Abstract
The family Apiosporaceae is a cosmopolitan family of fungi that occurs as endophytes, pathogens, and saprobes. Due to the morphological similarity of species within this family, both morphological and phylogenetic analyses are required to determine their taxonomic status. In this study, eleven specimens were collected from Yunnan and Guizhou provinces in China. Morphological and phylogenetic analyses based on ITS, LSU, tef1-α, and tub2 sequence data were performed to identify Apiospora (A.) and Nigrospora (N.) isolates. Five new species are described: A. huaxiensissp. nov., A. nanmingensissp. nov., A. qingzhenensissp. nov., A. tongrenensissp. nov., and Nigrospora neosaccharicolasp. nov. Six new host record species are reported: A. locuta-pollinis, A. setariae, N. chinensis, N. endophytica, N. osmanthi, and N. sphaerica. Furthermore, A. mediterraneasyn. nov. is synonymized under A. hispanica, and A. euphorbiaesyn. nov., A. magnisporasyn. nov., and A. malaysianasyn. nov. are reduced to synonyms of A. vietnamensis based on morphological comparison and phylogenetic analysis. In addition, Apiospora sinensis is reinstated, with Arthrinium sinense (Ar. sinense) syn. nov. treated as its synonym. Statistical analysis of host preferences indicates that Poaceae is the dominant host family. Analysis of their regional distributions showed that Apiospora is more prevalent in Europe, while Nigrospora is more widespread in Asia. This study refines the taxonomy and expands the phylogenetic understanding of Apiosporaceae in China.
Key words: Arthrinium-like, Ascomycota , multilocus analysis, phylogeny, Sordariomycetes , taxonomy
Introduction
The family Apiosporaceae was established by Hyde et al. (1998) to accommodate Apiospora Sacc., together with Appendicospora K.D. Hyde, Dictyoarthrinium S. Hughes, Endocalyx Berk. & Broome, Scyphospora L.A. Kantsch., and Spegazzinia Sacc., based on distinctive morphological characteristics, particularly the characters of multilocular ascomata and Arthrinium-like basauxic conidiogenous cell (Senanayake et al. 2015; Li et al. 2023; Liu et al. 2024b). Phylogenetic analyses of LSU and SSU sequence data subsequently placed the family within Xylariales (Smith 2003), a position later confirmed by Crous and Groenewald (2013), who demonstrated that Apiosporaceae and Amphisphaeriaceae form sister clades. Wijayawardene et al. (2020) subsequently reassigned Apiosporaceae to Amphisphaeriales (Ascomycota, Sordariomycetes) (Hyde et al. 2024b), where it is currently classified. Over time, molecular data have refined the generic boundaries within the family. Tanaka et al. (2015) transferred Spegazzinia to Didymosphaeriaceae based on multigene phylogenetic evidence, while Wang et al. (2017) included Nigrospora Zimm. within Apiosporaceae. Samarakoon et al. (2020) further transferred Dictyoarthrinium to Didymosphaeriaceae based on its close relationship with Spegazzinia in both morphology and molecular phylogeny. Konta et al. (2021) placed Endocalyx in Cainiaceae, and Samarakoon et al. (2022) established Appendicosporaceae to accommodate Appendicospora as an independent lineage. As a result of these revisions, Apiosporaceae is now circumscribed to three well-established genera, Apiospora, Arthrinium Kunze, and Nigrospora (Wang et al. 2017; Pintos and Alvarado 2021; Hyde et al. 2024b; Mukhopadhyay et al. 2025). These genera are morphologically and phylogenetically related but differ in their diagnostic reproductive characteristics. Apiospora is the sexually typified genus, producing multilocular ascomata beneath stromata and exhibiting Arthrinium-like basauxic conidiogenesis, whereas Arthrinium was for a period considered the asexual morph linked to Apiospora (Kirk et al. 2008). The genus Apiospora, once treated as a synonym of Arthrinium, is now recognized as an independent lineage within Apiosporaceae based on multilocus phylogenetic evidence (Pintos and Alvarado 2021; Han et al. 2024; Liu et al. 2024b; Yan and Zhang 2024; Chang et al. 2025; Yu et al. 2025). For decades, both names were used interchangeably until multilocus phylogenetic analyses demonstrated that Apiosporasensu stricto and Arthriniumsensu stricto represent two distinct but closely related lineages within Apiosporaceae (Pintos and Alvarado 2021). Although both genera share morphological similarities, they differ in host range, distribution, and conidial morphology; Apiospora conidia are generally rounded in face view and lenticular in side view, while those of Arthrinium are more variable, including angular, curved, fusiform, globose, and polygonal forms (Zeng et al. 2022). Species of Apiospora have been found worldwide, ranging from tropical and subtropical regions to Mediterranean, temperate, and cold climates, while Arthrinium s.str. is quite rarely reported in tropical and subtropical habitats (Pintos and Alvarado 2021; Liu et al. 2023a). Additionally, Apiospora has a broader host range, whereas Arthrinium s.str. specimens are exclusively associated with plants from the Cyperaceae and Juncaceae (Pintos and Alvarado 2021). Nigrospora forms the third major lineage, phylogenetically allied to the Apiospora–Arthrinium clade, and is characterized by spherical to subspherical conidiogenous cells producing black, globose to subglobose conidia (Zhang et al. 2024a; Zou et al. 2024). Nigrospora is another important genus comprising endophytic, saprobic, and pathogenic species that can cause diseases in economically significant plants (Zou et al. 2024). Despite their morphological distinctness, extensive convergence in conidial size, shape, and development has historically obscured genus boundaries, underscoring the necessity of molecular approaches for accurate identification and phylogenetic resolution within the family (Wang et al. 2017; Pintos and Alvarado 2021; de Queiroz Brito et al. 2023).
Members of Apiosporaceae are ecologically versatile, functioning as endophytes, saprobes, and plant pathogens on numerous hosts worldwide (Liao et al. 2023; Liu et al. 2023a). Several species are of agricultural and ecological importance, including A. arundinis, associated with root rot of Pseudostellaria heterophylla (Xiao et al. 2024), and N. aurantiaca, causing leaf spots on Myrica rubra, Pandanus amaryllifolius, and Castanea mollissima (Luo et al. 2020; Khoo et al. 2022; Fu et al. 2025). Moreover, most species in this family can produce secondary metabolites, demonstrating strong biological activity (Xu et al. 2022; Overgaard et al. 2023). Their ecological versatility, morphological plasticity, and close evolutionary relationships make Apiosporaceae one of the most taxonomically complex and diverse families within Amphisphaeriales (Liao et al. 2023; Liu et al. 2023a).
The Yunnan-Guizhou Plateau harbors numerous potential Apiospora species due to its suitable climate, demonstrating strong ecological adaptability of this genus (Zeng et al. 2022; Zhang et al. 2023; Han et al. 2024; Guo et al. 2025; Yu et al. 2025). Despite significant taxonomic progress, the diversity and distribution of Apiosporaceae in China remain insufficiently explored, particularly in ecologically rich provinces such as Yunnan and Guizhou. To address this gap, diseased leaves and decaying stems were collected from Southwestern China, from which 35 strains were isolated. These isolates were identified and characterized through detailed morphological observations and multilocus phylogenetic analyses based on ITS, LSU, tef1-α, and tub2 sequence data. Additionally, we statistically analyzed the species distribution and host range of Apiospora and Nigrospora and selected the most suitable culture medium for morphological observation by testing multiple media. The objectives of this study were to clarify the taxonomy of these fungi, describe their morphological diversity, and elucidate their phylogenetic relationships and ecological associations within these diverse ecosystems. Comprehensive descriptions, illustrations, and phylogenetic placements of the newly identified taxa are provided herein.
Materials and methods
Specimen collection and isolation
Samples were collected from different hosts, including Poaceae (bamboo), Zea mays L. (maize), and Juglans regia L. (walnut), in various localities across Southwestern China during 2023–2024. All sample collections were placed in sterile paper bags and brought back to the Plant Pathology Laboratory at Guizhou University for fungal observation and isolation. The collected materials were first examined under a stereomicroscope to observe conidiomata and conidia. Fungi were isolated using two complementary techniques depending on the specimen condition: 1) spore suspension isolation was carried out for the species exhibiting conidiomata; 2) tissue isolation was used for samples without conidiomata but exhibiting disease symptoms, both following the protocol of Senanayake et al. (2020). Subsequently, culture plates containing spore suspensions or sterilized leaf fragments were incubated at 25 °C for 24 h. A single germinated spore or hyphal tip was then transferred to potato dextrose agar (PDA) and incubated at 25 °C for approximately 10 days to obtain pure cultures. Pure cultures were transferred to fresh PDA plates for morphological observation and DNA extraction. The fungal cultures were stored in sterile water and 30% glycerol in a refrigerator at 4 °C. The specimens were dried and stored in the Herbarium of the Department of Plant Pathology, Agricultural College, Guizhou University, Guizhou, China (HGUP). The cultures were deposited in the Culture Collection at the Department of Plant Pathology, Agricultural College, Guizhou University, Guizhou, China (GUCC). Registration ID numbers were obtained from MycoBank (www.mycobank.org) for the novel species in this study.
Morphological observation
Morphological features of the fungal isolates were examined from cultures grown on PDA, water agar (WA), synthetic low-nutrient agar (SNA), oatmeal agar (OA), cornmeal agar (CMA), and malt extract agar (MEA) for 7–14 days at 25 °C under alternating light and dark conditions. Colony characteristics, including color, texture, growth rate, and pigmentation were recorded after 7 and 14 days of incubation. After 14 days, conidia formed on the culture medium were examined and documented. The colony color recognition was performed using the color charts of Rayner (1970). Microscopic structures were observed and photographed using a Zeiss Scope 5 compound microscope (Carl Zeiss Microscopy GmbH, Jena, Germany) with an attached camera, Axioscope 5 (AxioCam 208 color). The dimensions of the different spore structures were measured using ZEN 3.0 (blue edition) software (Jena, Germany). At least 30 measurements were taken for each significant morphological structure (e.g., conidia, conidiogenous cells, asci, and ascospores) to determine size ranges and mean values. Illustrations were prepared using Adobe Photoshop CS6 (Adobe Systems Inc., USA) to assemble images and adjust brightness and contrast without altering the original structure or color.
