. 2025 Jun:111:115-198.

doi: 10.3114/sim.2025.111.04. Epub 2025 Apr 17.

Bionectriaceae: a poorly known family of hypocrealean fungi with major commercial potential

L Zhao^{1

2}, J Z Groenewald¹, L W Hou¹, R C Summerbell^{3

4}, P W Crous^{1

2}

Affiliations

¹ Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, Utrecht, 3584 CT, The Netherlands.
² Microbiology, Department of Biology, Utrecht University, Padualaan 8, Utrecht, 3584 CH, The Netherlands.
³ Sporometrics, Toronto, ON, Canada.
⁴ Dalla Lana School of Public Health, University of Toronto, Toronto, ON, Canada.

PMID: 40371418
PMCID: PMC12070156
DOI: 10.3114/sim.2025.111.04

Bionectriaceae: a poorly known family of hypocrealean fungi with major commercial potential

L Zhao et al. Stud Mycol. 2025 Jun.

. 2025 Jun:111:115-198.

doi: 10.3114/sim.2025.111.04. Epub 2025 Apr 17.

Authors

L Zhao^{1

2}, J Z Groenewald¹, L W Hou¹, R C Summerbell^{3

4}, P W Crous^{1

2}

Affiliations

¹ Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, Utrecht, 3584 CT, The Netherlands.
² Microbiology, Department of Biology, Utrecht University, Padualaan 8, Utrecht, 3584 CH, The Netherlands.
³ Sporometrics, Toronto, ON, Canada.
⁴ Dalla Lana School of Public Health, University of Toronto, Toronto, ON, Canada.

