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23 May 2026

15 Pages

Untapped Diversity of Termite-Associated Ophiocordyceps and a New Species from China

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1
College of Agronomy, Chuxiong Normal University, Chuxiong 675000, China
2
Key Laboratory of East China Plant Conservation and Utilization, National Forestry and Grassland Administration, Shanghai Chenshan Botanical Garden, Shanghai 201602, China
*
Author to whom correspondence should be addressed.
Diversity2026, 18(6), 313;https://doi.org/10.3390/d18060313
This article belongs to the Section Phylogeny and Evolution
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Abstract

Termite-associated Ophiocordyceps species remain understudied despite the high diversity of the genus. Here we describe Ophiocordyceps minuta (holotype CXAC 0026) from termites (Termitidae, Macrotermitinae) collected in Yunnan, China. Phylogenetic analyses based on nrLSU (nuclear large subunit ribosomal RNA), tef1-α (translation elongation factor 1-alpha), and rpb2 (RNA polymerase II second largest subunit) sequences resolve this fungus as a distinct lineage. Notably, deep genetic divergence (65 bp in tef1-α) between the two ex-type strains of the allied O. fusiformis (BCC 93025 and BCC 93026) reveals cryptic diversity within that nominal species. Beyond the new species, a morphological assessment of termite-associated Ophiocordyceps indicates that perithecial immersion status, ascospore morphology, and conidial features are the most taxonomically valuable characters for this ecological group. These findings expand the known diversity in southwest China and underscore the importance of integrating phylogenetic data with key morphological traits for species delimitation in under-explored habitats.

1. Introduction

The genus Ophiocordyceps Petch (Ophiocordycipitaceae, Hypocreales) ranks among the most species-rich lineages of entomopathogenic fungi, with over 370 accepted names distributed globally (www.indexfungorum.org, accessed on 15 March 2026). Unlike their close relatives in Cordyceps sensu stricto, which typically produce brightly coloured, fleshy stromata, Ophiocordyceps species are characterized by dark-pigmented, wiry to pliant stromata and are associated with anamorphic genera including Hirsutella, Hymenostilbe, Paraisaria, and Syngliocladium [1,2,3]. Members of this genus exploit an exceptionally broad range of insect hosts, attacking various life stages across Blattodea, Coleoptera, Diptera, Hemiptera, Hymenoptera, Lepidoptera, Megaloptera, Neuroptera and Odonata [3,4,5,6,7]. However, the association with Blattodea is not restricted to termites; several Ophiocordyceps species, including the type species O. blattae, as well as O. asiatica, O. cf. blattae, and O. salganeicola, have been confirmed from non-termite Blattodea [8,9]. This ecological diversity is matched by morphological plasticity: perithecia may be superficial, immersed, or partially embedded, while ascospores either remain intact as multiseptate units or dissociate into individual part-spores [2,10].
Despite this overall diversity, the distribution of Ophiocordyceps species across host groups is markedly uneven [11,12,13]. Certain host associations, particularly with ants and beetle larvae, have yielded numerous described species, while others remain poorly documented [14,15]. Termites (Blattodea: Termitidae) represent one such understudied host group [16]. Termites were historically treated as a separate order, Isoptera, but molecular phylogenetic studies have firmly placed them within Blattodea [17], a placement further corroborated by recent genomic-scale analyses [18]. Termitidae is the most speciose termite family, comprising approximately 2000 described species [19]. As eusocial insects inhabiting subterranean nests in tropical and subtropical ecosystems worldwide, termites play critical roles in decomposition and nutrient cycling [20]. Their constant exposure to soil-borne microorganisms creates opportunities for fungal pathogens, yet the diversity of termite-associated Ophiocordyceps has only recently begun to receive sustained attention [8,21,22,23].
Historical records of termite-pathogenic Ophiocordyceps were limited to three species described over a span of six decades: O. bispora from East Africa, O. koningsbergeri from Java, and O. octospora from Mexico [24,25,26]. Following these early reports, progress remained slow until intensive surveys in Thailand during the past decade uncovered 11 additional species from forests [8,27]. The remaining species have been documented from other East and Southeast Asian localities: O. globiperitheciata, O. longistipes, O. ovatospora and O. taiwanensis from China, and O. puluongensis from Vietnam [21,22,23,28]. Thus, a total of 20 Ophiocordyceps species have been reported from termite hosts worldwide to date, not including the new species described in the present study [2,8,21,23,24,25,26,29]. These discoveries demonstrated that termite-pathogenic lineages harbor substantially greater diversity than previously recognized and suggested that similar surveys in other under-explored regions would likely yield additional taxa.
Southwest China, particularly Yunnan Province, has emerged as a promising area for discovering novel Cordyceps sensu lato species [30,31]. The region’s complex topography, elevational gradients, and high forest cover support diverse insect communities and their associated fungal pathogens [32,33]. Recent investigations have documented multiple new Ophiocordyceps taxa from Yunnan across various insect hosts, including two termite-associated species described in the past two years [21,22]. These findings indicate that the termite-pathogenic Ophiocordyceps diversity in this region remains incompletely sampled, with additional species awaiting discovery through continued field efforts. In the present study, we describe the new termite-associated species O. minuta using morphological and multigene phylogenetic evidence, reassess the key morphological characters for delimiting termite-associated Ophiocordyceps, and discuss the group’s diversity in southwest China.

