
Phylogenetic analyses of the Diaporthe arecae species complex
The concatenated dataset of five loci (ITS, cal, his3, tef1, and tub2) for the Diaporthe arecae species complex comprised 25 strains, including D. eres (AR5193 and DLR12a) as the outgroup. The alignment consisted of 2,581 characters (including gaps) partitioned as follows: cal (1–485), his3 (486–949), ITS (950–1483), tef1 (1484–1807), and tub2 (1808–2581). The alignment contained 640 distinct site patterns, with 30.49% of characters consisting of gaps or missing data. The best-fit substitution models were estimated as TNe+R2 for ITS, TNe+G4 for cal, HKY+F+I for his3, and HKY+F+G4 for both tef1 and tub2. The Maximum Likelihood (ML) analysis yielded a best tree with a log-likelihood value of –8485.05. The estimated gamma shape parameter (α) was 0.262. Bayesian Inference (BI) resulted in a topology congruent with the ML tree; thus, only the ML tree is presented with statistical support values (ML-BS/BI-PP) annotated at the nodes (Fig. 1).
Figure 1.
Phylogram of Diaporthe arecae and related species resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci. Numbers above the branches indicate ML bootstrap values (left, MLBS ≥ 50%) and Bayesian posterior probabilities (right, BPP ≥ 0.90). Ex-type strains are marked with *, and isolates from this study are in bold blue. Colored vertical bars on the right indicate species delimitation results based on the 5-locus phylogeny (5-loci), Poisson Tree Processes (PTP), and Multi-rate PTP (mPTP). The scale bar represents the expected number of nucleotide substitutions per site. The tree is rooted with D. eres (AR5193 and DLR12a).
In the phylogenetic tree, the isolates obtained in this study were resolved into three distinct lineages. (1) Two strains (CFCC 54324 and CFCC 55901) formed a highly supported clade (ML-BS/BI-PP = 100/1), described herein as D. liangxii. This species formed a sister lineage to D. zhaoqingensis (95/0.99). (2) Two strains (CFCC 54462 and CFCC 55903) formed a distinct, fully supported clade (100/1), described herein as D. cyclobalanopsidicola. This species appeared as a sister lineage to the clade containing D. aseana and D. xishuangbanica. (3) Isolate CFCC 55899 clustered within the D. arecae clade with strong support (100/1).
Species delimitation analyses yielded slightly conflicting results. The PTP analysis (Fig. 1) recognized D. liangxii and D. cyclobalanopsidicola as independent evolutionary lineages. However, the multi-rate PTP (mPTP) analysis was more conservative, delimiting D. liangxii and D. zhaoqingensis as a single molecular entity, and similarly merging D. cyclobalanopsidicola with D. aseana and D. xishuangbanica. Despite the mPTP results, distinct morphological traits and significant genetic distances (in tef1 and tub2) support their status as distinct species.
Phylogenetic analyses of the Diaporthe gardeniae species complex
The concatenated dataset (ITS, cal, his3, tef1, and tub2) for the Diaporthe gardeniae species complex comprised 18 strains, with D. irregularis (CGMCC 3.20092 and GZCC 19-0344) serving as the outgroup. The alignment contained 2,113 characters (including gaps) partitioned as follows: cal (1–469), his3 (470–929), ITS (930–1387), tef1 (1388–1719), and tub2 (1720–2113). The alignment comprised 372 distinct site patterns, with 24.58% gaps or missing data. Model selection utilizing ModelFinder determined the best-fit substitution models as K2P+R2 for ITS, K2P+G4 for cal, TN+F+G4 for his3, TIM2e+G4 for tef1, and HKY+F+I for tub2. The ML analysis yielded a best tree with a log-likelihood value of –5761.24, and the estimated gamma shape parameter was 0.183. Bayesian inference produced a topology consistent with the ML tree (Fig. 2).
Figure 2.
Phylogram of Diaporthe gardeniae species complex resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci. Numbers above the branches indicate ML bootstrap values (left, MLBS ≥ 50%) and Bayesian posterior probabilities (right, BPP ≥ 0.90). Ex-type strains are marked with *, and isolates from this study are in bold blue. Colored vertical bars on the right indicate species delimitation results based on the 5-locus phylogeny (5-loci), Poisson Tree Processes (PTP), and Multi-rate PTP (mPTP). The scale bar represents the expected number of nucleotide substitutions per site. The tree is rooted with D. irregularis (CGMCC 3.20092 and GZCC 19-0344).
In the phylogenetic tree, the isolates from this study clustered into two groups: (1) Four isolates (CFCC 54306, CFCC 52791, CFCC 54296, and CFCC 54333) clustered within the D. apiculata clade with strong statistical support (BS/PP = 100/1). (2) Two isolates (CFCC 54297 and CFCC 55913) formed a distinct, fully supported clade (100/1), described herein as D. lithocarpicola. This species formed a sister lineage to D. sennicola (CFCC 51634 and CFCC 51635).