PMID: 40371418
PMCID: PMC12070156
DOI: 10.3114/sim.2025.111.04

Abstract

The ascomycete family Bionectriaceae (Hypocreales) contains cosmopolitan species distributed throughout a broad range of environments, mainly occurring in terrestrial and freshwater ecosystems, with a less frequent occurrence in marine habitats. Members of the family are commonly used in industrial, pharmaceutical, and commercial applications. Applications utilise biodegraders and biocontrol agents, while certain taxa serve as a rich source of bioactive secondary metabolites. In recent years, several studies have proposed new taxonomic concepts within Bionectriaceae based on multi-gene phylogenetic inference. However, the status of several genera remains controversial or unclear, and many need to be re-collected and subjected to molecular analysis. The present study aims to improve our understanding of Bionectriaceae by re-examining CBS culture collection strains preliminarily identified as taxa within this family. Morphological and molecular phylogenetic analyses are based on alignments of the nuclear ribosomal subunits consisting of the internal transcribed spacer regions and intervening 5.8S nrDNA (ITS), as well as partial sequences for the 28S large subunit (LSU) nrDNA. Additional regions within protein-encoding genes were used, including the DNA-directed RNA polymerase II second largest subunit (RPB2), and translation elongation factor 1-alpha (TEF1) regions. The sequences generated were used to reconstruct a phylogenetic backbone of the family Bionectriaceae, and to delineate lineages and generic boundaries within it. Based on these results, seven new genera, 35 new species, and nine new combinations are proposed. A robustly supported phylogenetic framework is provided for Bionectriaceae, resolving 352 species and 50 well-supported genera. This study provides a solid foundation for more in-depth future studies on taxa in the family. Taxonomic novelties: New genera: Clavatomyces Lin Zhao & Crous, Collarimyces Lin Zhao & Crous, Vitreipilata Lin Zhao & Crous, Parageonectria Lin Zhao & Crous, Physaromyces Lin Zhao & Crous, Smyrniomyces Lin Zhao & Crous, Urticomyces Lin Zhao & Crous. New species: Acremonium paramultiramosum Lin Zhao & Crous, Clavatomyces prestoeae Lin Zhao & Crous, Clonostachys novocaledonica Lin Zhao & Crous, Clonostachys tropica Lin Zhao & Crous, Collarimyces guttiformis Lin Zhao & Crous, Emericellopsis mexicana Lin Zhao & Crous, Emericellopsis proliferata Lin Zhao & Crous, Emericellopsis soli Lin Zhao & Crous, Fusariella triangulispora Lin Zhao & Crous, Geonectria alni Lin Zhao & Crous, Geonectria quercus Lin Zhao & Crous, Geosmithia cupressina V. Meshram et al., Geosmithia magnispora Lin Zhao & Crous, Gliomastix olivacea Lin Zhao & Crous, Hydropisphaera armeniaca Lin Zhao & Crous, Hydropisphaera gossypina Lin Zhao & Crous, Hydropisphaera martinicensis Lin Zhao & Crous, Hydropisphaera solani Lin Zhao, L.W. Hou & Crous, Lasionectria chondroidea Lin Zhao & Crous, Lasionectria phormii Lin Zhao, L.W. Hou & Crous, Lasionectriopsis stereicola Lin Zhao & Crous, Nectriopsis cribrariae Lin Zhao & Crous, Nectriopsis floccosa Lin Zhao & Crous, Ovicillium theobromae Lin Zhao & Crous, Paracylindrocarpon jigongshanense Lin Zhao & Crous, Paracylindrocarpon spartinae Lin Zhao & Crous, Parageonectria arachispora Lin Zhao & Crous, Paragliomastix venezuelana Lin Zhao & Crous, Physaromyces sterilis Lin Zhao & Crous, Protocreopsis chlamydospora Lin Zhao & Crous, Protocreopsis gallica Lin Zhao & Crous, Roumegueriella echinulata Lin Zhao & Crous, Sesquicillium pouteriae Lin Zhao & Crous, Sesquicillium thailandense Lin Zhao & Crous, Smyrniomyces setaceus Lin Zhao & P.W. Crous. New combinations: Clavatomyces korfii (Lechat & J. Fourn.) Lin Zhao & Crous, Vitreipilata cirsii (Lechat & J. Fourn) Lin Zhao & Crous, Protocreopsis loweniae (Flakus et al.) Lin Zhao & Crous, Protocreopsis vulpina (Cooke) Lin Zhao & Crous, Proxiovicillium capsici (S.Q. Tong & Y.J. Wu) Lin Zhao & Crous, Sesquicillium shanghaiense (Zhang et al.) Lin Zhao & Crous, Urticomyces pseudoarenulus (Lechat & J. Fourn) Lin Zhao & Crous, Verruciconidia guizhouensis (S.Q. Tong & Y.J. Wu) Lin Zhao & Crous, Verruciconidia saulensis (Lechat & J. Fourn.) Lin Zhao & Crous. New synonyms: Clonostachys aquatica D.F. Bao et al., Emericellopsis microspora Backus & Orpurt. Citation: Zhao L, Groenewald JZ, Hou LW, Summerbell RC, Crous PW (2025). Bionectriaceae: a poorly known family of hypocrealean fungi with major commercial potential. Studies in Mycology 111: 115-198. doi: 10.3114/sim.2025.111.04.

Keywords: Bionectriaceae; industry; multi-locus phylogeny; new taxa; pharmaceutics; taxonomy.

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Figures

**Fig. 1**
Phylogenetic tree inferred from a Maximum Likelihood (RAxML) analysis based on a concatenated alignment of ITS, LSU, *RPB2*, and *TEF1* sequences of 713 strains representing *Bionectriaceae* and outgroup taxa. Numbers at branches indicate support values (RAxML-BS / IQ-TREE-BS / BI-PP) above 50 % / 90 % / 0.90. Fully supported branches (RAxML-BS = 100 % / IQ-TREE-BS = 100 % / BI-PP = 1) are thickened. The new genera and new species are printed in red font, the new combinations in blue font, and the ingroup genera are highlighted with coloured boxes. Strains with a type status are indicated with “T”. The tree is rooted to *Bullanockia australis* (CBS 142093), *Flammocladiella aceris* (CBS 138906), *F. anomiae* (CBS 142775), *F. decora* (CBS 142776), *Stromatonectria caraganae* (CBS 125579, CBS 127387), *Tilachlidium brachiatum* (CBS 505.67, CBS 363.97) and *Xanthonectria pseudopeziza* (CBS 140160, CBS 141245). The scale bar represents the expected number of changes per site.