DNA extraction, PCR amplification, and sequencing
Genomic DNA was extracted from 7–10–day-old pure cultures grown on PDA using a BIOMIGA Fungal Genomic DNA Extraction Kit (Biomiga#GD2416, San Diego, California, USA) according to the manufacturer’S protocol. The extracted DNA was stored at – 20 °C until further use. Four loci were amplified for phylogenetic analyses: the internal transcribed spacer (ITS) region, the large subunit ribosomal RNA gene (LSU), the translation elongation factor 1-α (tef1-α), and the beta tubulin (tub2) gene. The primer pairs and amplification conditions are summarized in Suppl. material 3: table SS1. The following primer pairs were used: ITS5/ITS4 for ITS (White et al. 1990), LR0R/LR5 for LSU (Vilgalys and Hester 1990), EF1-728F/EF2 for tef1-α (Carbone and Kohn 1999), and Bt2a /T1/Bt2b for tub2 (Glass and Donaldson 1995). Polymerase chain reactions (PCR) were performed in a total volume of 25 µL, containing 12.5 µL of 2 × T5 Super PCR Mix (Tsingke Biotech Co., China), 1 µL of each primer (10 µM), 1 µL of DNA template, and 9.5 µL of sterile distilled water. PCR amplifications were carried out in a thermocycler (Applied Biosystems, USA) under the conditions listed in Suppl. material 3: table SS1. The PCR products were visualized on 1% agarose gels stained with ethidium bromide, and successful amplicons were purified and sequenced by Sangon Biotech in Chengdu, China. Forward and reverse sequence reads were assembled and edited using Geneious Prime v2023.2.2 (Biomatters Ltd., New Zealand). Consensus sequences were deposited in GenBank, and accession numbers are provided in Table 1.
Table 1.
GenBank accession numbers of the phylogenetic analysis in this study. (Species marked in red are new species discovered in this study, while those marked in blue are species for which new host records were identified in this study or species for which the taxonomic name has changed). “T” indicates the type strain.
Phylogenetic analyses
All sequences generated in this study were initially compared against the NCBI nucleotide database using the BLAST tool (https://blast.ncbi.nlm.nih.gov/) to determine closely related taxa. Representative reference sequences of related genera and species were selected for subsequent phylogenetic analyses based on BLAST results and recent taxonomic publications (Liu et al. 2024a; Yan and Zhang 2024; Zou et al. 2024; Chang et al. 2025; Yu et al. 2025). GenBank accession numbers of all sequences used in this study are listed in Table 1. Raw sequence chromatograms were trimmed and edited using the Trim Ends function in Geneious Prime v2023.2.2 (Biomatters Ltd., New Zealand), and individual loci were assembled into consensus sequences. Each locus was aligned using MAFFT v7.520 (https://mafft.cbrc.jp/alignment/server/) with default settings (Katoh et al. 2019). The resulting alignments were manually checked and trimmed using TrimAl v1.3 (Capella-Gutiérrez et al. 2009) to remove ambiguously aligned regions. Concatenated datasets were generated with PhyloSuite v1.2.3 (Zhang et al. 2020).
Phylogenetic analyses were conducted using both Maximum Likelihood (ML) and Bayesian Inference (BI) algorithms. The best-fit substitution model for each gene partition was determined using ModelFinder implemented in IQ-TREE v2.2.0 and is summarized in Suppl. material 3: table S2. ML analyses were performed with 1,000 ultrafast bootstrap replications to assess branch support. Bayesian analyses were carried out in MrBayes v3.2.7a, running two parallel analyses of four Markov Chain Monte Carlo (MCMC) chains for 5,000,000 generations, sampling every 1,000 generations, and discarding the first 25% of trees as burn-in. Phylogenetic trees were visualized in FigTree v1.4.0 (http://tree.bio.ed.ac.uk/software/figtree) and annotated in Adobe Illustrator 2023 (Adobe Systems Inc., USA) for final presentation.
Host and geographical distribution of Nigrospora and Apiospora
To assess the host associations and geographic distribution of Apiospora and Nigrospora, species occurrence records for both genera were retrieved from the Global Biodiversity Information Facility (GBIF) database (GBIF.org, accessed 19 January 2026; Occurrence Downloads: https://doi.org/10.15468/dl.dx2b4u and https://doi.org/10.15468/dl.4ng2ze). The downloaded datasets were curated and standardized, and the following fields were extracted for analysis: GBIF ID, habitat, associated taxa (host), host family, country of collection, and accepted scientific name (Suppl. material 3: tables S3, S4). Records lacking information on habitat or associated taxa (host) were excluded from further analyses.
In total, 2,354 occurrence records for Apiospora and 1,785 for Nigrospora were obtained. Of these, 2,231 Apiospora records and 1,759 Nigrospora records contained country-level collection data and were retained for geographic analyses. Subsequent data processing, statistical analyses, and graphical visualization were conducted using R software.
Results
Phylogenetic analyses
For family Apiosporaceae (Fig. 1), a concatenated alignment comprising ITS (1–615), LSU (616–1361), tef1-α (1362–2368), and tub2 (2369–2613) loci was analyzed, including 194 strains with Neoarthrinium lithocarpicola (CFCC 54456 and CFCC 55883) designated as outgroup taxa. Eleven strains in this research were incorporated into the dataset. The ML topology served as the reference, with Bayesian posterior probabilities mapped onto corresponding branches. Both topologies were identical, and the best-scoring RAxML tree with a final ML optimization likelihood value of – 38398.438 is presented. Estimated base frequencies were as follows: A = 0.232, C = 0.258, G = 0.256, T = 0.254; substitution rates AC = 1.17641, AG = 2.69166, AT = 1.17641, CG = 1.0, CT = 4.67220, GT = 1.0. The phylogenetic tree clearly demonstrates that the three genera Apiospora, Nigrospora, and Arthrinium within Apiosporaceae form distinct and well-separated clades.
Figure 1.
Phylogenetic tree constructed using Maximum likelihood (ML) analyses based on the multilocus alignment (ITS, LSU, tef1-α, and tub2) for Apiosporaceae. The outgroup is Neoarthrinium lithocarpicola (CFCC 54456 and CFCC 55883). Maximum likelihood bootstrap proportions ≥ 70% (left) and Bayesian inference posterior probability ≥ 0.8 (right) are indicated at nodes (ML/BI). The type strains are followed by “T” and bold. New species in this study are red, and new records and merged species are blue.
Phylogenetic analyses further revealed several instances of minimal or no genetic divergence among certain taxa; A. hispanica (IMI 326877) and A. mediterranea (IMI 326875) clustered together without detectable genetic distance. Similarly, A. vietnamensis (IMI 99670, KUMCC 21-0428, KUMCC 21-0429), A. malaysiana (CBS 102053, CBS 251.29), A. magnispora (ZHKUCC 22-0001T), and A. euphorbiae (IMI 285638b) formed a closely related cluster with little genetic divergence (ML/BI = 96/0.63). Apiospora jinanensis (SAUCC DL1981) and A. thailandica (MFLUCC 15-0202) grouped together with no genetic distance, as did A. sorghi (URM 93000) and A. taeanensis (KUC21322). Notably, Ar. sinense (HKUCC 3143) clusters within the Apiospora clade, suggesting potential taxonomic implications that may warrant further investigation.
The two phylogenetic trees of Apiospora and Nigrospora were conducted separately for each genus to more precisely determine their interrelationships. For Apiospora (Fig. 2), a concatenated alignment comprising ITS (1–466), LSU (467–1220), tef1-α (1221–1935), and tub2 (1936–2303) loci was analyzed, including 282 strains with Ne. lithocarpicola (CFCC 54456 and CFCC 55883) designated as outgroup taxa. The ML topology served as the reference, with Bayesian posterior probabilities mapped onto corresponding branches. Both topologies were identical, and the best-scoring RAxML tree with a final ML optimization likelihood value of – 29198.663 is presented. Estimated base frequencies were as follows: A = 0.237, C = 0.250, G = 0.254, T = 0.259; substitution rates AC = 1.29435, AG = 3.16215, AT = 1.03614, CG = 0.85141, CT = 4.90526, GT = 1.0. Sequences from 15 isolates obtained in this study were incorporated into the dataset, forming six well-supported lineages within Apiospora, representing four new species (A. huaxiensis sp. nov., A. nanmingensis sp. nov., A. qingzhenensis sp. nov., and A. tongrenensis sp. nov.) and two known species (A. locuta-pollinis and A. setariae). Apiospora huaxiensis (GUCC 25-0068, GUCC 25-0069, and GUCC 25-0070) formed a sister clade with A. bambusirimae (GUCC12.51 and GUCC12.52) with high support (ML/BI = 100/1). Apiospora nanmingensis (GUCC 25-0087 and GUCC 25-0088), formed a sister clade with A. hysterina and A. sasae, with high support (ML/BI = 100/1). Apiospora qingzhenensis (GUCC 25-0077, GUCC 25-0078, GUCC 25-0079, GUCC 25-0080), formed a single clade with high support (ML/BI = 100/1). Apiospora tongrenensis (GUCC 25-0083 and GUCC 25-0084) formed a sister clade with A. saccharicola, with high support(ML/BI = 99/1). Apiospora locuta-pollinis and A. setariae each clustered separately from other strains of the species with high support values (ML/BI = 100/0.96 and ML/BI = 100/1). Arthrinium sinense (HKUCC 3143) clusters within the clade of A. bambusilentiginis, A. guizhouensis, A. armeniaca, A. elliptica, A. cannae, and A. sacchari with high support (ML/BI = 86/-). Apiospora hispanica (IMI 326877) and A. mediterranea (IMI 326875) clustered together on the phylogenetic tree with no genetic distance with a high support value (ML/BI = 100/0.93), which is consistent with the previous studies in Tian et al. (2021), Monkai et al. (2022), and Liao et al. (2023). Apiospora vietnamensis (IMI 99670, KUMCC 21-0428, KUMCC 21-0429), A. malaysiana (CBS 102053, CBS 251.29), A. magnispora (ZHKUCC 22-0001), and A. euphorbiae (IMI 285638b) clustered together with minimal genetic distance with a high support value (ML/BI = 100/1).
Figure 2.
Phylogenetic tree constructed using Maximum likelihood (ML) analyses based on the multilocus alignment (ITS, LSU, tef1-α, and tub2) for Apiospora. The outgroup is Neoarthrinium lithocarpicola (CFCC 54456 and CFCC 55883). Maximum likelihood bootstrap proportions ≥ 70% (left) and Bayesian inference posterior probability ≥ 0.8 (right) are indicated at nodes (ML/BI). The type strains are followed by “T” and bold. New species in this study are red, and new records and merged species are blue.