Both PTP and mPTP analyses lumped D. lithocarpicola and D. sennicola into a single evolutionary entity (Fig. 2). However, these two taxa formed well-supported, distinct clades in the multi-locus phylogeny (100% BS / 1.00 PP). Furthermore, a comparison of nucleotide sequences revealed significant genetic distinctions. For instance, D. lithocarpicola differs from D. sennicola by 13 base pairs (bp) in cal, 2 base pairs (bp) in his3, 14 base pairs (bp) in tef1 and 5 bp in tub2. Coupled with distinct morphological characteristics, we strictly defined D. lithocarpicola as a novel species following the GCPSR concept.
Phylogenetic analyses of the Diaporthe siamensis species complex
The concatenated dataset (ITS, cal, his3, tef1, and tub2) for the Diaporthe siamensis species complex comprised 31 strains, rooted with D. oxe (CBS 133186 and CBS 133187). The alignment consisted of 2,460 characters (including gaps) partitioned as follows: cal (1–452), his3 (453–902), ITS (903–1360), tef1 (1361–1690), and tub2 (1691–2460). The matrix contained 531 distinct site patterns, with 33.83% of characters consisting of gaps or missing data. Model selection identified the best-fit substitution models as K2P+R2 for ITS, K2P+G4 for cal, TN+F+I for his3, TIM2e+G4 for tef1, and HKY+F+G4 for tub2. The ML analysis resulted in a best tree with a log-likelihood value of –7544.83, and the estimated gamma shape parameter (α) was 0.229. Bayesian Inference (BI) recovered a topology congruent with the ML tree (Fig. 3).
Figure 3.
Phylogram of Diaporthe siamensis species complex resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci. Numbers above the branches indicate ML bootstrap values (left, MLBS ≥ 50%) and Bayesian posterior probabilities (right, BPP ≥ 0.90). Ex-type strains are marked with *, and isolates from this study are in bold blue. Colored vertical bars on the right indicate species delimitation results based on the 5-locus phylogeny (5-loci), Poisson Tree Processes (PTP), and Multi-rate PTP (mPTP). The scale bar represents the expected number of nucleotide substitutions per site. The tree is rooted with D. oxe (CBS 133186 and CBS 133187).
In the phylogenetic tree, the isolates obtained in this study were assigned to two lineages: (1) Three isolates (CFCC 54817, CFCC 55916, and CFCC 54295) clustered with the ex-type strain of D. siamensis (MFLUCC 10-0573a) with maximum support (BS/PP = 100/1). (2) Two isolates (CFCC 54311 and CFCC 55917) formed a distinct, highly-supported clade (100/1), described herein as D. changjiangensis. This species resides in a sister relationship with D. yunnanensis (CGMCC 3.18289 and LC8107), separated by high statistical support (94/0.99).
Similar to the D. gardeniae complex, species delimitation analyses for this group yielded results incongruent with the phylogeny. Both PTP and mPTP analyses suggested that D. changjiangensis and D. yunnanensis constitute a single species (Fig. 3). However, D. changjiangensis differs from D. yunnanensis by 23 bp in ITS, 9 bp in his3, and 3 bp in tef1. Based on these genetic distinctions and morphological differences, we recognize D. changjiangensis as a novel species.
Phylogenetic analyses of the Diaporthe virgiliae species complex
The concatenated dataset (ITS, cal, his3, tef1, and tub2) for the Diaporthe virgiliae species complex comprised 15 strains, rooted with D. shennongjiaensis (CNUCC 201905 and CNUCC 201906). The alignment consisted of 2,585 characters (including gaps) partitioned as follows: cal (1–413), his3 (414–879), ITS (880–1472), tef1 (1473–1798), and tub2 (1799–2585). The matrix contained 348 distinct site patterns, with 19.27% of characters comprising gaps or missing data. Model selection identified the best-fit substitution models as K2P+I for ITS, K2P for cal, TN+F+I for his3, HKY+F+R2 for tef1, and HKY+F+I for tub2. The ML analysis resulted in a best tree with a log-likelihood value of –5750.79, and the estimated gamma shape parameter (α) was 0.02. Bayesian Inference (BI) recovered a topology congruent with the ML tree (Fig. 4).
Figure 4.
Phylogram of Diaporthe virgiliae species complex resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci. Numbers above the branches indicate ML bootstrap values (left, MLBS ≥ 50%) and Bayesian posterior probabilities (right, BPP ≥ 0.90). Ex-type strains are marked with *, and isolates from this study are in bold blue. The scale bar represents the expected number of nucleotide substitutions per site. Colored vertical bars on the right indicate species delimitation results based on the 5-locus phylogeny (5-loci), Poisson Tree Processes (PTP), and Multi-rate PTP (mPTP). The tree is rooted with D. shennongjiaensis (CNUCC 201905 and CNUCC 201906).