2. Materials and Methods

2.1. Specimen Collection

Fungal specimens were collected from ground-dwelling termites (Termitidae, Macrotermitinae) in Taiyanghe National Nature Reserve, Yunnan Province, China. Specimens were carefully removed to preserve host integrity, placed in sterile plastic containers, and transported to the laboratory. Upon arrival, they were examined under a stereomicroscope (OLYMPUS SZ61, Olympus Corporation, Tokyo, Japan), cleaned of surface debris, assigned accession numbers CXAC 0026–0027, and air-dried. Hosts were identified using morphological characters and with guidance from termite taxonomists; finer identification was precluded by the condition of the fungus-infected specimens. Voucher specimens are deposited at the College of Agronomy Herbarium (CXAC), Chuxiong Normal University, China, for taxonomic identification and future studies.

2.2. Morphological Characterization

Specimen data, including host association and geographic origin, were documented. Sexual structures (perithecia, asci, ascospores, part-spores) were mounted in lactophenol cotton blue and measured under an Olympus BX53 compound microscope (Olympus Corporation, Tokyo, Japan). For each structure, 20–50 measurements were recorded, and the range was calculated.

2.3. Extraction of DNA, Polymerase Chain Reaction (PCR), and Molecular Sequencing

Genomic DNA was extracted from fungal mycelia using a commercial plant DNA isolation kit (FOREGENE, Chengdu, China) and amplified via polymerase chain reaction (PCR) for three genetic regions: nrLSU, tef1-α, and rpb2, with primers as following: the LSU region was amplified with the primer pair LR0R/LR5 [34,35], and the tef1-α and rpb2 genes using the primer sets EF1α-EF/EF1α-ER [2,36] and RPB2-5′F/RPB2-5′R [2,36] respectively. Each 25 μL PCR contained 2.5 μL of 10× buffer (2 mM Mg2+; Transgen Biotech, Beijing, China), 0.25 μL of Taq polymerase, 2 μL of dNTPs (2.5 mM each), 1 μL of DNA template (~500 ng/μL), 1 μL of each primer (10 μM), and 17.25 μL of deionized water. Amplification was conducted in a BIO-RAD T100™ Thermal Cycler following the gene-specific thermal cycling protocols detailed in Table 1. Successful amplification was verified by gel electrophoresis; the resulting amplicons were then purified prior to bidirectional Sanger sequencing at the Beijing Genomics Institute (Shenzhen, China) with the same PCR primers.
Table 1. PCR cycling conditions for each gene.