For Nigrospora (Fig. 3), phylogenetic relationships were inferred from a combined dataset of ITS (1–522), tef1-α (523–961), and tub2 (962–1752) loci, comprising 132 strains with A. pseudoparenchymatica (LC 7234) and A. malaysiana (CBS 102053) as outgroup taxa. The ML topology was used as the reference, with Bayesian posterior probabilities mapped onto corresponding branches, and the best-scoring RAxML tree with a final ML optimization likelihood value of – 17739.687 is presented. Estimated base frequencies were as follows: A = 0.215, C = 0.306, G = 0.243, T = 0.237; substitution rates AC = 1.0, AG = 3.25843, AT = 1.0, CG = 1.0, CT = 4.42079, GT = 1.0. Twenty newly obtained isolates clustered into five well-supported lineages, representing one novel species (N. neosaccharicola sp. nov.) and four known species (N. chinensis, N. endophytica, N. osmanthi, and N. sphaerica). Nigrospora neosaccharicola (GUCC 25-0124, GUCC 25-0125, GUCC 25-0126, GUCC 25-0127, GUCC 25-0128, and GUCC 25-0129) formed a sister clade with N. saccharicola (CGMCC 3.19362 and LC 4241) with a high support value (ML/BI = 100/1). GUCC 25-0130, GUCC 25-0131, GUCC 25-0132, and GUCC 25-0133 clustered to the N. osmanthi clade with high support values (ML/BI = 100/1). GUCC 25-0122 and GUCC 25-0123 clustered to N. endophytica with high support values (ML/BI = 96/-). GUCC 25-0120 and GUCC 25-0121 clustered to N. chinensis with high support values (ML/BI = 100/1). GUCC 25-0134, GUCC 25-0135, GUCC 25-0136, GUCC 25-0137, GUCC 25-0138, and GUCC 25-0139 clustered to N. sphaerica with high support values (ML/BI = 100/1).
Figure 3.
Phylogenetic tree constructed using Maximum likelihood (ML) analyses based on the multilocus alignment (ITS, tef1-α, and tub2) for Nigrospora. The outgroups are A. pseudoparenchymatica (LC 7234T) and A. malaysiana (CBS 102053). Maximum likelihood bootstrap proportions ≥ 70% (left) and Bayesian inference posterior probability ≥ 0.8 (right) are indicated at nodes (ML/BI). The type strains are followed by “T” and bold. New species discovered in this study are red, and new records are blue.
Host and geographical distribution of Apiospora and Nigrospora
According to GBIF data, based on host association, Poaceae represents the dominant host family for both genera (Fig. 4). For Apiospora, 1,710 records (72.64% of all host-associated records) were linked to Poaceae, with bamboo accounting for the largest proportion (868 records, 36.87% of all host-associated records). Additional host families with notable representation include Fabaceae, Cyperaceae, and Pinaceae. In contrast, 42 records were associated with non-plant substrates or atypical hosts, such as soil, animals, humans, and other fungi. Similarly, Nigrospora species were predominantly associated with Poaceae, comprising 697 records (39.05% of host-associated records). Additional host families with notable representation include Fabaceae, Rosaceae, and Asteraceae (Fig. 4). Regarding geographical distribution, Apiospora species are most frequently recorded in Europe, followed by Asia and North America (Fig. 5). In contrast, Nigrospora species are most widely reported from Asia, with Africa representing the second most recorded region (Fig. 5). At the country level, the highest numbers of Apiospora records originate from the USA, UK, and China (Fig. 6). For Nigrospora, the top three countries are China, India, and the USA (Fig. 6).
Figure 4.
Host-substratum diversity of Apiospora (A) and Nigrospora (B) species according to GBIF.
Figure 5.
Apiospora (A) and Nigrospora (B) species distribution by continent according to GBIF.
Figure 6.
Distribution of Apiospora (A) and Nigrospora (B) species by countries according to GBIF.
Taxonomy
Apiosporaceae K.D. Hyde, et al. 1998.
Apiospora Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 85. 1875.
Apiospora hispanica
(Larrondo & Calvo) Pintos & P. Alvarado, Fungal Systematics and Evolution 7: 205. 2021.
B8C1FFC2-09C4-5AAD-9B92-CFAD670D68F8
837674
Basionym.
Arthrinium hispanicum Larrondo & Calvo, Mycologia 84(3): 476. 1992.
Synonyms.
Arthrinium mediterranei Larrondo & Calvo, Mycologia 84(3): 476. 1992.
Apiospora mediterranea (Larrondo & Calvo) Pintos & P. Alvarado, Fungal Systematics and Evolution 7: 206. 2021. syn. nov.
Substrate and distribution.
Beach sand and air, Spain (Larrondo and Calvo 1992).
Notes.
Arthrinium hispanicum was originally described by Larrondo and Calvo (1992) from maritime sand in Spain, while Ar. mediterranei was isolated from the air in the same region (Larrondo and Calvo 1992). The two taxa were morphologically distinguished primarily by conidial dimensions (7.5–8.5 × 6.2–7.6 µm vs. 9–9.5 × 7.5–9 µm, respectively). Pintos and Alvarado (2021) later clarified the relationship between Apiospora and Arthrinium, transferring both Ar. hispanicum and Ar. mediterranei to Apiospora based on multilocus phylogenetic analyses. However, they did not explicitly evaluate whether the two taxa represent a single species. Our phylogenetic analyses reveal that strains of A. hispanica (IMI 326877) and A. mediterranea (IMI 326875) form a single clade without genetic distinction. Comparative analyses of ITS, LSU, and tub2 sequences showed no nucleotide differences between the two species. Based on the congruence of morphological features and the molecular divergence, we accept A. hispanica as the correct name and regard A. mediterranea syn. nov. as its synonym.
Apiospora huaxiensis
J. E. Sun, X. C. Wang, K.D. Hyde & Yong Wang bis sp. nov.
1BB615FD-02B8-5B35-8A55-B8EBD48E2250
860108
Figure 7.
Apiospora huaxiensis (GUCC 25-0068). A, B Appearance of the fungus on spot leaves of bamboo; C–H Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days. (I) Close up of PDA culture with conidia; J, K Conidia with conidiogenous cells. (L) Conidia. (M) Conidia with germ slit. Scale bars: 100 μm (B); 10 μm (J–M).
Etymology.
The species name refers to the collected location, Huaxi District, Guizhou Province.
Holotype.
China • Guizhou Province, Guiyang City, Huaxi District, Huaxi Park, on the leaf spots of bamboo, 23 November 2023. X.C. Wang, HGUP 25-0006 (holotype); ex-type GUCC 25-0068.
Description.
Associated with the leaf spots of bamboo. Lesions are black and brown spots with yellow halos on leaves, subglobose and ellipsoidal in shape, measuring 250–700 μm in diam. Sexual morph: Not observed. Asexual morph: on PDA, Hyphae 2.4–4.1 μm wide, branched, septate, hyaline to pale green. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3.6–6.5 × 2.3–5.8 μm (x̄ =5.2 × 3.2 μm, n=30), cylindrical or round, monoblastic, pale green. Conidia 13.8–17.6 × 11.3–16.4 μm (x̄ =15.8 × 13.7 μm, n=30), globose to ellipsoidal granules, pale green to dark brown, with longitudinal germ-slit.
Culture characteristics.
After 7 days at 25°C, on PDA, colonies reach 70–74 mm in diam., with irregular margins, flat, cottony, dense aerial mycelia, pale red coloration, and a red reverse; they produce scarlet pigment. On MEA, colonies reach 88–90 mm diam., with regular margins, flat, dense, floccose, surface pale salmon and reverse salmon. On CMA, colonies reach 85–87 mm diam., cottony, flat, dense, irregular margins with aerial mycelia, surface straw, and reverse pale luteous. On SNA, colonies reach 55–59 mm diam., flat, spreading, with sparse aerial mycelia and regular margin, surface and reverse white. On WA, colonies reach 22–26 mm diam., flat, aerial mycelia scant, surface and reverse white. On OA, colonies reach 72–74 mm diam., flat, cottony, with regular floccose margin, surface pale ochreous and reverse umber middle area and pale-yellow outer area. Sporulation was abundant only on PDA after 14 days.
Material examined.
China • Guizhou Province, Guiyang City, Huaxi District, Huaxi Park, on the leaf spots of bamboo, 23 November 2023, X.C. Wang, HGUP 25-0006 (holotype); GUCC 25-0068 (ex-type), GUCC 25-0069 and GUCC 25-0070.
Notes.
Apiospora huaxiensis (GUCC 25-0068, GUCC 25-0069, and GUCC 25-0070) formed a sister clade to A. bambusirimae (GUCC 12.51) with strong statistical support (100% ML/1.0 BI) (Fig. 2). Morphologically, the conidia of A. huaxiensis are smaller than those of A. bambusirimae (x̄ = 15.8 × 13.7 μm vs. x̄ = 20.5 × 13 μm; Yu et al. 2025). In culture, A. huaxiensis produced reddish pigment on PDA after 3 days, whereas A. bambusirimae did not on the same cultivation conditions (Yu et al. 2025). Moreover, pairwise sequence comparisons between the two species revealed nucleotide differences of 2.34% in ITS (11/469 bp, including six gaps) and 2.65% in tef1-α (12/453 bp, including nine gaps).
Apiospora locuta-pollinis
(F. Liu & L. Cai) Pintos & P. Alvarado, Fungal Systematics and Evolution 7: 206. 2021.
0A314D1F-8365-5336-894A-8283350FD193
837763
Figure 8.
Apiospora locuta-pollinis (GUCC 25-0071). A Appearance of the fungus on leaves of maize; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H, O Close up of WA; H and SNA; O cultures with conidia; I–M Conidia with conidiogenous cells on WA; N coiled hyphae on WA; P–U Conidia with conidiogenous cells on SNA. Scale bars: 100 μm (H, O); 10 μm (I–N, P–U).
Basionym.
Arthrinium locutum-pollinis F. Liu & L. Cai [as ‘locuta-pollinis’], in Zhao, Zhang et al., Mycosphere 9(6): 1094. 2018.
Synonym.
Arthrinium pseudomarii T.Z. Chen, Yong Wang bis & K.D. Hyde, in Chen, Zhang et al., Mycotaxon 136(1): 191. 2021.
Substrate and distribution.
On hive-stored pollen, Hubei Province, China (Zhao et al. 2018); Aristolochia debilis, China (Chen et al. 2021); Bamboo, China (Monkai et al. 2022); Musa sp., Thailand (Samarakoon et al. 2024); grass, China (Gao et al. 2025).
Description.
Associated with the leaf spots of maize. Lesions are yellow spots with pale brown halos on leaves, subglobose and spindle-shaped in shape, measuring 300–1000 μm in diam. Sexual morph: Not observed. Asexual morph: on WA, Hyphae 2.1–4.1 μm wide, occasionally coiled, septate, branched hyphae, hyaline. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3.9–5.1 × 1.9–3.2 μm (x̄ = 4.2 × 2.1 μm, n = 30), subglobose to ampulliform, mostly polyblastic, hyaline to pale green. Conidia 4.2–10.1 × 3.9–8.7 μm (x̄ = 7.7 × 5.9 μm, n = 30), globose to subglobose, smooth-walled, pale green when immature, brown when mature. On SNA, Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3.5–8.2 × 2.1–4.2 μm (x̄ = 4.9× 2.7 μm, n = 30), subglobose to ampulliform, mostly polyblastic, pale green. Conidia 4.8–13.9 × 3.7–7.6 μm (x̄ = 8.2× 6.2 μm, n = 30), globose, subcylindrical to ovate, smooth, brown.
Culture characteristics.