In the phylogenetic tree, the two isolates obtained in this study (CFCC 54374 and CFCC 55908) formed a highly supported, distinct clade (BS/PP = 100/1), described herein as D. castanopsidicola. Phylogenetically, D. castanopsidicola is sister to D. linzhiensis (CFCC 71057 and N266C).
Species delimitation analyses yielded remarkably distinct results. The PTP analysis successfully delimited D. castanopsidicola as an independent evolutionary lineage separate from D. linzhiensis (Fig. 4). In contrast, the mPTP analysis produced a highly conservative delimitation, lumping D. castanopsidicola into a large species aggregate that includes D. linzhiensis, D. virgiliae, D. alnicola, and D. heterophyllae. Given that D. virgiliae and D. heterophyllae are well-established biological species, the lumping result of mPTP is considered an artifact of the method’s conservatism in this dataset. Therefore, based on the robust phylogenetic separation (100% BS) and morphological distinctions, we establish D. castanopsidicola as a novel species.
Evaluation of recombination (PHI analysis)
To further validate the genealogical exclusivity of the novel lineages, the Pairwise Homoplasy Index (PHI) test was performed to detect potential recombination events between the new species and their phylogenetically closest relatives. Five specific datasets were constructed for assessment: (A) D. castanopsidicola vs. D. linzhiensis; (B) D. changjiangensis vs. D. yunnanensis; (C) D. lithocarpicola vs. D. sennicola; (D) D. cyclobalanopsidicola vs. D. aseana and D. xishuangbanica; and (E) D. liangxii vs. D. zhaoqingensis and D. arecae. The PHI test results revealed no statistically significant evidence of genetic recombination (p > 0.05) within any of the tested groups (Fig. 5). Specifically, comparisons for Clades A, B, and C yielded a p value of 1.0, indicating a lack of recombination signal. Similarly, Clades D (p = 0.447) and E (p = 0.958) showed no significant recombination. These findings confirm the absence of gene flow between the novel species and their sister taxa, reinforcing their validity as distinct evolutionary lineages.
Figure 5.
The split graphs of a PHI test result of Diaporthe species using the LogDet transformation and split decomposition options based on the combined ITS, cal, his3, tef1 and tub2 gene loci. Ap = 1.0. Bp = 1.0. Cp = 1.0. Dp = 0.4474. Ep = 0.9579.
Taxonomy
Diaporthe apiculata
Y.H. Gao, F. Liu & L. Cai, Syst. Biodivers. 14: 106 (2016)
0E95B2B8-1934-5002-8614-6D096E8F9295
Figure 6.
Morphology of Diaporthe apiculata (CFCC 52791). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C, D Conidiogenous cells giving rise to conidia. E–G Alpha conidia. Scale bars: 200 μm (B); 10 μm (C–G).
Description.
See Gao et al. (2016).
Distribution.
China.
Ecology.
Associated with leaf spot disease of Castanopsis chunii, C. hystrix, Lithocarpus amygdalifolius and L. naiadarum, branch canker disease of Rhus chinensis, and endophytic in Camellia sinensis.
Materials examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus naiadarum, 24 November 2019, Yong Li (culture CFCC 52791); • Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Castanopsis chunii, 26 November 2019, Dan-ran Bian (culture CFCC 54306); • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus amygdalifolius, 31 March 2019, Yong Li (culture CFCC 54333); • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Castanopsis hystrix, 31 March 2019, Yong Li (culture CFCC 54296).
Notes.
Diaporthe apiculata was originally described as an endophyte from Camellia sinensis in China (Gao et al. 2016) and subsequently reported causing branch cankers on Rhus chinensis (Yang et al. 2021a). In the present study, four isolates were recovered from leaf spots of Castanopsis chunii, C. hystrix, Lithocarpus amygdalifolius, and L. naiadarum. Phylogenetically, these isolates formed a highly supported clade (ML/BI = 100/1) with the ex-type strain of D. apiculata (CGMCC 3.17533) (Fig. 2). To our knowledge, this is the first report of D. apiculata associated with Castanopsis and Lithocarpus species, expanding the known host range of this fungus to Fagaceae.
Diaporthe arecae
(H.C. Srivast., Zakia & Govindar.) R.R. Gomes, Glienke & Crous, Persoonia 31: 16 (2013)
BAD6FC2C-9D00-509C-874D-3E9E01EB9438
Figure 7.
Morphology of Diaporthe arecae (CFCC 55899). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C Conidiogenous cells. D–G Alpha and beta conidia. Scale bars: 400 μm (B); 10 μm (C–G).
Description.
Conidiomata pycnidial, aggregated or solitary, erumpent, subglobose, black, 300–1200 μm diam., exuding white conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, attenuate towards the apex, phialidic, 10.5–28.5 × 1.5–3.5 μm. Alpha conidia aseptate, hyaline, smooth, multi-guttulate, fusoid, straight or slightly curved, base truncate, (5.5–)6.5–8.5(–9.5) × 2–2.5(–3) μm (n = 50), L/W = 2.2–4.6. Beta conidia aseptate, hyaline, smooth, guttulate or not, filiform, curved or not, (12–)13.5–19.5(–24.5) × 1.5–2 μm (n = 50), L/W = 5.7–15.5.
