Published February 17, 2026
| Version v1
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Reappraisal of the phylogenetic relationships of hirsutella-like fungi across Ophiocordycipitaceae and Polycephalomycetaceae (Hypocreales, Sordariomycetes), and the description of seven new species
Authors/Creators
- 1. State Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang, China|Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand|School of Science, Mae Fah Luang University, Chiang Rai, Thailand|Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University, Guiyang, China
- 2. State Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang, China|Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University, Guiyang, China|Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand|School of Science, Mae Fah Luang University, Chiang Rai, Thailand
- 3. State Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang, China|Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University, Guiyang, China
- 4. Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University, Guiyang, China|State Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang, China
- 5. Agricultural College, Guizhou University, Guiyang, China
- 6. College of Life Science and Oceanography, Shenzhen University, Shenzhen, China
- 7. Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand
- 8. School of Science, Mae Fah Luang University, Chiang Rai, Thailand
Description
Entomopathogens constitute a unique and specialized trophic group of fungi, most of which belong to Hypocreales (Sordariomycetes, Ascomycota). In this study, eight species were collected and isolated from China and Thailand. Through comprehensive morphological analyses and multigene phylogenetic studies (ITS, nrSSU, nrLSU, tef1-α, rpb1, rpb2), seven novel species (Ophiocordyceps jinguangensis sp. nov., O. northeastensis sp. nov., Polycephalomyces bannaensis sp. nov., Po. chiangraiensis sp. nov., Pleurocordyceps shibingensis sp. nov., Pl. tengchongensis sp. nov., and Dingleyomyces yunnanensis sp. nov.) and one known species (O. formicarum) were identified. Additionally, the pairwise homoplasy index (PHI) test results and morphological differences between the new species and their closely related taxa are provided. Notably, as the number of reported hirsutella-like species continues to increase, their phylogenetic placement has become increasingly unclear in previous classifications. To address this issue, this paper presents the first comprehensive summary of the distribution of hirsutella-like species within the families Ophiocordycipitaceae and Polycephalomycetaceae, along with an analysis of the similarities and differences in their phialidic characteristics. These findings significantly expand our knowledge of the diversity, taxonomy, and phylogenetic relationships of entomopathogenic fungi in these families, providing a valuable framework for future studies on their ecology and evolution.
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References
- Adachi K, Chiba K (2007) FTY720 story. Its discovery and the following accelerated development of sphingosine 1-phosphate receptor agonists as immunomodulators based on reverse pharmacology. Perspectives in medicinal chemistry 1: 11–23. https://doi.org/10.1177/1177391X0700100002
- Araújo JP, Hughes DP (2017) Myrmecophilous Ophiocordyceps as a Case Study. The fungal community: its organization and role in the ecosystem, 359 pp. https://doi.org/10.1201/9781315119496-25
- Araújo JPM, Evans HC, Kepler R et al. (2018) Zombie-ant fungi across continents: 15 new species and new combinations within Ophiocordyceps. I. Myrmecophilous hirsutelloid species. Studies in Mycology 90(1): 119–160. https://doi.org/10.1016/j.simyco.2017.12.002
- Araújo JPM, Evans HC, Fernandes IO et al. (2020) Zombie-ant fungi cross continents: II. Myrmecophilous hymenostilboid species and a novel zombie lineage. Mycologia 112(6): 1138–1170. https://doi.org/10.1080/00275514.2020.1822093
- Araújo JPM, Moriguchi MG, Uchiyama S et al. (2021) Ophiocordyceps salganeicola, a parasite of social cockroaches in Japan and insights into the evolution of other closely-related Blattodea-associated lineages. IMA Fungus 12: 3. https://doi.org/10.1186/s43008-020-00053-9
- Araújo JPM, Lebert BM, Vermeulen S et al. (2022) Masters of the manipulator: Two new hypocrealean genera, Niveomyces (Cordycipitaceae) and Torrubiellomyces (Ophiocordycipitaceae), parasitic on the zombie ant fungus Ophiocordyceps camponoti-floridani. Persoonia-Molecular Phylogeny and Evolution of Fungi 49(1): 171–194. https://doi.org/10.3767/persoonia.2022.49.05
- Ban S, Sakane T, Nakagiri A (2015) Three new species of Ophiocordyceps and overview of anamorph types in the genus and the family Ophiocordycipitaceae. Mycological Progress 14(1): 1017–1030. https://doi.org/10.1007/s11557-014-1017-8
- Ban S, Sakane T, Toyama K et al. (2009) Teleomorph–anamorph relationships and reclassification of Cordyceps cuboidea and its allied species. Mycoscience 50(4): 261–272. https://doi.org/10.1007/S10267-008-0480-Y
- Bissett J (1983) Notes on Tolypocladium and related genera. Canadian Journal of Botany 61: 1311–1329. https://doi.org/10.1139/b83-139
- Bischoff JF, Chaverri P, White Jr JF (2005) Clarification of the host substrate of Ascopolyporus and description of Ascopolyporus philodendrus sp. nov. Mycologia 97(3): 710–717. https://doi.org/10.1080/15572536.2006.11832800
- Borel JF (1983) Cyclosporine: historical perspectives. Transplantation Proceedings 15: 2219–2229.