After 7 days at 25°C, on PDA, colonies reach 88–90 mm in diam., flat, cottony, with regular margins; surface pale straw, reverse salmon. On MEA, colonies reach 70–74 mm diam., flat, with regular margins, surface, and reverse white. On CMA, colonies reaching 56–58 mm in diam., flat, with regular margins, surface, and reverse white. On SNA, colonies reach 77–79 mm in diam., flat, spreading, with scant aerial mycelia, and a regular margin, surface, and reverse that are white. On WA, colonies reach 42–45 mm diam., flat, spreading, with irregular margins, aerial mycelia scant, surface and reverse white. On OA, colonies reach 81–84 mm diam., circular, flat, cottony, with regular margin, surface white and reverse pale saffron. Sporulation was abundant on WA and SNA after 14 days.
Material examined.
• Guizhou Province, Liupanshui City, leaf spot on Zea mays, 16 September 2024, X.C. Wang, HGUP 25-0058; GUCC 25-0071; • ibid. GUCC 25-0072.
Notes.
GUCC 25-0071 and GUCC 25-0072 clustered with A. locuta-pollinis (LC 11683). Morphologically, the new collections also closely resemble A. locuta-pollinis (LC 11683), with globose to subglobose conidia (x̄ = 7.7 × 5.9 μm vs. x̄ = 7.1 × 6.4 μm) and subglobose to ampulliform conidiogenous cells (x̄ = 4.2 × 2.1 μm vs. x̄ = 4.9 × 3.8 μm) (Zhao et al. 2018). Nucleotide comparison with LC 11683 revealed that only tef1-α exhibited two nucleotide differences (2/430). Therefore, strains GUCC 25-0071 and GUCC 25-0072 are identified as A. locuta-pollinis. This is the first report of A. locuta-pollinis isolated from maize.
Apiospora nanmingensis
X. C. Wang, S. Q. Guo, K.D. Hyde & Yong Wang bis sp. nov.
E3288603-4944-526E-8BF5-08EDF78B43F7
861263
Figure 9.
Apiospora nanmingensis (GUCC 25-0087). A Appearance of the fungus on stem of bamboo; B Conidiomata on host; C–H Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; I Close up of SNA culture with conidia; J–Q Conidia with conidiogenous cells on SNA; R Conidia. Scale bars: 100 μm (B, I); 10 μm (J–R).
Etymology.
This species was named according to the location where the fungus was first collected, Nanming District, Guiyang City, Guizhou Province.
Holotype.
China • Guizhou Province, Guiyang City, Nanming District, Guiyang Forest Park, on the stem of bamboo, 1 April 2024. X.C. Wang, HGUP 25-0053 (holotype); ex-type GUCC 25-0087.
Description.
Isolated from decaying stems of bamboo. On the host, Conidiomata punctiform, brown, 150–400 µm × 125–200 µm (n = 30). Conidia 9.2 × 15.2–7.4 × 13.2 μm (x̄ = 13.2 × 11.4 μm, n=30), globose to subglobose, brown. Sexual morph: Not observed. Asexual morph: on SNA, Hyphae1.5–3.8 µm wide, branched, septate, smooth, hyaline. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3.9–11.3 × 2.5–3.9 μm (x̄ = 6.8 × 3.1 μm, n=30), cylindrical to globose, monoblastic to mostly polyblastic, aggregated, hyaline to pale green. Conidia 7.6–10.4 × 6.9–10.1 μm (x̄ = 8.9 × 7.5 μm, n=30), globose, subglobose to lenticular, with longitudinal germ slit, pale green to brown.
Culture characteristics.
Colonies for 7 days at 25°C: On PDA, colonies reach 88–90 mm diam., flat, spreading, with irregular margins, dense aerial mycelia, surface white and reverse pale salmon center and white margin. On MEA, colonies reach 57–60 mm diam., flat, floccose, cottony, with regular margins, surface and reverse white. On CMA, colonies reach 77–80 mm diam., flat, with regular margins, spreading, with moderate aerial mycelia, surface and reverse pale salmon. On SNA, colonies reach 87–90 mm diam., flat, aerial mycelia scant, with regular margins, surface and reverse white. On WA, colonies reach 63–65 mm diam., flat, aerial mycelia scant, with regular margins, surface and reverse white. On OA, colonies reach 70–72 mm diam., cottony, with dense aerial mycelia, regular margins, surface white and reverse pale straw. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Guizhou Province, Guiyang City, Nanming District, Guiyang Forest Park, on the stem of bamboo, 1 April 2024. X.C. Wang, HGUP 25-0053 (holotype); GUCC 25-0087 (ex-type), GUCC 25-0088.
Notes.
Based on the phylogenetic analyses, the two strains obtained in this study, GUCC 25-0087 and GUCC 25-0088, formed a distinct, well-supported lineage (100% ML/1.0 BI) with A. hysterina, A. sasae, and A. yunnana (Fig. 2). Morphologically, our strains can be readily distinguished from these closely related taxa by their smaller conidia, which measure 7.6–10.4 × 6.9–10.1 µm (vs. 15–18 × 14–16.5 µm in A. hysterina; Kwon et al. 2022; Saccardo 1893;17–18 × 16–17 µm in A. sasae; Crous et al. 2021; and 17.5–26.5 × 15.5–25 µm in A. yunnana; Dai et al. 2017). The conidiogenous cells are likewise smaller than those of A. yunnana (3.9–11.3 × 2.5–3.9 µm vs. 16.5–50 × 2–4 µm). Interestingly, in this clade, the shape and size of the conidia of A. hysterina, A. sasae, and A. yunnana are relatively similar, whereas the conidia of A. nanmingensis that we identified differ significantly in shape and size from those of the other species, whether observed on the host or on culture media. Apiospora species are variable in morphological characteristics depending on growth conditions, and molecular and phylogenetic analysis are the key to recognize species in this genus. Pairwise sequence comparisons revealed that strains GUCC 25-0087 and GUCC 25-0088 differ from A. hysterina (ICPM 6889) by 2.72% in ITS (14/441 bp, including two gaps), 1.86% in tef1-α (8/430 bp, four gaps), and 1.19% in tub2 (9/758 bp, one gap). Comparisons with A. sasae (CBS 146808) showed divergences of 2.02% in ITS (10/496 bp, three gaps) and 1.55% in tub2 (12/773 bp). Comparisons with A. yunnana (MFLUCC 15-1002) showed divergences of 4.77% in ITS (26/545 bp, seven gaps) and 0.34% in LSU (3/875 bp, one gap).
Apiospora qingzhenensis
X.C. Wang, K.D. Hyde & Yong Wang bis sp. nov.
5122DDF4-296D-5F00-971D-5CA83CCB81FC
860112
Figure 10.
Apiospora qingzhenensis (GUCC 25-0077). A, B Appearance of the fungus on leaf spots of bamboo; C–H Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; I, O Close up of WA; I and SNA; O cultures with conidia; J–M Conidia with conidiogenous cells on WA; N Conidia on WA; P, Q Conidia with conidiogenous cells on SNA; R Conidia and sterile cell on SNA; S Conidia on SNA. Scale bars: 100 μm (B, I, O); 10 μm (J–N, P–S).
Etymology.
The name refers to Qingzhen City in Guizhou Province, where the fungus was collected.
Holotype.
China • Guizhou Province, Qingzhen City, on the leaves of bamboo with spots, 18 January 2024, X.C. Wang, HGUP 25-0008 (holotype); ex-type GUCC 25-0077.
Description.
Associated with bamboo leaf spots. Lesions are black spots on leaves, subglobose in shape, measuring 250–620 μm in diam. Sexual morph: Not observed. Asexual morph: On WA, Hyphae 2.1–5.9 μm in diam, branched, septate, hyaline. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4.1–6.3 × 2.1–5.7 μm (x̄ = 5.7 × 3.6 μm, n=30), cylindrical, mostly polyblastic, aggregated, hyaline or pale green. Conidia 7.1–17.5 × 6.0–11.4 μm (x̄ = 10.7 × 8.7 μm, n=30), globose, subglobose to ovate with a straight germ-slit along spore length, green to dark brown. On SNA, Conidiogenous cells 5.2–8.3 × 2.7–6.1 μm (x̄ = 6.1 × 3.9 μm, n=30), cylindrical, mostly polyblastic, aggregated, hyaline or pale green. Conidia 6.5–14.3 × 6.0–12.4 μm (x̄ = 11.8 × 9.3 μm, n=30), globose to subglobose, pale green for immature and dark brown for mature. Sterile cells extremely rare, pale brown, rolled up, irregularly lobed.
Culture characteristics.
Colonies for 7 days at 25°C: On PDA, colonies reach 90 mm diam., flat, spreading, cottony, dense aerial mycelia, with regular margins, surface, and reverse pale flesh. On MEA, colonies reach 77–80 mm diam., flat, with moderate aerial mycelia, irregular margins, surface white, and reverse center pale salmon, with margin white. On CMA, colonies reach mm 71–74 diam., flat, spreading, with moderate aerial mycelia, regular margins, surface and reverse pale salmon. On SNA, colonies reach 90 mm diam., flat, aerial mycelia scant, with regular margins, surface and reverse pale white. On WA, colonies reach 53–57 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse white. On OA, colonies reach 81–84 mm diam., floccose, flat, spreading, with dense aerial mycelia, regular margins, surface white and reverse pale salmon. Sporulation was abundant on WA and SNA after 14 days.
Material examined.
China • Guizhou Province, Qingzhen City, on the leaves of bamboo with spots, 18 January 2024, X.C. Wang, HGUP 25-0008 (holotype); GUCC 25-0077 (ex-type), GUCC 25-0078, GUCC 25-0079, and GUCC 25-0080.
Notes.
Based on phylogenetic analyses (Fig. 2), four strains (GUCC 25-0077, GUCC 25-0078, GUCC 25-0079, and GUCC 25-0080) formed a distinct and well-supported clade within Apiosporasensu stricto. The closest related species are A. zhaotongensis, A. zhenxiongensis, and A. bambusiparasitica. Since A. zhaotongensis and A. zhenxiongensis have been reported only from their sexual morphs (Han et al. 2024), morphological comparisons were made solely with A. bambusiparasitica. Morphologically, the new species differs from A. bambusiparasitica by having smaller conidiogenous cells (4.1–6.3 × 2.1–5.7 μm vs. 7–17 × 2.0–4.5 μm) and slightly larger conidia (x̄ = 10.7 × 8.7 μm vs. x̄ = 9.2 ± 0.9 × 8.1 ± 1.1 μm; Chang et al. 2025). Occasionally, pale brown, elongated sterile cells can be observed in our species. Furthermore, pairwise sequence comparisons across four gene regions revealed clear nucleotide differences between the new species and its closest relatives. The new strains differ from A. zhaotongensis (GMBCC 1015) by 4.34% in tub2 (21/484 bp, including seven gaps) and 5.66% in tef1-α (25/442 bp, including five gaps); from A. zhenxiongensis (GMBCC 1017) by 5.15% in tub2 (25/485 bp, including eight gaps) and 5.67% in tef1-α (25/441 bp, including five gaps); and from A. bambusiparasitica (RCEF 20003) by 4.72% in tub2 (23/487 bp, including ten gaps) and 6.39% in tef1-α (28/438 bp, including six gaps).