Culture characteristics.
Colonies on PDA flat, spreading, flocculent, with entire edge, white to mouse grey, reaching 25 mm in diam. after 10 days at 25 °C; colonies on MPA flat, spreading, flocculent, with entire edge, white, reaching 25 mm in diam. after 10 days at 25 °C, forming black conidiomata with off-white masses; colonies on SNA flat, with sparse aerial mycelium and entire margin, white, reaching 25 mm in diam. after 20 days at 25 °C.
Distribution.
Worldwide.
Ecology.
Associated with leaf spot, branch canker and fruit rot diseases on various hosts, including Citrus sp., Cyclobalanopsis glauca, Musa sp., Pterocarpus indicus, Senna bicapsularis, Subramanella arecae, etc.
Materials examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis glauca, 30 March 2019, Yong Li (culture CFCC 55899).
Notes.
Diaporthe arecae was recently circumscribed based on a combination of phylogenetic analysis, GCPSR, and coalescence-based models (Dissanayake et al. 2024). In the current study, isolate CFCC 55899 clustered with the ex-type strain of D. arecae (CBS 161.64) in the multi-locus phylogeny (Fig. 1). To our knowledge, this is the first report of D. arecae causing leaf spots on Cyclobalanopsis glauca in China.
Diaporthe castanopsidicola
Ning Jiang sp. nov.
D6502957-32AA-59F3-9B9B-41868CFB6B64
841314
Figure 8.
Morphology of Diaporthe castanopsidicola (CFCC 54374). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C Conidiogenous cells giving rise to conidia. D–G Alpha conidia. Scale bars: 400 μm (B); 10 μm (C–G).
Etymology.
Referring to the host genus, Castanopsis, from which the holotype was isolated.
Diagnosis.
Distinct from its phylogenetically related species D. linzhiensis by larger conidiomata.
Typus.
CHINA • Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Castanopsis chunii, 26 November 2019, Dan-ran Bian (holotype CAF800027; ex-holotype culture CFCC 54374).
Description.
Conidiomata pycnidial, aggregated, erumpent, pulvinate, subglobose, umber, 800–2000 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to filiform, phialidic, 19.5–41.5 × 1.5–3.5 μm. Alpha conidia aseptate, hyaline, smooth, biguttulate, cylindrical, slightly constricted at the middle, straight, base truncate, (4.5–)5–6(–6.5) × (2–)2.5–3 μm (n = 50), L/W = 1.6–2.6.
Culture characteristics.
Colonies on PDA flat, spreading, flocculent, with entire edge, initially white, becoming mouse grey with age, reaching 25 mm in diam. after 10 days at 25 °C; colonies on MPA flat, spreading, flocculent, with entire edge, white to grey, reaching 25 mm in diam. after 10 days at 25 °C, forming umber conidiomata with cream conidial masses; colonies on SNA flat, spreading, with sparse aerial mycelium and undulate margin, white, reaching 25 mm in diam. after 20 days at 25 °C.
Additional material examined.
CHINA • Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Castanopsis chunii, 28 November 2019, Dan-ran Bian (culture CFCC 55908).
Distribution.
China, Guangdong Province.
Ecology.
Associated with leaf spot disease of Castanopsis chunii.
Notes.
Phylogenetically, Diaporthe castanopsidicola forms a highly supported sister clade to D. linzhiensis (Fig. 4). Although the mPTP analysis conservatively lumped these two taxa (along with D. virgiliae, D. alnicola, and D. heterophyllae), the PHI test (p = 1.0) confirmed no significant recombination between them (Fig. 5A), supporting their divergence. Morphologically, D. castanopsidicola can be readily distinguished from D. linzhiensis by its larger conidiomata (800–2000 μm in D. castanopsidicola vs. 300–500 μm in D. linzhiensis) (Li et al. 2025). Furthermore, comparisons of nucleotide sequences revealed significant differences between the two species: ITS (6/460 bp, 1.3%), cal (4/417 bp, 1.0%), his3 (9/428 bp, 2.1%), tef1 (4/335 bp, 1.2%), and tub2 (6/428 bp, 1.4%). Based on the distinct morphology, significant genetic distance, and genealogical exclusivity, we introduce D. castanopsidicola as a new species.
Diaporthe changjiangensis
Ning Jiang sp. nov.
F6951229-C2A9-5918-9C45-DA431CA5A6C0
848340
Figure 9.
Morphology of Diaporthe changjiangensis (CFCC 54311). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C, D Conidiogenous cells. D–G Alpha conidia. Scale bars: 200 μm (B); 10 μm (C–G).
Etymology.
Referring to Changjiang County, Hainan Province, where the type specimen was collected.