- Boucias DG, Scharf DW, Breaux SE et al. (2007) Studies on the fungi associated with the glassy-winged sharpshooter Homalodisca coagulata with emphasis on a new species Hirsutella homalodiscae nom. prov. Biocontrol 52(2): 231–258. https://doi.org/10.1007/s10526-006-9019-3
- Bruen TC, Philippe H, Bryant D (2006) A simple and robust statistical test for detecting the presence of recombination. Genetics 172: 2665–2681. https://doi.org/10.1534/genetics.105.048975
- Calvillo‐Medina RP, Ponce‐Angulo DG, Raymundo T et al. (2021) Purpureocillium roseum sp. nov. A new ocular pathogen for humans and mice resistant to antifungals. Mycoses 64(2): 162–173. https://doi.org/10.1111/myc.13198
- Capella-Gutiérrez S, Silla-Martínez JM, Gabaldón T (2009) TrimAl: a tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25(15): 1972–1973. https://doi.org/10.1093/bioinformatics/btp348
- Castlebury LA, Rossman AY, Sung GH et al. (2004) Multigene phylogeny reveals new lineage for Stachybotrys chartarum, the indoor air fungus. Mycological Research 108(8): 864–872. https://doi.org/10.1017/S0953756204000607
- Chaverri P, Bischoff JF, Liu M et al. (2005) A new species of Hypocrella, H. macrostroma, and its phylogenetic relationships to other species with large stromata. Mycological Research 109(11): 1268–1275. https://doi.org/10.1017/s0953756205003904
- Chen MJ, Chavez LJ, Kang JY et al. (2025) Harposporium incensis sp. nov. a South American cordycipitoid species exhibiting inter-phylum host-jumping and having potential as a biological control agent for pest management. Mycology 16(1): 238–249. https://doi.org/10.1080/21501203.2024.2350959
- Chen W, Tang Y, Liu T et al. (2024) Purpureocillium jiangxiense sp. nov.: Entomopathogenic Effects on Ostrinia furnacalis and Galleria mellonella. Microorganisms 12(6): 1041. https://doi.org/10.3390/microorganisms12061041
- Chen ZH, Dai YD, Yu H et al. (2013) Systematic analyses of Ophiocordyceps lanpingensis sp. nov., a new species of Ophiocordyceps in China. Microbiological Research 168: 525–532. https://doi.org/10.1016/j.micres.2013.02.010
- Clifton EH, Castrillo LA, Hajek AE (2021) Discovery of two hypocrealean fungi infecting spotted lanternflies, Lycorma delicatula: Metarhizium pemphigi and a novel species, Ophiocordyceps delicatula. Journal of Invertebrate Pathology 186: 107689. https://doi.org/10.1016/j.jip.2021.107689
- Crous PW, Wingfield MJ, Burgess TI et al. (2016) Fungal Planet description sheets: 469–557. Persoonia 37: 218–403. https://doi.org/10.3767/003158516X694499
- Crous PW, Wingfield MJ, Burgess TI et al. (2017) Fungal Planet description sheets: 625–715. Persoonia 39: 270–467 https://doi.org/10.3767/persoonia.2017.39.11
- Crous PW, Wingfield MJ, Lombard L et al. (2019) Fungal planet description sheets: 951–1041. Persoonia 43: 223–425. https://doi.org/10.3767/persoonia.2019.43.06
- Crous PW, Cowan DA, Maggs-Kölling G et al. (2020) Fungal Planet description sheets: 1112–1181. Persoonia: Molecular Phylogeny and Evolution of Fungi 45: 251–409. https://doi.org/10.3767/persoonia.2020.45.10
- Crous PW, Akulov A, Balashov S et al. (2023) New and Interesting Fungi. 6. Fungal Systematics and Evolution 11: 109–156. https://doi.org/10.3114/fuse.2023.11.09
- Crous PW, Wingfield MJ, Jurjević Ž et al. (2024) Fungal Planet description sheets: 1697–1780. Fungal Systematics and Evolution 14: 325–577. https://doi.org/10.3114/fuse.2024.14.19
- Dai Y, Chen S, Wang Y et al. (2024) Molecular phylogenetics of the Ophiocordyceps sinensis-species complex lineage (Ascomycota, Hypocreales), with the discovery of new species and predictions of species distribution. IMA Fungus 15: 2. https://doi.org/10.1186/s43008-023-00131-8
- Das K, Ryu JJ, Hong SM et al. (2023) Molecular phylogeny and morphology of Tolypocladium globosum sp. nov. isolated from soil in Korea. Mycobiology 51(2): 79–86. https://doi.org/10.1080/12298093.2023.2192614
- David SG, Jeffrey KStone (1999) Phylogenetic analysis of nuclear ribosomal DNA places the nematode parasite, Drechmeria coniospora, in Clavicipitaceae. Mycologia 91(6): 993–1000. https://doi.org/10.1080/00275514.1999.12061109
- Dong QY, Wang Y, Wang ZQ et al. (2022) Morphology and phylogeny reveal five novel species in the genus Cordyceps (Cordycipitaceae, Hypocreales) from Yunnan, China. Frontiers in Microbiology 13: 846909. https://doi.org/10.3389/fmicb.2022.846909
- Evans HC, Elliot SL, Hughes DP (2011) Hidden Diversity Behind the Zombie-Ant Fungus Ophiocordyceps unilateralis: Four New Species Described from Carpenter Ants in Minas Gerais, Brazil. PLoS ONE 6: e17024. https://doi.org/10.1371/journal.pone.0017024
- Evans HC, Whitehead PF (2005) Entomogenous fungi of arboreal Coleoptera from Worcestershire, England, including the new species Harposporium bredonense. Mycological Progress 4(2): 91–99. https://doi.org/10.1007/s11557-006-0112-x
- Fan J, Liu P, Zhao K et al. (2023) Three previously undescribed metabolites from Cordyceps cicadae JXCH-1, an entomopathogenic fungus. Natural Products and Bioprospecting 13(1): 46. https://doi.org/10.1007/s13659-023-00410-2
- Fei GX, Guan JQ, Tian HL et al. (2024) A new entomopathogenic fungus, Ophiocordyceps xifengensis sp. nov., from Liaoning, China. Phytotaxa 642(3): 211–224. https://doi.org/10.11646/phytotaxa.642.3.1
- Freire FM (2015) Taxonomia e distribuição de Ophiocordyceps dipterigena (Ophiocordycipitaceae, Hypocreales). Repositório Institucional da UFSC, 1–128. https://repositorio.ufsc.br/handle/123456789/160715
- Gallardo-Pillancari E, González C, Barahona-Segovia RM et al. (2023) Natural infection of Chiromyzinae larvae (Diptera: Stratiomyidae) in southern Chile by Tolypocladium valdiviae sp. nov. Fungal Biology 127(1–2): 845–853. https://doi.org/10.1016/j.funbio.2022.12.004
- Gams W (1971) Tolypocladium, eine Hyphomycetengattung mit geschwollenen Phialiden. Persoonia-Molecular Phylogeny and Evolution of Fungi 6(2): 185–191.