Apiospora setariae
(C.M. Tian & N. Jiang) X.G. Tian & Tibpromma, Life 11(no. 1071): 19. 2021.
2046CA96-5EC6-59E2-AF86-9CF13455A6F2
558561
Figure 11.
Apiospora setariae (GUCC 25-0081). A Appearance of the fungus on stem of bamboo stems; B Stromata on bamboo stems; C–H Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; I Transverse sections of the stromata; J, K Peridium; L Asci; M–P Ascospores. Scale bar: 100 μm (B, I); 20 μm (J–L); 10 μm (M–P).
Basionym.
Arthrinium setariae C.M. Tian & N. Jiang, in Jiang & Tian, Phytotaxa 484(2): 153. 2020.
Substrate and distribution.
Setaria viridis, China (Jiang and Tian 2021).
Description.
Associated with decaying stems of bamboo. Sexual morph: Stromata 400–1250 × 270–495 μm (n = 30), solitary to gregarious, partially immersed to erumpent, fusiform, raised on the host surface, with a slit-like opening, multi-loculate. Ascomata 70–240 μm (n = 30) diam., uniseriate or irregularly arranged beneath stromata, globose to subglobose. Peridium 5–20 μm (n = 30) thick, composed of 3–5 layers of brown to hyaline cells arranged in textura angularis. Hamathecium not observed. Asci 67.1–91.6 × 10.1–20.4 μm (x̄ = 79.9 × 15 μm, n = 30), 8-spored, broadly cylindrical, with an indistinct pedicel. Ascospores 22.9–29.4 × 6.9–10.9 μm (x̄ = 25.9 × 9 μm, n = 30), broadly ellipsoidal, composed of a large upper cell and a small lower cell, hyaline to pale green, smooth-walled. Asexual morph: refer to (Jiang and Tian 2021).
Culture characteristics—Colonies for 7 days at 25°C: On PDA, colonies reaching 88–90 mm diam., flat, spreading, dense aerial mycelia, with regular margins, surface and reverse scarlet, produce scarlet pigment. On MEA, colonies reach 55–58 mm diam., flat, spreading, with irregular margins, surface and reverse center scarlet and margin white, produce reddish pigment. On CMA, colonies reach 31–33 mm diam., flat, aerial mycelia scant, regular margins, surface and reverse white. On SNA, colonies reach 41–43 mm diam., flat, aerial mycelia scant, regular margins, surface and reverse white. On WA, colonies reach 30–32 mm diam., flat, aerial mycelia scant, regular margins, surface and reverse white. On OA, colonies reach 44–47 mm diam., flat, cottony, dense aerial mycelia, with regular margins, surface white and reverse salmon. Sporulation was absent on any medium after 14 days.
Material examined.
China • Guizhou Province, Guiyang City, Huaxi District, Huaxi Park, on the decaying bamboo stems, 17 April 2024, X.C. Wang, HGUP 25-0059; GUCC 25-0081 and GUCC 25-0082.
Notes.
In phylogenetic analyses, the newly generated strains in this study, GUCC 25-0081 and GUCC 25-0082, clustered with A. setariae (strains CFCC 54041 and beilin024) with ML/BI = 100/1 (Fig. 2). Morphologically, our new collections are very similar to A. setariae in the asci size (67.1–91.6 × 10.1–20.4 μm vs. 60–95 × 15–21 μm) and ascospores (22.9–29.4 × 6.9–10.9 μm vs. 18–23 × 8–11 μm, Jiang and Tian 2021). Therefore, both collections are identified as A. setariae. Previously, A. setariae was reported only from decaying culms of Setaria viridis (Jiang and Tian 2021). The comparison of nucleotide differences between strain GUCC 25-0081 and the type strain of A. setariae (CFCC 54041T) revealed clear sequence divergence: 0.69% in ITS (4/583 bp, including one gap), 0.41% in tub2 (3/730 bp, including one gap), and 0.46% in tef1-α (2/439 bp, no gaps). In this study, the species was isolated from bamboo stems, representing a new host record for A. setariae.
Apiospora sinense
K.D. Hyde, J. Fröhl. & Joanne E. Taylor, Sydowia 50(1): 27. 1998 syn. nov.
12F85AAE-322C-5344-9B79-DFBFD131F025
816836
Synonym.
Arthrinium sinense (K.D. Hyde, J. Fröhl. & Joanne E. Taylor) Crous & J.Z. Groenew., in Réblová et al., IMA Fungus 7(1): 140. 2016.
Substrate and distribution.
On dead petiole of Trachycarpus fortunei, Hubei Province, China (Hyde 1998).
Morphological description.
refer to Hyde et al. (1998).
Notes.
Apiospora sinensis was described initially from dead petiole of Trachycarpus fortunei by Hyde et al. (1998). Due to uncertainty regarding the taxonomic boundaries between Apiospora and Arthrinium, Crous and Groenewald in 2016 subsequently transferred this species to Arthrinium based on phylogenetic analysis (Réblová et al. 2016). Pintos and Alvarado (2021) clarified the generic delimitation between Apiospora and Arthrinium, and reassigned most species previously placed in Arthrinium to Apiospora. Currently, only LSU sequence data (AY083831) is available for this species in GenBank. In our phylogenetic analyses, Ar. sinensis (HKUCC 3143) clustered within Apiosporasensu stricto and formed an independent branch distinct from other taxa. Compared to the reported morphology of this species, its asexual form, the conidia, exhibit the following characteristics: 9–12 × 6–8 μm, mainly rounded in face view, mainly lenticular, brown, with an equatorial germ slit, smooth (Hyde et al. 1998), which shares the same conidial characteristics as those reported for Apiospora (Pintos and Alvarado 2021). Therefore, Ar. sinense is reinstated in Apiospora to accommodate this taxon.
Apiospora tongrenensis
S.Q. Guo, X.C. Wang, K.D. Hyde & Yong Wang bis sp. nov.
4FF98D5A-A06B-55AB-BA61-F8085117334B
860113
Figure 12.
Apiospora tongrenensis (GUCC 25-0083). A Collecting location; B Appearance of the fungus on spot leaves of bamboo; C–H Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; I Close up of SNA culture with conidia. (J, L, O–Q) Conidia with conidiogenous cells on SNA. (K, M, N) Elongated conidia (sterile cells) with conidiogenous cells on SNA; R Conidia; S Elongated conidia (sterile cells). Scale bars: 100 μm (B); 10 μm (J–S).
Etymology.
Named according to Tongren, Guizhou Province, China, where the species was collected.
Holotype.
China • Guizhou Province, Tongren City, Foding Mountain, leaf spots on bamboo, 10 July 2024, S.Q. Guo, HGUP 25-0009 (holotype), ex-type living culture GUCC 25-0083.
Description.
Associated with leaf spots on bamboo. Lesions as pale-yellow spots on leaves, subglobose and ellipsoidal in shape, measuring 500–2000 × 240–1200 μm. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 1.9–3.6 μm wide, branched, coiled hyphae, septate, hyaline to pale green. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 6–18.4 × 3.1–6.6 μm (x̄ = 12.4 × 4.2 μm, n=30), cylindrical, aggregated, monoblastic to polyblastic, smooth, hyaline or brown. Conidia immature pale green, mature brownish, smooth-walled, globose or subglobose, 10–14.9 × 8.5–11.4 μm (x̄ = 13 × 9.7 μm, n=30), sometimes with longitudinal germ slit. Elongated conidia (sterile cells) pale brown, elongate, 17.6–32.5 × 4.5–8.8 μm (x̄ = 22.3 × 6.9 μm, n=30).
Culture characteristics.
Colonies after 7 days at 25°C: On PDA, colonies reach 90 mm diam., flat, spreading, dense aerial mycelia, with regular margins, surface white and reverse pale salmon. On MEA, colonies reach 80–82 mm diam., flat, cottony, dense aerial mycelia, with regular margins, surface white and reverse pale salmon. On CMA, colonies reach 77–80 mm diam., flat, spreading, with regular margins, surface white, and reverse pale salmon. On SNA, colonies reach 83–85 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse white. On WA, colonies reach 47–50 mm diam., flat, center dense aerial mycelia, and margin scant, surface and reverse white. On OA, colonies reach 64–67 mm diam., flat, cottony, dense aerial mycelia, with regular margins, surface white and reverse salmon. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Guizhou Province, Tongren City, Foding Mountain, leaf spots on bamboo, 10 July 2024, S.Q. Guo, HGUP 25-0009 (holotype); GUCC 25-0083 (ex-type), GUCC 25-0083 and GUCC 25-0084.
Notes.
Phylogenetic analyses (Fig. 2) revealed that the new species, A. tongrenensis (GUCC 25-0083 and GUCC 25-0084) formed a distinct and well-supported branch with A. saccharicola (99% ML/1.0 BI). Morphologically, the two species are clearly different, with A. tongrenensis possessing larger conidiogenous cells (6–18.4 × 3.1–6.6 μm vs. 5–12 × 2.5–4 μm; Crous and Groenewald 2013). The conidia of A. tongrenensis are also larger than those of A. saccharicola (Crous and Groenewald 2013). Pairwise sequence comparisons between A. tongrenensis (GUCC 25-0083) and A. saccharicola (CBS 191.73) revealed nucleotide differences of 1% in ITS (6/600, no gaps), 0.82% in LSU (10/1225, no gaps), 3.47% in tub2 (28/806, including six gaps), and 0.92% in tef1-α (4/435 bp, no gaps).
Apiospora vietnamensis
(Hol.-Jech.) Pintos & P. Alvarado, Fungal Syst. Evol. 7: 207. 2021.
7D2B2152-B4C6-57DE-87D3-4FC1C946A6D4
837737
Basionym.
Nigrospora vietnamensis Hol.-Jech., Česká Mykol. 17(1): 19. 1963.
Synonyms.
Arthrinium euphorbiae M.B. Ellis, Mycol. Pap. 103: 6. 1965.
Arthrinium malaysianum Crous, in Crous & Groenewald, IMA Fungus 4(1): 144. 2013. syn. nov.
Arthrinium vietnamense (Hol.-Jech.) Mei Wang & L. Cai [as ‘vietnamensis’], in Wang, Liu, et al., Persoonia 39: 139. 2017. syn. nov.
Apiospora euphorbiae (M.B. Ellis) X.G. Tian & Tibpromma, in Tian, Karunarathna, et al., Life 11(no. 1071): 17. 2021. syn. nov.