Diagnosis.
Distinct from its phylogenetically related species D. yunnanensis by wider alpha conidia.
Typus.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis kerrii, 30 March 2019, Yong Li (holotype CAF800077; ex-holotype culture CFCC 54311).
Description.
Conidiomata pycnidial, aggregated, erumpent, pulvinate, subglobose, brown, 150–450 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical, phialidic, 9–46 × 2.5–5.5 μm. Alpha conidia aseptate, hyaline, smooth, biguttulate, cylindrical, straight, base truncate, (5–)5.5–6.5(–7.5) × 2.5–3 μm (n = 50), L/W = 1.8–2.8.
Culture characteristics.
Colonies on PDA flat, spreading, flocculent, with entire edge, white, reaching 25 mm in diam. after 10 days at 25 °C, forming brown conidiomata with luteous conidial masses; colonies on MPA flat, spreading, flocculent, with entire edge, white, forming dark brown irregular zone, reaching 25 mm in diam. after 10 days at 25 °C, forming brown conidiomata with luteous conidial masses; colonies on SNA flat, spreading, with sparse aerial mycelium and undulate margin, white, reaching 25 mm in diam. after 15 days at 25 °C, forming brown conidiomata with luteous conidial masses on the pine needles.
Additional material examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis kerrii, 30 March 2019, Yong Li (culture CFCC 55917).
Distribution.
China, Hainan Province.
Ecology.
Associated with leaf spot disease of Cyclobalanopsis kerrii.
Notes.
In the phylogenetic analysis, Diaporthe changjiangensis forms a sister lineage to D. yunnanensis (Fig. 3). Although the PTP and mPTP analyses produced ambiguous species boundaries for this pair (Fig. 3), the PHI test (p = 1.0) strongly supported the lack of recombination between them (Fig. 5B). Morphologically, D. changjiangensis differs from D. yunnanensis (described from Coffea sp.) by having distinctly wider alpha conidia (2.5–3 μm vs. 1–2.5 μm) (Gao et al. 2017). Most notably, comparison of nucleotide sequences revealed remarkable genetic divergence between the two species, particularly in the ITS region (23/462 bp, 5.0%), as well as in his3 (9/453 bp, 2.0%) and tef1 (3/338 bp, 0.9%). Such a high degree of sequence variation in the ITS region provides evidence for the establishment of D. changjiangensis as a novel species.
Diaporthe cyclobalanopsidicola
Ning Jiang sp. nov.
8AC20099-3ED5-5258-923B-B147C3CD1249
841316
Figure 10.
Morphology of Diaporthe cyclobalanopsidicola (CFCC 54462). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C Conidiogenous cells giving rise to conidia. D–G Alpha and beta conidia. Scale bars: 400 μm (B); 10 μm (C–G).
Etymology.
Named after the host genus Cyclobalanopsis and “-cola” = “inhabiting”.
Diagnosis.
Distinct from D. aseana by longer alpha conidia and from D. xishuangbanica by larger conidiomata.
Typus.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis blakei, 30 March 2019, Yong Li (holotype CAF800029; ex-type culture CFCC 54462).
Description.
Conidiomata pycnidial, aggregated or solitary, erumpent, subglobose, brown, 500–1300 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical, phialidic, 11.5–57 × 1.5–2.5 μm. Alpha conidia aseptate, hyaline, smooth, guttulate, fusoid, straight or slightly curved, base truncate, (8–)8.5–12.5(–15) × (2–)2.5–3(–3.5) μm (n = 50), L/W = 2.5–5. Beta conidia aseptate, hyaline, smooth, guttulate, filiform, straight or curved, (17.5–)20.5–29.5(–32) × 1.5–2 μm (n = 50), L/W = 10.3–19.9.
Culture characters.
Colonies on PDA flat, spreading, flocculent, with entire edge, initially white, becoming olivaceous grey with age, reaching 25 mm in diam. after 7 days at 25 °C; colonies on MPA flat, spreading, flocculent, with entire edge, white to pale luteous, reaching 25 mm in diam. after 7 days at 25 °C, forming brown conidiomata with amber conidial masses; colonies on SNA flat, feathery, with sparse aerial mycelium and undulate margin, white.
Additional material examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis blakei, 28 March 2019, Yong Li (culture CFCC 55903).
Distribution.
China, Hainan Province.
Ecology.
Associated with leaf spot disease of Cyclobalanopsis blakei.
Notes.