2.4. Phylogenetic Analyses

To establish the phylogenetic placement of our taxa, we analyzed a three-gene matrix (nrLSU, tef1-α, and rpb2). Newly obtained sequences were confirmed via BLAST searches and aligned with reference sequences from GenBank (Table 2) using MAFFT v7.526 (L-INS-i strategy) (http://mafft.cbrc.jp/alignment/server/, accessed 15 March 2026) and refined manually in BioEdit v7.7.1. These alignments were concatenated into a supermatrix with FASconCAT-G v1.06 [37]. The partition homogeneity test in PAUP v5.0 (1000 reps, p > 0.01) revealed no significant conflict between gene partitions. Optimal partitioning schemes and models of evolution were selected under the BIC criterion using PartitionFinder2 v2.0.0 (greedy algorithm) [38]. Phylogenetic relationships were then inferred using two complementary methods. First, a Maximum Likelihood (ML) analysis was performed in IQ-TREE v3.0.1 under the optimal partition scheme (GTR + G + I model), with nodal support assessed via 1000 ultrafast bootstrap replicates [39]. Second, Bayesian Inference (BI) was conducted in MrBayes v3.2.7 using the GTR + G + I model (selected by MrModeltest v2.2) [40]. MCMC chains were run for 5 million generations, sampling every 1000 generations [41]. Convergence (ESS > 200) was verified in Tracer v1.7.2, after which the first 25% of samples were discarded as burn-in [42]. The resulting trees were visualized and annotated in FigTree v1.4.4 (https://tree.bio.ed.ac.uk/software/figtree/ (accessed on 18 may 2026), with the final artwork prepared in Adobe Illustrator CS6 according to the standards of Xie et al. [43].
Table 2. Specimen information and GenBank accession numbers for sequences used in this study.

3. Results

3.1. Phylogenetic Analyses

Phylogenetic analyses of Ophiocordyceps were based on a dataset of 92 sequences (Table 2). Tolypocladium inflatum OSC 71235 and T. ophioglossoides CBS 100239 were used as the outgroups. The final concatenated matrix comprised 2927 characters (including gaps), with three partitions: nrLSU 908 bp, tef1-α 957 bp, and rpb2 1062 bp. Both Bayesian inference (BI) and maximum likelihood (ML) analyses produced congruent trees, recovering Ophiocordyceps as distinct, well-supported clades (Figure 1). The new species Ophiocordyceps minuta formed a distinct lineage within a well-supported clade, clearly separated from its close relatives (Figure 1).
Figure 1. Molecular phylogenetic analyses using the ML and BI based on combined nrLSU, tef1-α, and rpb2 sequence data. Statistical support values (BS ≥ 80% and PP ≥ 0.80) are shown at the nodes for ML bootstrap support (BS) and BI posterior probabilities (PP). Tolypocladium inflatum OSC 71235 and T. ophioglossoides CBS 100239 were used as outgroup taxa. Species in red type are those analyzed in this study. The scale bar represents the expected number of changes per site.