Apiospora malaysiana (Crous) Pintos & P. Alvarado, Fungal Syst. Evol. 7: 206. 2021. syn. nov.
Apiospora magnispora H.J. Zhao, Manawas. & W. Dong, in Zhao, Dong, et al., Curr. Res. Envir. & App. Myc. 13(1): 9. 2023. syn. nov.
Substrate and distribution.
On decayed fruit of Citrus sinensis, Czech Republic (Holubová-Jechová 1963); On dead stems of Euphorbia, Zambia (Ellis 1965); Macaranga hullettii, Malaysia (Crous and Groenewald 2013); Cinnamomum camphora (Crous and Groenewald 2013); Ficus septica, Taiwan (China) (Tennakoon et al. 2021); Bambusa textilis, China (Zhao et al. 2023); Bats, China (Liu et al. 2023b). Other distributions such as Vietnam, Indonesia, Brazil, Portugal, India, United Kingdom, and The United Republic of Tanzania record in GBIF.
Notes.
The species A. vietnamensis was originally described as N. vietnamensis from decayed fruit of Citrus sinensis (Jechová 1963) and later transferred to Arthrinium by Wang et al. (2017) before being reclassified under Apiospora based on DNA sequence data (Pintos and Alvarado 2021). Our phylogenetic analyses indicate that A. vietnamensis, A. euphorbiae, A. malaysiana, and A. magnispora form a single clade without evident genetic divergence (Fig. 2). Pairwise sequence comparisons further reveal negligible nucleotide differences among the type strains of these species. Morphologically, A. euphorbiae (Ellis 1965; Tian et al. 2021), A. malaysiana (Crous & Groenewald, 2013), and A. vietnamensis (Wang et al. 2017) share nearly identical conidial dimensions (4–5.5 × 3–4 µm, 5–6 × 3–4 µm, and 5–6 × 3–4 µm, respectively). In contrast, A. magnispora produces distinctly larger conidia (20–35 × 15–25 µm; Zhao et al. 2023), which may represent host-associated morphological plasticity rather than a taxonomically meaningful difference. Pairwise sequence comparisons revealed that type strain IMI 99670T of A. vietnamensis differs from A. malaysiana (CBS 102053T) by 0.18% in ITS (1/570 bp, no gaps), 0.12% in LSU (1/812 bp, one gap), and no differences in tub2. Comparisons with A. euphorbiae (IMI 285638b) showed divergences of 0.18% in ITS (1/520 bp, no gaps), 0.31% in LSU (1/318 bp), and no difference in tub2. Comparisons with A. magnispora (ZHKUCC 22-0001T) showed divergences of 0.18% in ITS (1/570 bp, no gaps), 0.12% in LSU (1/812 bp, one gap), and no differences in tub2. Given the absence of genetic differentiation and the strong morphological congruence among these taxa, we consider A. euphorbiae, A. malaysiana, and A. magnispora to be conspecific with A. vietnamensis, and thus propose them as its synonyms.
Nigrospora Zimm., Centbl. Bakt. ParasitKde, Abt. II 8: 220 (1902).
Nigrospora chinensis
Mei Wang & L. Cai, in Wang, Liu, et al., Persoonia 39: 129. 2017.
57ED8FD0-92F9-58B0-86A4-EB51185AA911
820732
Figure 13.
Nigrospora chinensis (GUCC 25-0120) A Appearance of the fungus on leaves of Juglans regia; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H Close up of SNA culture with conidia; I–P Conidia with conidiogenous cells on SNA. Scale bars: 100 μm (H); 10 μm (I–P).
Substrate and distribution.
Found mostly in China on Zanthoxylum bungeanum (Cheng et al. 2025); dragon fruit (Guo et al. 2024); Camellia oleifera (Qin et al. 2021); rice, (Liu et al. 2024c); Camellia sinensis, Musa paradisiaca, Lindera aggregate, Aucuba japonica, Machilus duthiei, Osmanthus sp., Quercus sp., Smilax ocreata, (Wang et al. 2017). And on Eleiodoxa conferta, Thailand (Karimi et al. 2025).
Description.
Associated with leaf spots on Juglans regia. Circular or irregular lesions, ranging from yellowish-brown to dark brown, primarily concentrated along the leaf margins. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 2.5–5.1 µm wide, hyphae, branched, septate, hyaline to pale green, smooth. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5.4–9.5 × 4.8–6.1 µm (x̄ = 7.4 × 5.3 μm, n=30), ampulliform to subglobose, monoblastic, hyaline. Conidia 10.3–13.5 × 8.5–12.8 µm (x̄ = 11.9 × 10.3 μm, n=30), globose to subglobose, solitary, aseptate, black, smooth-walled, rarely longitudinal germ-slit.
Culture characteristics.
Colonies after 7 days at 25 °C: On PDA, colonies reach mm 90 diam., flat, spreading, with regular margins, surface white and reverse pale salmon. On MEA, colonies reach 88–90 mm diam., flat, spreading, with regular margins, surface and reverse white. On CMA, colonies reach 70–72 mm diam., floccose, spreading, with regular margins, surface and reverse white. On SNA, colonies reach 85–87 mm diam., flat, spreading, with regular margins, surface and reverse white. On WA, colonies reach 53–57 mm diam., flat, spreading, with erose margin, surface and reverse white. On OA, colonies reach 90 mm diam., flat, cottony, with regular margins, surface pale mouse grey and reverse pale salmon. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Guizhou Province, Tongren City, Dejiang County, on the diseased leaves of Juglans regia with spots, 5 May 2024, M.T. Zou, HGUP 25-0060; living cultures GUCC 25-0120 and GUCC 25-0121.
Notes.
Our phylogenetic analysis revealed that the two strains GUCC 25-0120 and GUCC 25-0121 clustered together with N. chinensis, showing no significant genetic distance and high support rates (100% ML/1.0 BI; Fig. 3). Morphological comparison indicated that the characteristics of our strains are consistent with those reported for N. chinensis with the size of conidia and conidiogenous cells (Wang et al. 2017). This species is reported for the first time on the leaves of Juglans regia.
Nigrospora endophytica
A.C.Q. Brito & A.R. Machado, in Brito, Mello, et al., Mycological Progress 22(6, no. 37): 5. 2023.
B5809F7E-410E-5E2B-933D-EE12653B7682
845749
Figure 14.
Nigrospora endophytica (GUCC 25-0122) A Appearance of the fungus on leaves of Juglans regia; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H Close up of SNA culture with conidia; I–L Conidia with conidiogenous cells on SNA. Scale bars: 100 μm (H); 10 μm (I–L).
Substrate and distribution.
Manihot esculenta, Brazil (de Queiroz Brito et al. 2023).
Description.
Associated with leaf spots on Juglans regia. Circular or irregular lesions, ranging from yellowish-brown to dark brown. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 1.5–4 µm wide, hyphae, ranched, sometimes coiled, septate, hyaline or pale green to dark brown. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 7.4–9.1 × 4.9–7.8 µm (x̄ = 8.2 × 6.2 μm, n=30), ampulliform to subglobose, solitary, monoblastic, pale green. Conidia 11.1–16.3 × 7.4–13.4 µm (x̄ = 13.6 × 10.9 μm, n=30), subglobose, solitary, aseptate, pale brown to black, smooth.
Culture characteristics.
Colonies for 7 days at 25 °C: On PDA, colonies reach 90 mm diam., flat, spreading, with regular margins, surface center fuscous black and margin white, reverse center black and margin pale salmon. On MEA, colonies reach 90 mm diam., flat, spreading, with regular margins, surface center and reverse vinaceous grey and margin greyish lilac. On CMA, colonies reach 90 mm diam., flat spreading, with regular margins, surface, and reverse greyish lilac. On SNA, colonies reaching 90 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse pale vinaceous grey. On WA, colonies reach 40–42 mm diam., flat, aerial mycelia scant, with erose margin, surface and reverse greyish. On OA, colonies reach 90 mm diam., flat, spreading, with regular margins, surface greyish lilac and reverse center black and margin pale grey. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Yunnan Province, Lincang City, on the diseased leaves of Juglans regia with spots, 21 June 2024, M.T. Zou, HGUP 25-0061; GUCC 25-0122 and GUCC 25-0123.
Notes.
Our phylogenetic analysis indicated that strains GUCC 25-0122 and GUCC 25-0123 clustered together with N. endophytica (strains A.R.M 973 and A.R.M 687) with statistical support (96% ML/0.69 BI) (Fig. 3). Comparison of nucleotide differences between GUCC 25-0122 and N. endophytica (A.R.M 973) revealed minor variations: 0.83% in ITS (4/481bp, including one gap), 1.05% in tef1-α (5/478 bp, including one gap), and 0.05% in tub2 (2/405 bp, without gaps). Although slight sequence differences were observed, the morphological characteristics of our strains are closely similar to those of N. endophytica (de Queiroz Brito et al. 2023). Therefore, strains GUCC 25-0122 and GUCC 25-0123 are identified as N. endophytica. Previously, N. endophytica was reported only as an endophyte from the stem tissue of Manihot esculenta (de Queiroz Brito et al. 2023). This study represents the first record of this species from J. regia leaves, indicating a new host association.
Nigrospora neosaccharicola
M. T. Zou, X. C. Wang, K.D. Hyde & Yong Wang bis sp. nov.
41880857-C8AB-5139-A19A-60DB52983537
860114
Figure 15.
Nigrospora neosaccharicola (GUCC 25-0124) A Appearance of the fungus on leaves of Juglans regia; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H Close up of SNA culture with conidia; I–M Conidia with conidiogenous cells on SNA. Scale bars: 100 μm (H); 10 μm (I–M).
Etymology.
The species was named based on morphological similarity to Nigrospora saccharicola.
Holotype.
China • Yunnan Province, Chuxiong City, on the diseased leaves of Juglans regia with spots, 26 March 2024, M.T. Zou, HGUP 25-0010 (holotype), ex-type GUCC 25-0124.
Description.
Associated with leaf spots on Juglans regia. Leaves lose their green coloration, developing nearly circular lesions that appear yellowish-brown with a surrounding yellow halo. In later stages, the center of the lesions turns pale yellow. Small spots merge into larger ones, eventually causing leaf death. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 3.2–5.8 µm wide, branched, septate, hyaline to dark brown, smooth. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4.5–10.7 × 5.4–8.9 µm (x̄ = 7.4 × 6.7 μm, n=30), subglobose to pot-shaped, aseptate, pale brown. Conidia 11.4–16.7 × 8.1–13.4 µm (x̄ = 13.7 × 10.6 μm, n=30), globose to subglobose, solitary, aseptate, brown to black, smooth-walled.
Culture characteristics.