Phylogenetically, Diaporthe cyclobalanopsidicola resides in a subclade with D. aseana and D. xishuangbanica (Fig. 1). Although the mPTP analysis conservatively recognized this group as a single species, the PHI test (p = 0.447) detected no significant recombination (Fig. 5D), supporting their status as independent lineages. Morphologically, D. cyclobalanopsidicola is easily distinguished from D. aseana by its longer alpha conidia (8.5–12.5 × 2.5–3 μm in D. cyclobalanopsidicola vs. 6–9 × 2–3 μm in D. aseana) and from D. xishuangbanica by markedly larger conidiomata (500–1300 μm in D. cyclobalanopsidicola vs. 180–310 μm in D. xishuangbanica) (Hyde et al. 2016; Gao et al. 2017). Moreover, pairwise sequence alignment revealed substantial nucleotide divergence. Diaporthe cyclobalanopsidicola differs from D. aseana by 17 bp in ITS, 24 bp in cal, 19 bp in his3, 13 bp in tef1, and 38 bp in tub2. Similarly, it differs from D. xishuangbanica by 11 bp in ITS, 31 bp in his3, 31 bp in tef1, and 33 bp in tub2. These significant genetic distances provide robust evidence to justify D. cyclobalanopsidicola as a distinct species.
Diaporthe liangxii
Ning Jiang sp. nov.
0AEB6BF0-E93C-503D-8615-B8219FD4470C
841318
Figure 11.
Morphology of Diaporthe liangxii (CFCC 54324). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidioma. C Conidiogenous cells giving rise to conidia. D–G Beta conidia. Scale bars: B = 200 μm; C–G = 10 μm.
Etymology.
In honor of Prof. Dr. Liang Xi, in recognition of his distinguished contributions to forestry research in China.
Diagnosis.
Distinct from its phylogenetically related species of D. zhaoqingensis by host plant.
Typus.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus naiadarum, 30 March 2019, Yong Li (holotype CAF800078; ex-holotype culture CFCC 54324).
Description.
Conidiomata pycnidial, aggregated, erumpent, pulvinate, subglobose, dark brown, 350–1000 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, filiform, phialidic, 9.5–22.5 × 2–3.5 μm. Beta conidia aseptate, hyaline, smooth, guttulate or not, filiform, straight or hamate, base truncate, (13.5–)20.5–30(–31) × (1–)1.5–2 μm (n = 50), L/W = 14.1–23.2.
Culture characters.
Colonies on PDA flat, spreading, flocculent, with entire edge, white to buff, reaching 25 mm in diam. after 10 days at 25 °C; colonies on MPA flat, spreading, flocculent, with undulate edge, initially white, forming grey zones in the middle and producing dark brown conidiomata with cream conidial masses, reaching 25 mm in diam. after 10 days at 25 °C; colonies on SNA flat, spreading, with sparse aerial mycelium and undulate margin, white, forming concentric zones with age, reaching 25 mm in diam. after 15 days at 25 °C.
Additional material examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus naiadarum, 30 March 2019, Yong Li (culture CFCC 55901).
Distribution.
China, Hainan Province.
Ecology.
Associated with leaf spot disease of Lithocarpus naiadarum.
Notes.
Phylogenetically, Diaporthe liangxii clusters with D. zhaoqingensis (Fig. 1). Due to their close relationship, the mPTP analysis lumped them into a single entity; however, the PHI test (p = 0.958) revealed no significant recombination (Fig. 5E), supporting their separation as distinct lineages. Morphologically, D. liangxii (producing only beta conidia in this study) shares similar conidial dimensions with D. zhaoqingensis (Luo et al. 2022). Nevertheless, they can be distinguished by host association (Lithocarpus naiadarum vs. Morinda officinalis). More importantly, the two species exhibit remarkable genetic divergence. Diaporthe liangxii differs from D. zhaoqingensis by 12 bp in ITS, 45 bp in cal, 3 bp in his3, and 34 bp in tub2. The substantial sequence variations in protein-coding genes (cal and tub2) strongly justify the recognition of D. liangxii as a novel species despite their morphological resemblance.
Diaporthe lithocarpicola
Ning Jiang sp. nov.
CB5B039E-33E5-5113-BB6C-C14828CC4B58
841319
Figure 12.
Morphology of Diaporthe lithocarpicola (CFCC 54297). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C Conidiogenous cells giving rise to conidia. D–G Beta conidia. Scale bars: 400 μm (B); 10 μm (C–G).
Etymology.
Named after the host genus Lithocarpus and “-cola” = “inhabiting”.
Diagnosis.
Distinct from its phylogenetically related species D. sennicola by wider conidiogenous cells.
Typus.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus naiadarum, 28 March 2019, Yong Li (holotype CAF800079; ex-holotype culture CFCC 54297).
Description.
Conidiomata pycnidial, aggregated or solitary, erumpent, subglobose, amber, 400–950 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, attenuate towards the apex, phialidic, 4.5–13.5 × 1.5–2.5 μm. Beta conidia aseptate, hyaline, smooth, guttulate or not, filiform, curved, (24.5–)28.5–38(–43.5) × 1.5–2 μm (n = 50), L/W = 15.4–27.9.
Culture characters.