- Gazis R, Skaltsas D, Chaverri P (2014) Novel endophytic lineages of Tolypocladium provide new insights into the ecology and evolution of Cordyceps-like fungi. Mycologia 106(6): 1090–1105. https://doi.org/10.3852/13-346
- Gernandt DS, Stone JK (1999) Phylogenetic analysis of nuclear ribosomal DNA places the nematode parasite, Drechmeria coniospora, in Clavicipitaceae. Mycologia 91(6): 993–1000. https://doi.org/10.1080/00275514.1999.12061109
- Giné A, Sorribas FJ (2017) Effect of plant resistance and BioAct WG (Purpureocillium lilacinum strain 251) on Meloidogyne incognita in a tomato–cucumber rotation in a greenhouse. Pest Management Science 73(5): 880–887. https://doi.org/10.1002/ps.4357
- Gouli V, Gouli S, Marcelino JA et al. (2013) Entomopathogenic fungi associated with exotic invasive insect pests in northeastern forests of the USA. Insects 4(4): 631–645. https://doi.org/10.3390/insects4040631
- Hawksworth DL, Crous PW, Redhead SA et al. (2011) The Amsterdam declaration on fungal nomenclature. IMA fungus 2(1): 105–111. https://doi.org/10.5598/imafungus.2011.02.01.14
- Hodge KT (1998) Revisionary studies in Hirsutella (Anamorphic Hypocreales: Clavicipitaceae). Cornell University.
- Hodge KT, Viaene NM, Gams W (1997) Two Harposporium species with Hirsutella synanamorphs. Mycological Research 101(11): 1377–1382. https://doi.org/10.1017/S0953756297004152
- Hughes DP, Evans HC, Hywel et al. (2009) Novel fungal disease in complex leaf‐cutting ant societies. Ecological Entomology 34(2): 214–220. https://doi.org/10.1111/j.1365-2311.2008.01066.x
- Huson DH, Bryant D (2006) Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution 23: 254–267. https://doi.org/10.1093/molbev/msj030
- Hyde KD, Hongsanan S, Jeewon R et al. (2016) Fungal diversity notes 367–490: taxonomic and phylogenetic contributions to fungal taxa. Fungal Divers 80: 1–270. https://doi.org/10.1007/s13225-016-0373-x
- Hyde KD, Norphanphoun C, Abreu VP et al. (2017) Fungal diversity notes 603–708: Taxonomic and phylogenetic notes on genera and species. Fungal Diversity 87: 1–235. https://doi.org/10.1007/s13225-017-0391-3
- Hyde KD, Noorabadi MT, Thiyagaraja V et al. (2024) The 2024 Outline of Fungi and fungus-like taxa. Mycosphere 15(1): 5146–6239. https://doi.org/10.5943/mycosphere/15/1/25
- Jayasiri SC, Hyde KD, Ariyawansa HA et al. (2015) The faces of fungi database: fungal names linked with morphology, phylogeny and human impacts. Fungal Diversity 74: 3–18. https://doi.org/10.1007/s13225-015-0351-8
- Johnson D, Sung GH, Hywel-Jones NL et al. (2009) Systematics and evolution of the genus Torrubiella (Hypocreales, Ascomycota). Mycological Research 113(3): 279–289. https://doi.org/10.1016/j.mycres.2008.09.008
- Johnston PR, Park D (2023) Dingleyomyces lloydii (Hypocreales, Polycephalomycetaceae), gen. nov., comb. nov. New Zealand Journal of Botany 1–8. https://doi.org/10.1080/0028825X.2023.2291453
- Kepler R, Ban S, Nakagiri A et al. (2013) The phylogenetic placement of hypocrealean insect pathogens in the genus Polycephalomyces: an application of One Fungus One Name. Fungal Biology 117(9): 611–622. https://doi.org/10.1016/j.funbio.2013.06.002
- Kepler RM, Luangsa-ard JJ, Hywel-Jones NL et al. (2017) A phylogenetically-based nomenclature for Cordycipitaceae (Hypocreales). IMA Fungus 8: 335–353. https://doi.org/10.5598/imafungus.2017.08.02.08
- Kepler RM, Sung GH, Harada Y et al. (2012) Host jumping onto close relatives and across kingdoms by Tyrannicordyceps (Clavicipitaceae) gen. nov. and Ustilaginoidea (Clavicipitaceae). American Journal of Botany 99(3): 552–561. https://doi.org/10.3732/ajb.1100124
- Khonsanit A, Luangsa-ard JJ, Thanakitpipattana D et al. (2019) Cryptic species within Ophiocordyceps myrmecophila complex on formicine ants from Thailand. Mycological Progress 18: 147–161. https://doi.org/10.1007/s11557-018-1412-7
- Kobayasi Y (1941) The genus Cordyceps and its allies. Science Reports of the Tokyo Bunrika Daigaku 5: 53–260.