Colonies after 7 days at 25 °C: On PDA, colonies reach 90 mm diam., flat, dense aerial mycelia, with regular margins, surface pale olivaceous gray, reverse center black, and margin pale flesh. On MEA, colonies reach 90 mm in diam., flat, spreading, and dense, with regular margins; surface dark mouse grey and reverse umber. On CMA, colonies reach 90 mm diam., flat, spreading, with regular margins, surface, and reverse center grey and margin greyish lilac. On SNA, colonies reach 90 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse pale vinaceous grey. On WA, colonies reach 77–79 mm diam., flat, aerial mycelia scant, with erose margin, surface and reverse greyish. On OA, colonies reach 90 mm diam., cottony, dense aerial mycelia, with regular margins, surface mouse grey, reverse center dark mouse grey, and margin saffron. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Yunnan Province, Chuxiong City, on the diseased leaves of Juglans regia with spots, 26 March 2024, M.T. Zou, HGUP 25-0010 (holotype), GUCC 25-0124, (ex-type), GUCC 25-0125, GUCC 25-0126, GUCC 25-0127, GUCC 25-0128 and GUCC 25-0129.
Notes.
In the phylogenetic analyses (Fig. 3), six strains (GUCC 25-0124, GUCC 25-0125, GUCC 25-0126, GUCC 25-0127, GUCC 25-0128, and GUCC 25-0129) formed a sister clade to N. saccharicola (type strain CGMCC 3.19362) with strong statistical support (100% ML/1.0 BI). The comparison of nucleotide differences between strain GUCC 25-0124 and the type strain of N. saccharicola revealed clear sequence divergence: 0.31% in ITS (1/324 bp, no gaps), 3.32% in tub2 (12/361 bp, including seven gaps), and 5.58% in tef1-α (24/430 bp, including three gaps). Morphologically, our isolates differ slightly from N. saccharicola in both conidial and conidiogenous cell dimensions. Nigrospora neosaccharicola produces smaller conidia (mean = 13.7 × 10.6 μm vs. 15.24 × 11.69 μm) and narrower conidiogenous cells (mean = 7.4 × 6.7 μm vs. 8.65 × 6.8 μm) (Raza et al. 2019). Furthermore, sterile (elongated) cells, which are present in N. saccharicola, were not observed in our isolates. While N. saccharicola was originally described from Saccharum officinarum (Raza et al. 2019), our isolates were obtained from leaves of Juglans regia.
Nigrospora osmanthi
Mei Wang & L. Cai, Persoonia 39: 135 (2017).
A2EDC710-48D0-56E4-A238-21197035B59E
820736
Figure 16.
Nigrospora osmanthi (GUCC 25-0130). A Appearance of the fungus on leaves of Juglans regia; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H Close up of SNA culture with conidia; I–M Conidia with conidiogenous cells on SNA; N Coiled hyphae on SNA. Scale bars: 100 μm (H); 10 μm (I–N).
Substrate and distribution.
Ficus pandurata, China (Liu et al. 2019); Hedera nepalensis, China (Wang et al. 2017); Water Lettuce, China (Lin et al. 2023); Stenotaphrum secundatum China (Mei et al. 2019); Orthosiphon stamineus, Malaysia (Ismail et al. 2022); Artemisia argyi, China (Yang et al. 2025); Cirsium setosum, Phyllostachys nigra, Phragmites australis, Rosa chinensis, China (Hao et al. 2020). Other distributions such as Colombia, Croatia, Iran, Malaysia, Saudi Arabia, South Africa, and United States recorded in GBIF.
Description.
Associated with leaf spots of Juglans regia. Circular or irregular lesions, ranging from yellowish-brown to dark brown with yellow halos. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 3.5–7 µm wide, branched, septate, hyaline to dark brown, smooth. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5.8–8.7 × 4.1–6.6 µm (x̄ = 7.0 × 5.2 μm, n=30), subglobose, monoblastic, pale green to brown. Conidia 10.2–12.3 × 8.3–10.9 µm (x̄ = 11.4 × 9.5 μm, n=30), globose to subglobose, aseptate, solitary, brown to black, smooth, shiny.
Culture characteristics.
Colonies after 7 days at 25 °C: On PDA, colonies reach 90 mm diam., flat, spreading, dense aerial mycelia, with regular margins, surface and reverse white. On MEA, colonies reach 90 mm diam., flat, spreading, dense aerial mycelia, regular margins, surface and reverse pale vinaceous grey. On CMA, colonies reach 90 mm diam., flat, spreading, regular margins, surface and reverse pale vinaceous grey. On SNA, colonies reach 88–90 mm diam., flat, spreading, aerial mycelia scant, regular margins filamentous, surface and reverse pale vinaceous grey. On WA, colonies reach 52–55 mm diam., flat, spreading, with erose and irregular margin, surface and reverse pale vinaceous grey. On OA, colonies reach 90 mm diam., flat, spreading, dense aerial mycelia, with regular margins, surface mouse grey and reverse center dark mouse grey and margin saffron. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Yunnan Province, Chuxiong City, on the diseased leaves of Juglans regia with spots, 23 March 2024, M.T. Zou, HGUP 25-0062; GUCC 25-0130, GUCC 25-0131, GUCC 25-0132 and GUCC 25-0133.
Notes.
Nigrospora osmanthi was first observed on Osmanthus sp. in Jiangxi Province, China (Wang et al. 2017). In this study, four strains (GUCC 25-0130, GUCC 25-0131, GUCC 25-0132, and GUCC 25-0133) were isolated from leaf spots of J. regia. Phylogenetic analysis revealed that these two strains clustered with the type strain of N. osmanthi (CGMCC 3.18126) and strain LC4467, showing no significant genetic divergence. Morphologically, our isolates exhibited characteristics consistent with those described for N. osmanthi (Wang et al. 2017). Based on the combined morphological and phylogenetic evidence, the four strains are identified as N. osmanthi. This represents the first record of N. osmanthi from J. regia. However, further pathogenicity tests are required to confirm the pathogenic causes of leaf spot symptoms.
Nigrospora sphaerica
(Sacc.) E.W. Mason, Trans. Br. Mycol. Soc. 12(2-3): 158 (1927).
8708EAD7-9D28-5539-BE6C-8941DF9AD3F8
254776
Figure 17.
Nigrospora sphaerica (GUCC 25-0134). A Appearance of the fungus on leaves of Juglans regia; B–G Upper view and reverse view of culture on PDA, MEA, CMA, SNA, WA, and OA after 7 days; H Close up of SNA culture with conidia; I–M Conidia with conidiogenous cells on SNA. Scale bars: 100 μm (H); 10 μm (I–M).
Substrate and distribution.
Pathogens with a wide host range, primarily causing leaf spot or leaf blight.
Description.
Associated with leaf spots of Juglans regia. Lesions irregular in shape and dark brown in color. Sexual morph: Not observed. Asexual morph: On SNA, Hyphae 2.5–5 µm in diam., branched, septate, contracted at septum, hyaline to dark brown, smooth. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 6.0–10.5 × 5.0–8.1 µm (x̄ = 7.8 × 6.6 μm, n=30), subglobose, monoblastic, pale brown. Conidia 11.8–15.8 × 9.9–13.7 µm (x̄ = 14.1 × 11.6 μm, n=30), globose to subglobose, solitary, aseptate, black, smooth.
Culture characteristics.
Colonies after 7 days at 25 °C: On PDA, colonies reach 90 mm diam., flat, spreading, with regular margins, surface white and reverse salmon. On MEA, colonies reach 27–30 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse white. On CMA, colonies reach 56–60 mm diam., cottony, dense aerial mycelia, with regular margins, surface white and reverse pale salmon. On SNA, colonies reach 20–22 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse white. On WA, colonies reach 20–22 mm diam., flat, aerial mycelia scant, regular margins filamentous, surface and reverse white. On OA, colonies reach 73–75 mm diam., flat, spreading, dense aerial mycelia, with regular margins, surface greyish lilac and reverse center black and margin pale salmon. Sporulation was abundant only on SNA after 14 days.
Material examined.
China • Yunnan Province, Lincang City, on the diseased leaves of Juglans regia with spots, 7 May 2024, M.T. Zou, HGUP 25-0063; GUCC 25-0134, GUCC 25-0135, GUCC 25-0136, GUCC 25-0137, GUCC 25-0138 and GUCC 25-0139.
Notes.
In the phylogenetic analyses (Fig. 3), the six strains obtained in this study (GUCC 25-0134, GUCC 25-0135, GUCC 25-0136, GUCC 25-0137, GUCC 25-0138, and GUCC 25-0139) clustered with the N. sphaerica species clade with strong statistical support (100% ML/1.0 BI). Morphologically, our isolates closely resemble N. sphaerica in their overall diagnostic characteristics. The conidia of our strains are comparable in shape but slightly smaller (14.1 × 11.6 µm) than those reported for N. sphaerica (18.22 × 18.22 µm), while the conidiogenous cells also show similar dimensions (7.8 × 6.6 µm vs. 7.97 × 7.97 µm; Wang et al. 2017; Hyde et al. 2024a). This study represents the first record of N. sphaerica isolated from leaves of J. regia.
Discussion
The family Apiosporaceae is widely distributed across terrestrial ecosystems and exhibits broad ecological adaptability, with members recorded from tropical, subtropical, temperate, and boreal regions and colonizing diverse hosts with no clear host specificity (Jiang et al. 2019; Phukhamsakda et al. 2022). The Yunnan–Guizhou Plateau in Southwest China represents a well-known biodiversity hotspot where numerous novel fungi have been discovered in recent years (Wijayawardene et al. 2021), including members of Apiosporaceae, such as A. aseptata, A. dematiacea, A. dicranopteridis, A. globosa (Zhang et al. 2023), A. olivata (Zhang et al. 2024b), and N. weininensis (Liu et al. 2024a). In this study, all specimens were collected from Guizhou and Yunnan provinces, further confirming that the plateau remains an important reservoir of Apiosporaceae diversity.
The multilocus phylogenetic analysis using ITS, LSU, tef1-α, and tub2 sequences confirms the novelty of these taxa. Four single-gene phylogenetic trees were constructed to evaluate variation in the ITS and protein-coding genes (tef1-α and tub2). In the ITS phylogeny (Suppl. material 1: fig. S1), several Apiospora species exhibited low genetic divergence, whereas this phenomenon is less frequent in tub2 and tef1-α phylogenies (Suppl. material 1: figs S3, S4). These results indicate that tub2 and tef1-α provide higher resolution for species delimitation than ITS, consistent with Monkai et al. (2022). Similarly, in a study of Hypoxylaceae, another family within the Xylariomycetidae to which Apiospora belongs, Stadler et al. (2020) demonstrated that ITS alone is insufficient for delimiting taxa, as ITS and other rDNA markers exhibit polymorphism both between and within species. Therefore, species identification in Apiospora should rely on multigene phylogenetic frameworks, with a particular emphasis on tub2 and tef1-α genes, while ITS is primarily suitable for genus-level identification.