3.2. Taxonomy

Ophiocordyceps minuta Q.Y. Dong & C. D. Xu, sp. nov.
Figure 2
Figure 2. Morphological features of Ophiocordyceps minuta (holotype CXAC 0026). (A). Habitat; (B,C). Fungus on termite; (D–F). Ascospores; (G). Perithecia; (H–J). Ascospores. Scale bars: (B,C) = 10 mm; (D–F,H–J) = 20 µm; (G) = 200 µm.
MycoBank no: 862337
Etymology: minuta = minute, referring to the distinctly small size of the perithecia, a key diagnostic feature that distinguishes this species from its closely related congeners.
Holotype: China, Yunnan Province, Pu’er City, Simao District, Taiyanghe National Nature Reserve, on termite (Termitidae, Macrotermitinae), 15 August 2025, Quanying Dong (holotype CXAC 0026).
Stromata morph: Stromata arising from the segment between the prothorax and mesothorax of termite hosts buried in soil, solitary, simple, cylindrical, flexible, leathery, 6–14 cm long, 0.5–1.0 mm wide, brown to yellowish brown. Fertile parts cylindrical, yellowish brown, 3.5–5.5 cm long, 4–5 mm wide, located on the upper part of the stromata, with a spinous surface and a sterile apical tip 13–28 mm long, 0.5–1.0 mm wide. Perithecia superficial, pale yellow when young, turning brown at maturity, ovoid to flask-shaped, 151.5–221 × 115–188 µm, densely and irregularly scattered over the mid portion of the stromata. Ascospores dimorphic: type I fusiform, 80–92.5 × 5.0–6.5 µm, whole, hyaline, multiseptate, septa 8–14.5 µm long; type II vermiform or lanceolate, 32.5–52.5 × 2.0–4.5 µm.
Asexual morph: Undetermined.
Host: Parasitic on termites buried in soil.
Known distribution: Yunnan Province, China.
Additional specimens examined: China, Yunnan Province, Pu’er City, Simao District, Taiyanghe National Nature Reserve, on termite (Termitidae, Macrotermitinae), 15 August 2025, Quanying Dong (paratype CXAC 0027).
Commentary: The diagnostic morphology of Ophiocordyceps minuta conforms to the generic concept of Ophiocordyceps. The species produces dimorphic ascospores comprising two types: cylindrical, with pointed ends, and vermiform. The species can be distinguished by the following combination of morphological characteristics: perithecia distinctly small (151.5–221 × 115–188 µm), which is the smallest among its closely related congeners (Table 3); host specificity to termites (Termitidae, Macrotermitinae); producing dimorphic ascospores, type I fusiform and multiseptate (80–92.5 × 5.0–6.5 µm), type II vermiform or lanceolate (32.5–52.5 × 2.0–4.5 µm); and asci with conspicuous apical caps (3.5–5.5 μm high, 6.5–8.0 μm wide).
Table 3. Morphological comparison between Ophiocordyceps species associated with termites.
Our multi-locus phylogenetic data (Figure 1) indicate that Ophiocordyceps minuta is related to O. fusiformis Tasan. et al. and O. longistipes Y.B. Wang et al. However, it forms a distinct clade separate from these taxa (Figure 1). Morphologically, O. minuta differs from related species in the following ways. Ophiocordyceps fusiformis, a species described from Thailand, also similar to O. minuta in appearance, has relatively larger perithecia 300–360 × 180–270 µm, longer asci 141–227 × 7–15 µm and shorter ascospores 6–78 × 5–6.5 µm, shorter septa 5–12 (19) µm long [27]. Moreover, in our phylogenetic tree, the two strains of O. fusiformis (BCC 93025 and BCC 93026) did not form a single clade but instead separated into two distinct branches. Comparison of the tef1-α gene sequences between the two strains revealed a 65 bp difference (out of a total length of 903 bp). Ophiocordyceps longistipes can be distinguished from O. minuta by its grayish white to yellowish brown with relatively longer stromata, larger perithecia measuring 390–420 × 295–350 µm; filiform asci, 160–195 × 4.5–6.5 µm; and filiform shorter ascospores 70–85 × 3.5–4.5 µm [21].