Colonies on PDA flat, spreading, flocculent, with entire edge, initially white, becoming olivaceous grey with age, reaching 25 mm in diam. after 10 days at 25 °C; colonies on MPA flat, spreading, flocculent, with entire edge, white to amber, reaching 25 mm in diam. after 10 days at 25 °C, forming amber conidiomata with brown conidial masses; colonies on SNA flat, with sparse aerial mycelium and undulate margin, white, reaching 25 mm in diam. after 20 days at 25 °C.
Additional material examined.
CHINA • Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus naiadarum, 30 March 2019, Yong Li (culture CFCC 55913).
Ecology.
Associated with leaf spot disease of Lithocarpus naiadarum.
Notes.
Phylogenetically, Diaporthe lithocarpicola is closely related to D. sennicola (Fig. 2). Both PTP and mPTP analyses produced ambiguous species boundaries between D. lithocarpicola and D. sennicola. However, the PHI test (p = 1.0) provided strong evidence for the lack of recent recombination (Fig. 5C), supporting their genealogical independence. Morphologically, D. lithocarpicola differs significantly from D. sennicola (on Senna bicapsularis) by having much wider conidiogenous cells (1.5–2.5 μm vs. 0.9–1.4 μm) (Yang et al. 2017). Genetic comparison further justifies their separation, despite high similarity in the ITS region (1/462 bp difference). Diaporthe lithocarpicola exhibits distinct variations in protein-coding genes, differing from D. sennicola by 13 bp in cal, 14 bp in tef1, 5 bp in tub2, and 2 bp in his3.
Diaporthe siamensis
Udayanga, Xing Z. Liu & K.D. Hyde, Cryptog. Mycol. 33(3): 298 (2012)
CAAC3F4F-2F58-5446-B5D1-5DFF3C6C1B66
Figure 13.
Morphology of Diaporthe siamensis (CFCC 54295). A Colonies on PDA, MPA and SNA after 20 d at 25 °C. B Conidiomata. C Conidiogenous cells. D–G Alpha and beta conidia. Scale bars: 300 μm (B); 10 μm (C–G).
Description.
Distribution.
China, Thailand.
Ecology.
Associated with leaf spot disease of Cyclobalanopsis kerrii, Dasymaschalon sp., and Lithocarpus levis, and fruit rot disease of Citrus sinensis.
Materials examined.
CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis kerrii, 30 March 2019, Yong Li (cultures CFCC 54817, CFCC 55916). Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus levis, 30 March 2019, Yong Li (culture CFCC 54295).
Notes.
Diaporthe siamensis was originally described from Dasymaschalon sp. (Annonaceae) in Thailand (Udayanga et al. 2012). In the current study, three isolates recovered from Cyclobalanopsis kerrii and Lithocarpus levis clustered with the ex-type strain of D. siamensis (MFLUCC 10-0573a) with maximum statistical support (ML/BI = 100/1) (Fig. 3). These findings constitute new host records for D. siamensis on Cyclobalanopsis and Lithocarpus species, broadening the known host range of this pathogen to the family Fagaceae.
Discussion
Diaporthe is widely recognized as one of the most speciose and ecologically diverse genera within the Ascomycota. While extensive surveys have been conducted on economic crops (e.g., Citrus, Helianthus, Vitis) and ornamental plants (Gomzhina and Gannibal 2022; Wan et al. 2023), the diversity of Diaporthe associated with forest ecosystems, particularly in subtropical regions, remains significantly underestimated. Fagaceae species (e.g., Castanopsis, Lithocarpus, Cyclobalanopsis) are dominant canopy trees in the evergreen broad-leaved forests of southern China. Our study revealed a remarkable diversity of Diaporthe species inhabiting foliar lesions of these trees. From a relatively localized survey, we identified eight distinct lineages of Diaporthe, five of which are described here as new to science. This high rate of novelty (62.5%) suggests that Fagaceae forests serve as a substantial reservoir for Diaporthe diversity. Our findings corroborate the hypothesis that the “hyper-diversity” of Diaporthe is largely driven by its association with diverse woody hosts in natural ecosystems, warranting broader sampling efforts in forestry pathology.
Accurate species delimitation in Diaporthe has long been impeded by morphological plasticity and the lack of informative barcodes. In recent years, coalescence-based methods (GMYC, PTP/mPTP) have become standard tools for establishing species boundaries (Hilário et al. 2020, 2021a, 2021b; Dissanayake et al. 2024; Pereira et al. 2023; Pereira and Phillips 2024). However, our study highlights the limitations of relying solely on automated delimitation algorithms. In the D. virgiliae species complex, the mPTP model conservatively lumped D. castanopsidicola with clearly distinct species such as D. virgiliae and D. alnicola (Fig. 4). Similar incongruences were observed in the D. arecae, D. gardeniae, and D. siamensis complexes. While mPTP is generally considered robust, it can be prone to “over-lumping” when analyzing datasets with uneven sampling or recent rapid radiations (Kapli et al. 2017). To resolve these conflicts, we adopted a rigorous integrative taxonomic approach. We prioritized GCPSR based on well-supported clades in multi-locus phylogeny (ML/BI), supported by pairwise homoplasy index (PHI) tests which confirmed the absence of recent recombination (p > 0.05). Furthermore, significant nucleotide divergence in protein-coding genes (particularly tub2 and tef1) and distinct morphological traits (e.g., conidiomata size in D. castanopsidicola vs. D. linzhiensis) provided irrefutable evidence for the validity of the five new species. This study serves as a case study demonstrating that algorithmic delimitation results should be interpreted with caution and verified against biological and genetic distance data.