- Kobayasi Y, Shimizu D (1980) Cordyceps species from Japan 3. Bulletin of the National Science Museum Series B 6: 125–146.
- Kuo CC, Chen P, Chen CY (2008) Harposporium from Taiwan. Fungal Science 23(1–4): 35–41.
- Lai YP, Zhang DF (2011) Progress of Applicant Study on Beauveria bassiana and Metarhizium anisopliae to control agricultural pests. Science and Technology of Qinghai Agriculture and Forestry 1: 40–42. https://www.doc88.com/p-9925963263209.html
- Larsson A (2014) AliView: a fast and lightweight alignment viewer and editor for large datasets. Bioinformatics 30: 3276–3278. https://doi.org/10.1093/bioinformatics/btu531
- Li C, Hywel-Jones N, Cao Y et al. (2018) Tolypocladium dujiaolongae sp. nov. and its allies. Mycotaxon 133(2): 229–241. https://doi.org/10.5248/133.229
- Li CR, Xia CR, Lin YR et al. (2005) Hirsutella huangshanensis sp. nov., the anamorph of Cordyceps formosana and its infection on Tenebrio molitor. Mycosystema 24: 349–355. https://doi.org/10.13346/j.mycosystema.2005.03.008
- Li GJ, Hyde KD, Zhao RL et al. (2016) Fungal diversity notes 253–366: taxonomic and phylogenetic contributions to fungal taxa. Fungal Diversity 78: 1–237. https://doi.org/10.1007/s13225-016-0366-9
- Li SY, Lu YL, Zhao J et al. (2025) Morphology and phylogeny reveal a novel species, Tolypocladium phycosomatis (Ophiocordycipitaceae, Hypocreales), from Yunnan, China. Phytotaxa 684(2): 254–264. https://doi.org/10.11646/phytotaxa.684.2.8
- Lim SK, Kallol D, Choi JW et al. (2025) Microascus microspora sp. nov. and Tolypocladium terrae sp. nov.; Two Novel Fungal Species Isolated from Soil in Korea. Mycobiology 53(3): 280–288. https://doi.org/10.1080/12298093.2025.2474291
- Liu L, Cao YR, Zhang CC et al. (2019) An efficient gene disruption system for the nematophagous fungus Purpureocillium lavendulum. Fungal Biology 123(4): 274–282. https://doi.org/10.1016/j.funbio.2018.10.009
- Liu XJ, Guo YL, Yu YX et al. (1989) Isolation and identification of the anamorphic state of Cordyceps sinensis (Berk.) Sacc. Acta Mycologica Sinica 8(1): 35–40.
- Liu YJ, Whelen S, Hall BD (1999) Phylogenetic relationships among ascomycetes: evidence from an RNA polymerse II subunit. Molecular Biology and Evolution 16: 1799–1808. https://doi.org/10.1093/oxfordjournals.molbev.a026092
- Liu Z, Tang D, Lu Y et al. (2024) Morphology and phylogeny of four new species within Polycephalomycetaceae (Hypocreales) parasitising Ophiocordyceps species. MycoKeys 105: 179–202. https://doi.org/10.3897/mycokeys.105.119893
- Liu ZY, Liang ZQ, Liu AY et al. (2002) Molecular evidence for teleomorph–anamorph connections in Cordyceps based on ITS-5.8S rDNA sequences. Mycological Research 106(9): 1100–1108. https://doi.org/10.1017/S0953756202006378
- Long FY, Qin LW, Xiao YP et al. (2021) Multigene phylogeny and morphology reveal a new species, Ophiocordyceps vespulae, from Jilin Province, China. Phytotaxa 478(1): 33–48. https://doi.org/10.11646/phytotaxa.478.1.2
- Luangsa-ard J, Houbraken J, van Doorn T et al. (2011) Purpureocillium, a new genus for the medically important Paecilomyces lilacinus. FEMS microbiology letters 321(2): 141–149. https://doi.org/10.1111/j.1574-6968.2011.02322.x
- Luangsa-ard J, Tasanathai K, Thanakitpipattana D et al. (2018) Novel and interesting Ophiocordyceps spp. (Ophiocordycipitaceae, Hypocreales) with superficial perithecia from Thailand. Studies in Mycology 89(1): 125–142. https://doi.org/10.1016/j.simyco.2018.02.001
- Mains EB (1958) North American entomogenous species of Cordyceps. Mycologia 50: 169–222. https://doi.org/10.1080/00275514.1958.12024722
- Matočec N, Kušan I, Ozimec R (2014) The genus Polycephalomyces (Hypocreales) in the frame of monitoring Veternica cave (Croatia) with a new segregate genus Perennicordyceps. Ascomycete.org 6(5): 125–133.
- Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE) 1–8. New Orleans, LA, USA. https://doi.org/10.1109/GCE.2010.5676129
- Molnár I, Gibson DM, Krasnoff SB (2010) Secondary metabolites from entomopathogenic Hypocrealean fungi. Natural Product Reports 27(9): 1241–1275. https://doi.org/10.1039/c001459c
- Mongkolsamrit S, Noisripoom W, Arnamnart N et al. (2019) Resurrection of Paraisaria in the Ophiocordycipitaceae with three new species from Thailand. Mycological Progress 18: 1213–1230. https://doi.org/10.1007/s11557-019-01518-x
- Mongkolsamrit S, Noisripoom W, Tasanathai K et al. (2020) Molecular phylogeny and morphology reveal cryptic species in Blackwellomyces and Cordyceps (Cordycipitaceae) from Thailand. Mycological Progress 19: 957–983. https://doi.org/10.1007/s11557-020-01615-2
- Mongkolsamrit S, Noisripoom W, Pumiputikul S et al. (2021) Ophiocordyceps flavida sp. nov. (Ophiocordycipitaceae), a new species from Thailand associated with Pseudogibellula formicarum (Cordycipitaceae), and their bioactive secondary metabolites. Mycological Progress 20(4): 477–492. https://doi.org/10.1007/s11557-021-01683-y
- Mongkolsamrit S, Sandargo B, Ebada SS et al. (2023) Bhushaniella gen. nov. (Cordycipitaceae) on spider eggs sac: a new genus from Thailand and its bioactive secondary metabolites. Mycological Progress 22(9): 64. https://doi.org/10.1007/s11557-023-01915-3
- Mongkolsamrit S, Noisripoom W, Tasanathai T et al. (2024) Uncovering cryptic species diversity of Ophiocordyceps (Ophiocordycipitaceae) associated with Coleoptera from Thailand. Fungal Systematics and Evolution 14: 223–250. https://doi.org/10.3114/fuse.2024.14.15
- Nikoh N, Fukatsu T (2000) Interkingdom host jumping underground: phylogenetic analysis of entomoparasitic fungi of the genus Cordyceps. Molecular Biology and Evolution 17: 629–638. https://doi.org/10.1093/oxfordjournals.molbev.a026341
- Petch T (1931) Notes on entomogenous fungi. Transactions of the British Mycological Society 16(1): 55–75. https://doi.org/10.1016/S0007-1536(31)80006-3
- Peng XC, Wen TC, Wei DP et al. (2024) Two new species and one new combination of Ophiocordyceps (Hypocreales, Ophiocordycipitaceae) in Guizhou. MycoKeys 102: 245–266. https://doi.org/10.3897/mycokeys.102.113351
- Perdomo H, Cano J, Gené J et al. (2013) Polyphasic analysis of Purpureocillium lilacinum isolates from different origins and proposal of the new species Purpureocillium lavendulum. Mycologia 105(1): 151–161. https://doi.org/10.3852/11-190
- Pérez-González O, Rodríguez-Villarreal RA, López-Arroyo JI et al. (2015) Mexican strains of Hirsutella isolated from Diaphorina citri (Hemiptera: Liviidae): morphologic and molecular characterization. Florida Entomologist, 290–297. https://doi.org/10.1653/024.098.0147
- Qu JJ, Yu LQ, Zhang J et al. (2018a) A new entomopathogenic fungus, Ophiocordyceps ponerus sp. nov., from China. Phytotaxa 343(2): 116–126. https://doi.org/10.11646/phytotaxa.343.2.2
- Qu JJ, Zhou Y, Yu J et al. (2018b) Estimated divergence times of Hirsutella (asexual morphs) in Ophiocordyceps provides insight into evolution of phialide structure. BMC Evolutionary Biology 18(1): 111. https://doi.org/10.1186/s12862-018-1223-0
- Qu JJ, Zou X, Cao W et al. (2021) Two new species of Hirsutella (Ophiocordycipitaceae, Sordariomycetes) that are parasitic on lepidopteran insects from China. MycoKeys 82: 81–96. https://doi.org/10.3897/mycokeys.82.66927
- Qu JJ, Zou X, Yu JP et al. (2017) Re-description of two entomopathogenic fungi in Hirsutella Pat. Microbiology China 44(11): 2615–2623. https://doi.org/10.13344/j.microbiol.china.170020
- Quandt CA, Kepler RM, Gams W et al. (2014) Phylogenetic-based nomenclatural proposals for Ophiocordycipitaceae (Hypocreales) with new combinations in Tolypocladium. IMA Fungus 5: 121–134. https://doi.org/10.5598/imafungus.2014.05.01.12
- Rambaut A (2014) FigTree 1.4. 2 software. Institute of Evolutionary Biology, Univ. Edinburgh.