Multigene phylogenetic analyses demonstrate that A. mediterranea and A. hispanica are conspecific, forming a single clade with no genetic divergence and showing minimal morphological differentiation (Monkai et al. 2022; Liao et al. 2023). Similarly, A. euphorbiae, A. magnispora, and A. malaysiana should be treated as synonyms of A. vietnamensis, as they cluster together with negligible nucleotide differences and minimal morphological variation. Phylogenetic analyses in Nigrospora revealed that N. sphaerica is divided into two lineages, one of which includes only strains LC2839 and LC2840. The type strain lacks molecular data, and the sequences provided by Wang et al. (2017) indicate that LC2840 forms a separate branch from the main N. sphaerica clade. Comparative sequence analyses reveal differences of 1.94% in tub2 (8/412 bp, five gaps) and 2.55% in tef1-α (12/470 bp, no gap) between LC2840 and LC7295. BLAST analyses further indicate that these strains share <99% identity with known Nigrospora sequences. Therefore, resequencing of LC2840 and LC2839 is recommended to determine whether these strains represent a distinct taxon or belong to N. sphaericasensu stricto.
To investigate spore production in Apiospora and Nigrospora, six commonly used media (PDA, SNA, WA, OA, CMA, and MEA) were evaluated, and conidia production was examined after 14 days. Apiospora produced conidia readily on WA and SNA, consistent with the report by Zhang et al. (2023). Strains of A. setariae failed to produce conidia on any medium, even after 30 days. For Nigrospora, conidia production occurred only on SNA medium after 14 days, though at low levels. In addition, these observations suggest that certain cultural characteristics may serve as diagnostic features for species identification. For example, A. huaxiensis and A. setariae produce red pigments on PDA. Similar pigment production has been reported in A. dehongensis (Han et al. 2024). Therefore, red pigment production on PDA may represent a useful taxonomic character.
The four new species of Apiospora were isolated from symptomatic bamboo leaves or stems, supporting a strong association with Poaceae, particularly bamboo hosts (Han et al. 2024; Liu et al. 2024b). Two additional taxa, A. locuta-pollinis and A. setariae were recorded from new host species. Strains of A. locuta-pollinis, previously reported in hive-stored pollen (Zhao et al. 2018), Aristolochia debilis (Chen et al. 2021), bamboo (Monkai et al. 2022), Musa sp. (Samarakoon et al. 2024), and grass (Gao et al. 2025), were isolated from maize leaves. Moreover, A. setariae, previously known from dead culms of Setaria viridis (Jiang and Tian 2021), is recorded here for the first time from bamboo. All Nigrospora isolates in this research were obtained from walnut. To date, only one species, N. yunnanensis, has been reported from walnut (Zou et al. 2024). Analyses of host preferences and species distributions of Apiospora and Nigrospora indicate that for Apiospora, the primary host family is Poaceae, with bamboo species constituting the largest proportion, consistent with Monkai et al. (2022). For the species distributions of this genus, Monkai et al. (2022) identified China as the country with the highest number of recorded species for this genus. Our findings, however, indicate that the United States has the highest number of recorded species, followed by the United Kingdom and China. This discrepancy likely reflects differences between databases, as GBIF was used, whereas Monkai et al. (2022) used USDA data, which contain fewer records and are primarily disease-related. For Nigrospora, GBIF data indicate that Poaceae is the most common host family, followed by Fabaceae and Cyperaceae. In terms of geographic distribution, China has the highest number of hosts records, followed by India and the United States.
Overall, this study significantly expands current knowledge of Apiosporaceae diversity in Southwest China, clarifies several taxonomic relationships within Apiospora and Nigrospora, and highlights the Yunnan–Guizhou Plateau as a rich reservoir of new taxa. The discovery of five new species and six new host species records highlights the ecological adaptability of Apiosporaceae and underscores the importance of integrating morphology, molecular data, and host associations for accurate species identification and improved understanding of their taxonomy and evolution.
Supplementary Material
Citation
Wang X-C, Zhang R-N, Norphanphoun C, Wang S-B, Zou M-T, Guo S-Q, Sun J-E, Wang J-P, Wijayawardene NN, Hyde KD, Wang Y, Liu F-Q (2026) Novel species and new host records of Apiospora and Nigrospora (Amphisphaeriales, Apiosporaceae) from Yunnan-Guizhou Plateau, China. IMA Fungus 17: e177246. https://doi.org/10.3897/imafungus.17.177246
Funding Statement
This research is supported by the following projects: National Key Research and Development Program of China (No.2024YFE0214300), the Guizhou Plant Bacteria and Biological Control Science and Technology Innovation Talent Team Development (Qian Ke He Talent-BQW[2025]003), the National Natural Science Foundation of China (No. 31972222), the Program of Introducing Talents of Discipline to Universities of China (111 Program, D20023), and the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia.
Footnotes
Xing-Chang Wang and Rui-Nan Zhang contributed equally to this work.
Contributor Information
Yong Wang, Email: yongwangbis@aliyun.com.
Feng-Quan Liu, Email: fqliu20011@sina.com.
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 research is supported by the following projects: National Key Research and Development Program of China (No.2024YFE0214300), the Guizhou Plant Bacteria and Biological Control Science and Technology Innovation Talent Team Development (Qian Ke He Talent-BQW[2025]003), the National Natural Science Foundation of China (No. 31972222), the Program of Introducing Talents of Discipline to Universities of China (111 Program, D20023), and the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia.
Author contributions
Conceptualization: Xing-Chang Wang, Yong Wang; Data curation: Meng-Ting Zou, Shi-Qi Guo, Jia-Ping Wang; Formal analysis: Xing-Chang Wang, Rui-Nan Zhang, Shi-Qi Guo, Chada Norphanphoun, Yong Wang; Funding acquisition: Feng-Quan Liu, Yong Wang, Kevin D. Hyde; Investigation: Xing-Chang Wang, Meng-Ting Zou, Shi-Qi Guo, Si-Bo Wang; Project administration: Yong Wang, Feng-Quan Liu; Resources: Yong Wang, Feng-Quan Liu; Supervision: Yong Wang, Feng-Quan Liu; Validation: Xing-Chang Wang, Si-Bo Wang, Jing-E Sun; Visualization: Xing-Chang Wang, Jing-E Sun, Si-Bo Wang, Yong Wang; Writing – original draft: Xing-Chang Wang, Chada Norphanphoun, Shi-Qi Guo; Writing – review & editing: Yong Wang, Kevin D. Hyde, Nalin N. Wijayawardene, Feng-Quan Liu
All authors have edited the manuscript and approved to publish the research findings. This research has not been published and is not under consideration for publication elsewhere.
Author ORCIDs
Xing-Chang Wang https://orcid.org/0009-0008-4913-5316
Rui-Nan Zhang https://orcid.org/0009-0009-6010-9570
Chada Norphanphoun https://orcid.org/0000-0002-5756-7206
Si-Bo Wang https://orcid.org/0009-0004-0060-3247
Meng-Ting Zou https://orcid.org/0009-0005-5564-4131
Shi-Qi Guo https://orcid.org/0009-0002-4997-2276
Jing-E Sun https://orcid.org/0000-0002-5226-5743
Jia-Ping Wang https://orcid.org/0009-0002-7558-4760
Nalin N. Wijayawardene https://orcid.org/0000-0003-0522-5498
Kevin D. Hyde https://orcid.org/0000-0002-2191-0762
Yong Wang https://orcid.org/0000-0003-3831-2117
Feng-Quan Liu https://orcid.org/0000-0001-9325-1500
Data availability
All data generated or analyzed during this study are included in this published article or in the supplementary materials. All sequences during this study have been submitted to GenBank.
Supplementary materials
Single gene trees for Apiospora and Nigrospora
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
zip
Explanation note
fig. S1. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on ITS for Apiospora. fig. S2. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on LSU for Apiospora. fig. S3. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tef1-α for Apiospora. fig. S4. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tub2 for Apiospora. fig. S5. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on ITS for Nigrospora. fig. S6. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tef1-α for Nigrospora. fig. S7. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tub2 for Nigrospora.
Suppl. data 1-7: the sequences after mafft for Apiospora and Nigrospora
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
zip
Explanation note
suppl. data 1. The ITS sequences for Apiospora after Mafft. suppl. data 2. The LSU sequences for Apiospora after Mafft. suppl. data 3. The tef1-α sequences for Apiospora after Mafft. suppl. data 4. The tub2 sequences for Apiospora after Mafft. suppl. data 5. The ITS sequences for Nigrospora after Mafft. suppl. data 6. The tef1-α sequences for Nigrospora after Mafft. suppl. data 7. The tub2 sequences for Nigrospora after Mafft.
Primer pairs and amplification procedures used in this study, the best models of BI and ML used for each gene in the phylogenetic tree, and the records of Apiospora and Nigrospora in the GBIF database
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
xlsx
Explanation note
table SS1. Primer pairs and amplification procedures used in this study. table S2. The best models of BI and ML used for each gene in the phylogenetic tree. table S3. Records of Apiospora in GBIF. table S4. Records of Nigrospora in GBIF.
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Single gene trees for Apiospora and Nigrospora
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
zip
Explanation note
fig. S1. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on ITS for Apiospora. fig. S2. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on LSU for Apiospora. fig. S3. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tef1-α for Apiospora. fig. S4. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tub2 for Apiospora. fig. S5. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on ITS for Nigrospora. fig. S6. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tef1-α for Nigrospora. fig. S7. Phylogenetic trees constructed using Maximum likelihood (ML) analyses based on tub2 for Nigrospora.
Suppl. data 1-7: the sequences after mafft for Apiospora and Nigrospora
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
zip
Explanation note
suppl. data 1. The ITS sequences for Apiospora after Mafft. suppl. data 2. The LSU sequences for Apiospora after Mafft. suppl. data 3. The tef1-α sequences for Apiospora after Mafft. suppl. data 4. The tub2 sequences for Apiospora after Mafft. suppl. data 5. The ITS sequences for Nigrospora after Mafft. suppl. data 6. The tef1-α sequences for Nigrospora after Mafft. suppl. data 7. The tub2 sequences for Nigrospora after Mafft.
Primer pairs and amplification procedures used in this study, the best models of BI and ML used for each gene in the phylogenetic tree, and the records of Apiospora and Nigrospora in the GBIF database
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.
Xing-Chang Wang, Rui-Nan Zhang, Chada Norphanphoun, Si-Bo Wang, Meng-Ting Zou, Shi-Qi Guo, Jing-E Sun, Jia-Ping Wang, Nalin N. Wijayawardene, Kevin D. Hyde, Yong Wang, Feng-Quan Liu
Data type
xlsx
Explanation note
table SS1. Primer pairs and amplification procedures used in this study. table S2. The best models of BI and ML used for each gene in the phylogenetic tree. table S3. Records of Apiospora in GBIF. table S4. Records of Nigrospora in GBIF.
Data Availability Statement
All data generated or analyzed during this study are included in this published article or in the supplementary materials. All sequences during this study have been submitted to GenBank.