Cryptic speciation is a prevalent phenomenon in Diaporthe (Yang et al. 2018; Zhu et al. 2024). Traditional identification based on host association or ITS sequences often fails to distinguish closely related taxa (Yang et al. 2018; Dissanayake et al. 2024; Zhu et al. 2024). For instance, in our study, D. lithocarpicola and D. sennicola share high similarity in the ITS region (only 1 bp difference), which might lead to misidentification if analyzing ITS alone. However, the inclusion of cal and tef1 revealed substantial divergence (13–14 bp), disentangling these cryptic lineages. Similarly, D. liangxii and D. zhaoqingensis are morphologically indistinguishable in their anamorphs but are genetically distinct in cal (45 bp) and tub2 (34 bp). Conversely, D. changjiangensis exhibited an unexpectedly high variation in ITS (23 bp) compared to its sister species D. yunnanensis. These findings reinforce the necessity of using a five-locus concatened dataset (ITS, cal, his3, tef1, tub2) for reliable species resolution in Diaporthe, as proposed by recent taxonomic frameworks (Dissanayake et al. 2024).
The host range of Diaporthe species varies from strictly host-specific to broadly generalist (Mostert et al. 2001; Santos and Phillips 2009; Dissanayake et al. 2024). In this study, the five novel species (D. castanopsidicola, D. changjiangensis, D. cyclobalanopsidicola, D. liangxii, and D. lithocarpicola) were isolated exclusively from specific Fagaceae hosts (Castanopsis, Cyclobalanopsis, and Lithocarpus), suggesting potential host preference. However, given the limited sampling, it is premature to conclude strict host specificity. In contrast, we also recovered D. apiculata and D. siamensis from Fagaceae leaf spots. Diaporthe apiculata has been previously reported on Camellia (Theaceae) and Rhus (Anacardiaceae), and D. siamensis on Citrus (Rutaceae) and Dasymaschalon (Annonaceae). The identification of these known species on Fagaceae represent new host records and suggest they are generalist pathogens capable of cross-infection between forest trees and economic crops. This “host-jumping” potential poses a challenge for disease management in mixed-plantation forestry and highlights the need for further pathogenicity tests to evaluate their virulence across different host families.
Supplementary Material
Citation
Zhong X, Voglmayr H, Xue H, Li Y, Jiang N (2026) Unveiling cryptic diversity of Diaporthe associated with leaf spots of Fagaceae in China using an integrative taxonomic approach. IMA Fungus 17: e186438. https://doi.org/10.3897/imafungus.17.186438
Funding Statement
This study was supported by Fundamental Research Funds of CAF CAFYBB2021GA001 and CAFYBB2023PA002, and the National Natural Science Foundation of China (32501665).
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Artificial Intelligence (AI) use
The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.
No AI tools were used in the preparation of this manuscript.
Adherence to national and international regulations
All the fungal strains used in this study have been legally obtained, respecting the Convention on Biological Diversity (Rio Convention).
Funding
This study was supported by Fundamental Research Funds of CAF CAFYBB2021GA001 and CAFYBB2023PA002, and the National Natural Science Foundation of China (32501665).
Author contributions
Conceptualization (NJ, HV, HX and YL), Methodology (XZ, NJ, HV and HX), Software (NJ), Validation (NJ), Formal analysis (XZ and NJ), Investigation (NJ and YL), Resources (NJ, HX and YL), Data Curation (NJ, HX and YL), Writing - Original draft (NJ and XZ), Writing - Review and Editing (NJ, HV, HX and YL), Visualization (NJ), Supervision (YL), Project administration (NJ, HX and YL), Funding Acquisition (NJ, HX and YL).
Author ORCIDs
X. Zhong https://orcid.org/0009-0005-6817-2006
H. Voglmayr https://orcid.org/0000-0001-7666-993X
H. Xue https://orcid.org/0000-0003-0414-6237
Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary materials
Phylogram of Diaporthe resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
Single-gene phylogenies
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
docx
Alignments of arecae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of gardeniae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of siamensis species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of virgiliae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Phylogram of Diaporthe resulting from a maximum likelihood analysis based on the ITS, cal, his3, tef1 and tub2 gene loci
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
Single-gene phylogenies
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
docx
Alignments of arecae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of gardeniae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of siamensis species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Alignments of virgiliae species complex
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Ning Jiang
Data type
fas
Data Availability Statement
All of the data that support the findings of this study are available in the main text or Supplementary Information.