- Rasool S, Vidkjær NH, Hooshmand K et al. (2021) Seed inoculations with entomopathogenic fungi affect aphid populations coinciding with modulation of plant secondary metabolite profiles across plant families. The New Phytologist 229(3): 1715–1727. https://doi.org/10.1111/nph.16979
- Rossman AY (1977) The genus Ophionectria (Euascomycetes, Hypocreales). Mycologia 69(2): 355–391. https://doi.org/10.1080/00275514.1977.12020067
- Saltamachia SJ, Araújo JP (2020) Ophiocordyceps desmidiospora, a basal lineage within the "Zombie-Ant Fungi" clade. Mycologia 112(6): 1171–1183. https://doi.org/10.1080/00275514.2020.1732147
- Samarakoon MC, Hsu KC, Chiu CI et al. (2024) Ophiocordyceps taiwanensis sp. nov. (Ophiocordycipitaceae, Hypocreales) on Odontotermes formosanus (Termitidae, Blattodea). Phytotaxa 660(2): 97–111. https://doi.org/10.11646/phytotaxa.660.2.1
- Sanjuan TI, Franco-Molano AE, Kepler RM et al. (2015) Five new species of entomopathogenic fungi from the Amazon and evolution of neotropical Ophiocordyceps. Fungal Biology 119(10): 901–916. https://doi.org/10.1016/j.funbio.2015.06.010
- Schoch CL, Seifert KA, Huhndorf S et al. (2012) Fungal Barcoding Consortium Author List. Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proceedings of the National Academy of Sciences of the United States of America 109(16): 6241–6246. https://doi.org/10.1073/pnas.1117018109
- Simmons DR, Kepler RM, Renner SA et al. (2015) Phylogeny of Hirsutella species (Ophiocordycipitaceae) from the USA: remedying the paucity of Hirsutella sequence data. IMA Fungus 6(2): 345–356. https://doi.org/10.5598/imafungus.2015.06.02.06
- Soares JM, Karlsen-Ayala E, Salvador-Montoya CA et al. (2023) Two novel endophytic Tolypocladium species identified from native pines in south Florida. Fungal Systematics and Evolution 11: 51–61. https://doi.org/10.3114/fuse.2023.11.04
- Spatafora JW, Quandt CA, Kepler RM et al. (2015) New 1F1N species combinations in Ophiocordycipitaceae (Hypocreales). IMA Fungus 6(2): 357–362. https://doi.org/10.5598/imafungus.2015.06.02.07
- Spatafora JW, Sung GH, Sung JM et al. (2007) Phylogenetic evidence for an animal pathogen origin of ergot and the grass endophytes. Molecular Ecology 16(8): 1701–1711. https://doi.org/10.1111/j.1365-294X.2007.03225.x
- Speare AT (1920) On certain entomogenous fungi. Mycologia 12(2): 62–76. https://doi.org/10.1080/00275514.1920.12016820
- Suh SO, Spatafora JW, Ochiel GR et al. (1998) Molecular phylogenetic study of a termite pathogen Cordycepioideus bisporus. Mycologia 90(4): 611–617. https://doi.org/10.1080/00275514.1998.12026950
- Sun JZ, Liu XZ, McKenzie EH et al. (2019) Fungicolous fungi: terminology, diversity, distribution, evolution, and species checklist. Fungal Diversity 95: 337–430. https://doi.org/10.1007/s13225-019-00422-9
- Sun T, Chen Y, Wang D et al. (2024) Mitogenomics, phylogeny and morphology reveal two new entomopathogenic species of Ophiocordyceps (Ophiocordycipitaceae, Hypocreales) from south-western China. MycoKeys 109: 49–72. https://doi.org/10.3897/mycokeys.109.124975
- Sun T, Zou W, Dong Q et al. (2022) Morphology, phylogeny, mitogenomics and metagenomics reveal a new entomopathogenic fungus Ophiocordyceps nujiangensis (Hypocreales, Ophiocordycipitaceae) from Southwestern China. MycoKeys 94: 91–108. https://doi.org/10.3897/mycokeys.94.89425
- Sung GH, Hywel-Jones NL, Sung JM et al. (2007) Phylogenetic classification of Cordyceps and the clavicipitaceous fungi. Studies in Mycology 57: 5–59. https://doi.org/10.3114/sim.2007.57.01
- Sung GH, Spatafora JW, Zare R et al. (2001) A revision of Verticillium sect. Prostrata. II. Phylogenetic analyses of SSU and LSU nuclear rDNA sequences from anamorphs and teleomorphs of the Clavicipitaceae. Nova Hedwigia 72(3–4): 311–328. https://doi.org/10.1127/nova.hedwigia/72/2001/311
- Tang DX, Zhu JY, Luo LJ et al. (2022) Ophiocordyceps ovatospora sp. nov. (Ophiocordycipitaceae, Hypocreales), pathogenic on termites from China. Phytotaxa 574: 105–117. https://doi.org/10.11646/phytotaxa.574.1.8
- Tang DX, Zhao J, Lu YL et al. (2023) Morphology, phylogeny and host specificity of two new Ophiocordyceps species belonging to the "zombie-ant fungi" clade (Ophiocordycipitaceae, Hypocreales). MycoKeys 99: 269. https://doi.org/10.3897/mycokeys.99.107565
- Tasanathai K, Noisripoom W, Chaitika T et al. (2019) Phylogenetic and morphological classification of Ophiocordyceps species on termites from Thailand. MycoKeys 56: 101–129. https://doi.org/10.3897/mycokeys.56.37636
- Tasanathai K, Thanakitpipattana D, Himaman W et al. (2020) Three new Ophiocordyceps species in the Ophiocordyceps pseudoacicularis species complex on Lepidoptera larvae in Southeast Asia. Mycological Progress 19: 1043–1056. https://doi.org/10.1007/s11557-020-01611-6
- Tasanathai K, Khonsanit A, Noisripoom W et al. (2022) Hidden species behind Ophiocordyceps (Ophiocordycipitaceae, Hypocreales) on termites: Four new species from Thailand. Mycological Progress 21: 86. https://doi.org/10.1007/s11557-022-01837-6
- Thanakitpipattana D, Tasanathai K, Mongkolsamrit S et al. (2020) Fungal pathogens occurring on Orthopterida in Thailand. Persoonia - Molecular Phylogeny and Evolution of Fungi 44: 140–160. https://doi.org/10.3767/persoonia.2020.44.06
- Thiyagaraja V, Hyde KD, Piepenbring M et al. (2025) Orders of Ascomycota. Mycosphere 16(1): 536–1411. https://doi.org/10.5943/mycosphere/16/1/8
- Vaidya G, Lohman DJ, Meier R (2011) SequenceMatrix: concatenation software for the fast assembly of multi‐gene datasets with character set and codon information. Cladistics 27: 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x
- Van Vooren N, Audibert C (2005) Révision du complexe "Cordyceps sphecocephala". 1re partie: les guêpes végétales. Publications de la Société Linnéenne de Lyon 74(7): 221–254. https://doi.org/10.3406/linly.2005.13604
- Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990
- Vu D, Groenewald M, de Vries M et al. (2019) Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in Mycology 92: 135–154. https://doi.org/10.1016/j.simyco.2018.05.001
- Wang C, Mo M, Li X et al. (2007) Morphological characteristics and infection processes of nematophagous Harposporium with reference to two new species. Fungal Diversity 26(1): 287–304.