**Fig. 2**
*Physaromyces sterilis* (ex-type CBS 139048). **A–C.** Colonies on OA, MEA and PDA after 14 d at 25 °C in darkness. **D–J.** Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 3**
*Ovicillium theobromae* (ex-type CBS 110153). **A–C.** Colonies on OA, MEA and PDA after 14 d at 25 °C in darkness. **D–I.** Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars: D, E = 50 μm; F–I = 10 μm.

**Fig. 4**
*Geosmithia cupressina* (ex-type CBS 147103). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J.** Morphological structures from OA culture. **D–H.** Conidiophores. **I, J.** Conidia. Scale bars = 10 μm.

**Fig. 5**
*Geosmithia magnispora* (ex-type CBS 139790). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J.** Morphological structures from OA culture. **D–H.** Conidiophores. **I, J.** Conidia. Scale bars = 10 μm.

**Fig. 6**
*Acremonium paramultiramosum* (ex-type CBS 359.77). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I.** Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars: E, F = 50 μm; G–I = 10 μm.

**Fig. 7**
*Emericellopsis mexicana* (ex-type CBS 125295). **A–C**. Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. D. Cleistothecium. E. Asci. F. Ascospores. **G–I**. Conidiophores. J. Conidia. Scale bars: D = 100 μm; E–J = 10 μm.

**Fig. 8**
*Emericellopsis proliferata* (ex-type CBS 228.59). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars: D, H = 50 μm; E–G, I, J = 10 μm.

**Fig. 9**
*Emericellopsis salmosynnemata* (culture CBS 132049). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D, E.** Cleistothecium. **F, G.** Asci. **H, I.** Ascospores. Scale bars = 10 μm.

**Fig. 10**
*Emericellopsis soli* (ex-type CBS 489.73). **A–C**. Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. D. Cleistothecium. E. Asci. F. Ascospores. **G–I**. Conidiophores. J. Conidia. Scale bars: D = 50 μm; E–J = 10 μm.

**Fig. 11**
*Nectriopsis cribrariae* (ex-type CBS 828.70). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars = 10 μm.

**Fig. 12**
*Nectriopsis floccosa* (ex-type CBS 126110). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. D–I. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars = 10 μm.

**Fig. 13**
*Collarimyces guttiformis* (ex-type CBS 133486). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. D–I. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars = 10 μm.

**Fig. 14**
*Sesquicillium pouteriae* (ex-type CBS 136497). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–G.** Conidiophores. H, I. Conidia. Scale bars = 10 μm.

**Fig. 15**
*Sesquicillium thailandense* (ex-type CBS 139546). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 16**
*Clonostachys novocaledonica* (ex-type CBS 496.90). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–H**. Morphological structures from OA culture. **D, E, G.** Conidiophores. F. Conidia imbricate chains. H. Conidia. Scale bars: D, E = 50 μm; F–H = 10 μm.

**Fig. 17**
*Clonostachys tropica* (ex-type CBS 157.96). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–G.** Conidiophores. **H, I.** Conidia. Scale bars = 10 μm.

**Fig. 18**
*Protocreopsis chlamydospora* (ex-type CBS 144254). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness (CBS 141859). D–J. Morphological structures from OA culture. **D–G.** Conidiophores. H, I. Chlamydospores. J. Conidia. Scale bars = 10 μm.

**Fig. 19**
*Protocreopsis gallica* (ex-type CBS 135079). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars = 10 μm.

**Fig. 20**
*Protocreopsis vulpina* (CBS 565.76). **A–C.** Colonies on OA, MEA, PDA after 14 d in room temperature. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 21**
*Lasionectriopsis stereicola* (ex-type CBS 123299). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars: D = 50 μm; E–I = 10 μm.

**Fig. 22**
*Clavatomyces korfii* (CBS 138710). **A–C**. Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from SNA culture. D. Perithecium. E. Section through perithecium. F. Lateral perithecial wall showing two regions. **G, H.** Asci. I. Ascospores. Scale bars: E = 50 μm; F–I = 10 μm.