3.3. Morphological Characters and Their Taxonomic Significance in Termite-Associated Ophiocordyceps Species

Analysis of termite-associated Ophiocordyceps (Table 3) shows that morphological characters differ in taxonomic usefulness, a pattern seen in other Ophiocordycipitaceae. Host association is uniform (all on termites, Isoptera), so it does not help distinguish species but defines the group as a specialized ecological guild. Stromatal characters provide moderate to high resolution. Stromatal color (orange-brown, brown, yellowish-brown, grayish-white to yellowish-brown, or pale to red-brown) aids species recognition. Stromatal size shows clear extremes: diminutive in O. octospora and O. taiwanensis, elongate in O. asiatica, O. longistipes, O. pseudocommunis, and O. pseudorhizoidea. Multiple stromata (O. bispora: 20–30; O. globiperitheciata: 2–5) are also diagnostic. Perithecial immersion status (superficial, immersed, or pseudo-immersed) reliably separates species, as does perithecial size (large in O. communis and O. pseudocommunis; small in O. ovatospora and O. khokpasiensis). Ascus characters are less informative. Ascospores vary strikingly: most species have filiform, septate, whole ascospores; exceptions include elliptical to ellipsoidal–fusiform, non-disarticulating ascospores (O. bispora, O. octospora) and dimorphic ascospores (O. minuta), the latter being a unique autapomorphy. Asexual morphs (Hirsutella- or Hymenostilbe-like), conidiogenous cells (monophialidic or polyphialidic), and conidia (fusiform, globose, oval, or citriform) offer additional characters. In summary, perithecial immersion status, ascospore architecture, and conidial features are the most taxonomically valuable traits.

4. Discussion

The description of Ophiocordyceps minuta from termite hosts in Yunnan Province adds a new species to the entomopathogenic fungal diversity of Southeast Asia. The morphological and phylogenetic data presented here contribute to ongoing discussions on species delimitation, ascospore variation, and biogeography within the genus Ophiocordyceps.
Taxonomic implications of O. fusiformis strain divergence. In our phylogenetic analysis, the two ex-type strains of O. fusiformis (BCC 93025 and BCC 93026) occupied distinct positions (Figure 1), with a 65 bp divergence in the tef1-α region. Although these strains were originally described as conspecific from Thailand [27], this level of sequence divergence suggests that the current concept of O. fusiformis may encompass multiple species. Confirming species boundaries would require re-examination of type material and additional collections from the type locality. This finding reinforces the importance of including multiple representatives per putative species in phylogenetic studies, as single-strain typification may not fully capture intraspecific variation or detect cryptic diversity [48,60].
Ophiocordyceps minuta produces two morphologically distinct ascospore types. The vermiform Type II spores (32.5–52.5 × 2.0–4.5 µm) are similar in size and shape to microconidia produced by some Hirsutella species associated with termite-pathogenic Ophiocordyceps [8,21,22]. Whether the two ascospore types differ in biological function remains unknown; infection assays comparing their relative infectivity would help clarify their respective roles in the life cycle of this species.
Biogeographic patterns. The discovery of O. minuta in Yunnan adds to the growing number of termite-associated Ophiocordyceps species recorded from the Indochinese Peninsula, including recent reports from Thailand and Vietnam [21,22,27]. Multiple species from this ecological group have now been documented in the Mekong River basin and adjacent areas [6]. The 65 bp divergence observed between the two O. fusiformis strains, both collected in Thailand, further indicates that sequence variation exists within what is currently recognized as a single species. This observation highlights the need for more intensive sampling across the region, particularly collecting multiple specimens per morphospecies and from different host colonies [61].
Ophiocordyceps minuta is a morphologically and phylogenetically distinct species. The genetic divergence detected among O. fusiformis strains and the presence of dimorphic ascospores in O. minuta point to several directions for future research, including taxonomic revision of the O. fusiformis complex and functional studies of ascospore variation in O. minuta. These findings underscore the value of continued exploration of termite-associated Ophiocordyceps in Southeast Asia.

Author Contributions

Conceptualization, Q.-Y.D.; methodology, J.-L.L. and C.-K.L. validation, C.H. and W.-L.X.; formal analysis C.-D.X. and J.Y.; investigation, Q.-Y.D.; writing—review and editing, Q.-Y.D. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Joint Program of Basic Research for Provincial Undergraduate Schools, Yunnan Provincial Department of Science and Technology (202401BA070001-139), the Ph.D. Research Start-Up Project of Chuxiong Normal University (BSQD2307), and the 2025 Self-funded Science and Technology Project of Chuxiong Prefecture (cxzc2025019).

Institutional Review Board Statement

Not applicable for studies not involving humans or animals.

Data Availability Statement

Data are contained within the article.

Conflicts of Interest

The authors declare no conflicts of interest.

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