- Wang L, Li HH, Chen YQ et al. (2014) Polycephalomyces lianzhouensis sp. nov., a new species, co-occurs with Ophiocordyceps crinalis. Mycological Progress 13(4): 1089–1096. https://doi.org/10.1007/s11557-014-0996-9
- Wang WJ, Wang XL, Li Y et al. (2012) Molecular and morphological studies of Paecilomyces sinensis reveal a new clade in clavicipitaceous fungi and its new systematic position. Systematics and Biodiversity 10(2): 221–232. https://doi.org/10.1080/14772000.2012.690784
- Wang Y, Liu YF, Tang DX et al. (2022) Tolypocladium reniformisporum sp. nov. and Tolypocladium cylindrosporum (Ophiocordycipitaceae, Hypocreales) co-occurring on Ophiocordyceps sinensis. Mycological Progress 21(1): 199–214. https://doi.org/10.1007/s11557-021-01675-y
- Wang Y, Wei DP, Peng XC et al. (2024) Interesting mycoparasites and Paradingleyomyces lepidopterorum gen. et sp. nov. (Hypocreales, Polycephalomycetaceae) from Yunnan Province, China. MycoKeys 110: 185–210. https://doi.org/10.3897/mycokeys.110.134132
- Wang YB, Wang Y, Fan Q et al. (2020) Multigene phylogeny of the family Cordycipitaceae (Hypocreales): new taxa and the new systematic position of the Chinese cordycipitoid fungus Paecilomyces hepiali. Fungal Diversity 103(1): 1–46. https://doi.org/10.1007/s13225-020-00457-3
- Wang YB, Yu H, Dai YD et al. (2015a) Polycephalomyces agaricus, a new hyperparasite of Ophiocordyceps sp. infecting melolonthid larvae in southwestern China. Mycological Progress 14: 70–78. https://doi.org/10.1007/s11557-015-1090-7
- Wang YB, Yu H, Dai YD et al. (2015b) Polycephalomyces yunnanensis (Hypocreales), a new species of Polycephalomyces parasitizing Ophiocordyceps nutans and stink bugs (hemipteran adults). Phytotaxa 208(1): 34–44. https://doi.org/10.11646/phytotaxa.00.0.0
- Wang YB, Nguyen TT, Dai YD et al. (2018) Molecular phylogeny and morphology of Ophiocordyceps unituberculata sp. nov. (Ophiocordycipitaceae), a pathogen of caterpillars (Noctuidae, Lepidoptera) from Yunnan, China. Mycological Progress 17: 745–753. https://doi.org/10.1007/s11557-017-1370-5
- Wang YH, Ban S, Wang WJ et al. (2021) Pleurocordyceps gen. nov. for a clade of fungi previously included in Polycephalomyces based on molecular phylogeny and morphology. Journal of Systematics and Evolution 59(5): 1065–1080. https://doi.org/10.1111/jse.12705
- Wang YW, Hong TW, Tai YL et al. (2015c) Evaluation of an epitypified Ophiocordyceps formosana (Cordyceps s.l.) for its pharmacological potential. Evidence-Based Complementary and Alternative Medicine, 1–13. https://doi.org/10.1155/2015/189891
- Wei DP, Gentekaki E, Wanasinghe DN et al. (2022) Diversity, molecular dating and ancestral characters state reconstruction of entomopathogenic fungi in Hypocreales. Mycosphere 13: 281–351. https://doi.org/10.5943/mycosphere/si/1f/8
- Wei DP, Wanasinghe DN, Hyde KD et al. (2020) Ophiocordyceps tianshanensis sp. nov. on ants from Tianshan mountains, PR China. Phytotaxa 464(4): 277–292. https://doi.org/10.11646/phytotaxa.464.4.2
- Wei DP, Wanasinghe DN, Xu JC et al. (2021) Three novel entomopathogenic fungi from China and Thailand. Frontiers in Microbiology 11: 608991. https://doi.org/10.3389/fmicb.2020.608991
- Wen TC, Zhu RC, Kang JC et al. (2013) Ophiocordyceps xuefengensis sp. nov. from larvae of Phassus nodus (Hepialidae) in Hunan Province, southern China. Phytotaxa 123(1): 41–50. https://doi.org/10.11646/phytotaxa.123.1.2
- White TJ, Bruns T, Lee S et al. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR protocols: a guide to methods and applications 18: 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1
- Wraight SP, Galaini-Wraight S, Castrillo LA et al. (2018) Collection, isolation, in vitro culture, and laboratory transmission of Hirsutella eleutheratorum (Hypocreales: Ophiocordycipitaceae) from coffee berry borer on Hawai 'i Island. Journal of Invertebrate Pathology 157: 53–66. https://doi.org/10.1016/j.jip.2018.08.002
- Xiao YP, Hongsanan S, Hyde KD et al. (2019) Two new entomopathogenic species of Ophiocordyceps in Thailand. MycoKeys (47): 53. https://doi.org/10.3897/mycokeys.47.29898
- Xiao YP, Wen TC, Hongsanan S et al. (2018) Multigene phylogenetics of Polycephalomyces (Ophiocordycipitaceae, Hypocreales), with two new species from Thailand. Scientific Reports 8(1): 18087. https://doi.org/10.1038/s41598-018-36792-4
- Xiao YP, Wen TC, Hongsanan S et al. (2017) Introducing Ophiocordyceps thanathonensis, a new species of entomogenous fungi on ants, and a reference specimen for O. pseudolloydii. Phytotaxa 328(2): 115–126. https://doi.org/10.11646/phytotaxa.328.2.2