**Fig. 23**
*Clavatomyces prestoeae* (ex-type CBS 101691). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–G.** Detail of a portion of the synnemata. H. Conidiogenous cells and conidia. I. Conidia. Scale bars: D = 50 μm; E–I = 10 μm.

**Fig. 24**
*Lasionectria chondroidea* (ex-type CBS 565.73). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from PDA culture. **D–H.** Conidiophores. I. Conidia. Scale bars: D = 50 μm; E–I = 10 μm.

**Fig. 25**
*Lasionectria phormii* (ex-type CBS 102040). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–G.** Conidiophores. **H, I.** Conidia. Scale bars = 10 μm.

**Fig. 26**
*Paragliomastix venezuelana* (ex-type CBS 102074). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–G**. Morphological structures from OA culture. **D–G.** Sterile conidiophore-like structures. Scale bars = 10 μm.

**Fig. 27**
*Parageonectria arachispora* (ex-type CBS 118.87). **A–C**. Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–O**. Morphological structures from OA culture. D. Perithecia. E. Section through perithecium. F. Lateral perithecial wall showing two regions. G. Stroma. **H–J.** Asci. K. Ascospores. **L–N.** Conidiophores. O. Conidia. Scale bars: E = 100 μm; H, M, N = 50 μm; F, G, I–K, O = 10 μm.

**Fig. 28**
*Geonectria alni* (ex-type CBS 140756) **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 29**
*Geonectria quercus* (ex-type CBS 137308). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars: H = 50 μm; D–G, I, J = 10 μm.

**Fig. 30**
*Hydropisphaera armeniaca* (ex-type CBS 135905). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–M**. Morphological structures from PDA culture. D. Perithecia. E. Section through perithecium. F. Lateral perithecial wall showing two regions. **G, H.** Asci. I. Ascospores. **J–L.** Conidiophores. M. Conidia. Scale bars: E = 100 μm; G, H = 50 μm; F, I–M = 10 μm.

**Fig. 31**
*Hydropisphaera gossypina* (ex-type CBS 140581). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from MEA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 32**
*Hydropisphaera martinicensis* (ex-type CBS 136679). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D–I.** Conidiophores. J. Conidia. Scale bars = 10 μm.

**Fig. 33**
*Hydropisphaera solani* (ex-type CBS 147425). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–N**. Morphological structures from PDA culture. D. Perithecium. E. Section through perithecium. F. Lateral perithecial wall showing two regions. **G–I.** Asci. J. Ascospores. K. Percurrently proliferated conidiophores. **L, M.** Conidiophores. N. Conidia. Scale bars: E = 100 μm; F–H = 50 μm; I–N = 10 μm.

**Fig. 34**
*Roumegueriella echinulata* (ex-type CBS 276.59). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–K**. Morphological structures from MEA culture. D. Cleistothecia. E. Asci. **F, G.** Ascospores. **H–J.** Conidiophores. K. Conidia. Scale bars: E–G = 50 μm; H–K = 10 μm.

**Fig. 35**
*Gliomastix olivacea* (ex-type CBS 101736). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–K**. Morphological structures from OA culture. **D–J.** Conidiophores. K. Conidia. Scale bars = 10 μm.

**Fig. 36**
*Paracylindrocarpon jigongshanense* (ex-type CBS 128365). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–I**. Morphological structures from OA culture. **D–H.** Conidiophores. I. Conidia. Scale bars = 10 μm.

**Fig. 37**
*Smyrniomyces setaceus* (ex-type CBS 130334). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–J**. Morphological structures from OA culture. **D, E.** Setae with conidiophores. **F–I.** Conidiophores. J. Conidia. Scale bars: E = 50 μm; F–J = 10 μm.

**Fig. 38**
*Fusariella triangulispora* (ex-type CBS 144361). **A–C.** Colonies on OA, MEA, PDA after 14 d at 25 °C in darkness. **D–K**. Morphological structures from OA culture. **D–J.** Conidiophores. K. Conidia. Scale bars = 10 μm.

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Bionectriaceae: a poorly known family of hypocrealean fungi with major commercial potential

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Bionectriaceae: a poorly known family of hypocrealean fungi with major commercial potential

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