- Xiao YP, Wang YB, Hyde KD et al. (2023) Polycephalomycetaceae, a new family of clavicipitoid fungi segregates from Ophiocordycipitaceae. Fungal Diversity, 1–76. https://doi.org/10.1007/s13225-023-00517-4
- Xiao YP, Yang Y, Jayawardena RS et al. (2024) Four novel Pleurocordyceps (Polycephalomycetaceae) species from China. Frontiers in Microbiology 14: 1256967. https://doi.org/10.3389/fmicb.2023.1256967
- Xie SW, Wei DP, Qiu JZ et al. (2025) Overview of hirsutella-like anamorphs in Ophiocordyceps (Sordariomycetes, Ophiocordycipitaceae): introducing two new species and one new record from China. MycoKeys 119: 95–121. https://doi.org/10.3897/mycokeys.119.145174
- Xu ZH, Tran NL, Wang Y et al. (2022) Phylogeny and morphology of Ophiocordyceps puluongensis sp. nov. (Ophiocordycipitaceae, Hypocreales), a new fungal pathogen on termites from Vietnam. Journal of Invertebrate Pathology 192: 107771. https://doi.org/10.1016/j.jip.2022.107771
- Xu ZS, Deng LP, Wang HY et al. (2025) Description of two new species of Ophiocordyceps: O. sinocampes and O. cystidiata (Ophiocordycipitaceae, Hypocreales) from typical karst landform forests in Guizhou, China. MycoKeys 114: 1–27. https://doi.org/10.3897/mycokeys.114.134323
- Yadav B, Singh UB, Malviya D et al. (2023) Nematophagous fungi: biology, ecology and potential application. In Detection, Diagnosis and Management of Soil-borne Phytopathogens 309–328. Singapore: Springer Nature Singapore, 309–328. https://doi.org/10.1007/978-981-19-8307-8_12
- Yang E, Karunarathna SC, Hasith AK (2024) Microfungi associated with dead caterpillars and spiders in Xishuangbanna Tropical Botanical Garden, PR China. Phytotaxa 644(4): 243–257. https://doi.org/10.11646/phytotaxa.644.4.1
- Yang JI, Stadler M, Chuang WY et al. (2020) In vitro inferred interactions of selected entomopathogenic fungi from Taiwan and eggs of Meloidogyne graminicola. Mycological Progress 19(1): 97–109. https://doi.org/10.1007/s11557-019-01546-7
- Yang Y, Xiao YP, Jayawardena RS et al. (2025) Three new species of Ophiocordyceps (Hypocreales, Ophiocordycipitaceae) and a new host record for Hirsutella vermicola from China. MycoKeys 117: 289. https://doi.org/10.3897/mycokeys.117.144875
- Yang ZL, Qin J, Xia C et al. (2015) Ophiocordyceps highlandensis, a new entomopathogenic fungus from Yunnan, China. Phytotaxa 204(4): 287–295. https://doi.org/10.11646/phytotaxa.204.4.5
- Yu FM, Thilini Chethana KW, Wei DP et al. (2021) Comprehensive Review of Tolypocladium and Description of a Novel Lineage from Southwest China. Pathogens 10(11): 1389. https://doi.org/10.3390/pathogens10111389
- Yu Y, Hou SY, Sun ZL et al. (2018) Drechmeria panacis sp. nov., an endophyte isolated from Panax notoginseng. International Journal of Systematic and Evolutionary Microbiology 68(10): 3255–3259. https://doi.org/10.1099/ijsem.0.002971
- Zare R, Gams W (2001) A revision of Verticillium section Prostrata. Vl. The genus Haptocillium. Nova Hedwigia 73: 271–292. https://doi.org/10.1016/j.mycres.2008.01.019
- Zare R, Gams W, Culham A (2000) A revision of Verticillium sect. Prostrata - I. Phylogenetic studies using ITS sequences. Nova Hedwigia 71: 465–480. https://doi.org/10.1127/nova/71/2000/465
- Zeng XY, Tan TJ, Tian FH et al. (2023) OFPT: a one-stop software for fungal phylogeny. Mycosphere 14: 1730–1741. https://doi.org/10.5943/mycosphere/14/1/20
- Zha LS, Kryukov VY, Ding JH et al. (2021) Novel taxa and species diversity of Cordyceps sensu lato (Hypocreales, Ascomycota) developing on wireworms (Elateroidea and Tenebrionoidea, Coleoptera). MycoKeys 78: 79–117. https://doi.org/10.3897/mycokeys.78.61836
- Zhao J, Wang N, Chen YQ et al. (1999) Molecular identification for the asexual stage of Cordyceps sinensis. Acta Scientiarum Naturalium Universitatis Sunyatseni 38(1): 121–123.
- Zhong X, Li SS, Peng QY et al. (2016) A Polycephalomyces hyperparasite of Ophiocordyceps sinensis leads to shortened duration of production and reduced numbers of host ascospores. Fungal Ecology 21: 24–31. https://doi.org/10.1016/j.funeco.2016.03.002
- Zou WQ, Tang DX, Xu ZH et al. (2022) Multigene phylogeny and morphology reveal Ophiocordyceps hydrangea sp. nov. and Ophiocordyceps bidoupensis sp. nov. (Ophiocordycipitaceae). MycoKeys 92: 109. https://doi.org/10.3897/mycokeys.92.86160
- Zou X, Zhou JX, Liang ZQ et al. (2016) Hirsutella shennongjiaensis, a new entomopathogenic species infecting Earwig (Dermaptera). Mycosystema 35(9): 1070–1079.