Bionectriaceae, Samuels & Rossmanm, 1999

Zhang, Sheng-Nan, Hyde, Kevin D. & Liu, Jian-Kui, 2024, Morpho-phylogenetic evidence reveals Pseudobionectria phoenicis gen. et sp. nov. (Bionectriaceae) from a mangrove habitat, Phytotaxa 668 (3), pp. 207-225 : 214-218

Genera within Bionectriaceae

Acremonium Link 1809 , type species: Acremonium alternatum Link ; includes over 200 species; primarily characterized by their asexual morphs; most sexually typified species were reassigned to Nectria . members are mainly soilborne saprobes or facultative to obligate pathogens of plants, animals or humans ( Gams 1971; Summerbell et al. 2011). A molecular phylogeny reappraisal of acremonium-like species can be found in Hou et al. (2023).

Alloacremonium L. W. Hou, L. Cai & Crous 2023, type species: Alloacremonium humicola L.W. Hou, L. Cai & Crous ; two species; known only from its acremonium-like asexual morph; found in soil and clinical sample. Molecular data are available ( Hou et al. 2023).

Amphichorda Fr. 1825 , type species: Amphichorda felina (DC.) Fr. ; nine species; represented by its asexual morph (synnematous conidiophores and flask-shaped conidiogenous cells producing holoblastic conidia); typically coprophilous, entomopathogenic or isolated from bat wing surfaces, marine sediments, caves, and animal faces; sequences available. It was previously placed in Cordycipitaceae but has been recently accepted in Bionectriaceae ( Leão et al. 2024; Wang et al. 2024).

Anthonectria Döbbeler 2010 , type species: Anthonectria mammispora Döbbeler ; one species; holomorphic; parasitic on plant leaves, this genus lacks available sequences; original descriptions and illustrations can be found in Döbbeler (2010) and Perera et al. (2023).

Aphanotria Döbbeler 2007 , type species: Aphanotria paradoxa Döbbeler ; one species; known for its sexual morph (fusiform ascospores, 5-septate); parasitic on leaves and lacking molecular data. Original descriptions and illustrations are in Döbbeler (2007) and Perera et al. (2023).

Battarrina (Sacc.) Clem. & Shear 1931 , type species: Battarrina inclusa (Berk. & Broome) Clem. & Shear ; one species; characterized by its cleistothecial morphology; fungicolous on Tuber; with no available sequences. Illustrated in Rossman et al. (1999) and Perera et al. (2023).

Bryocentria Döbbeler 2004 , type species: Bryocentria brongniartii (P. Crouan & H. Crouan) Döbbeler ; 19 species; holomorphic ( Döbbeler et al. 2023; Greiff et al. 2024); biotrophic parasite with bryophytes; sequences available (Stenroos et al. 2010); herbarium specimens are illustrated in Perera et al. (2023).

Bryotria Döbbeler & P.G. Davison 2017 , type species: Bryotria urophora Döbbeler & P.G. Davison ; four species; known for its sexual morph; parasitic lifestyle on liverworts; no molecular data are available ( Perera et al. 2023).

Bulbithecium Udagawa & T. Muroi 1990 , type species: Bulbithecium hyalosporum Udagawa & T. Muroi ; nine species; known for its sexual morph and the acremonium-like asexual morph; isolated from soil, dung or skin of humans, some species are plant-associated; sequences are available ( Hou et al. 2023).

Caespitomonium Crous 2021 , type species: Caespitomonium euphorbiae Crous ; three species; represented by acremonium-like asexual morph; associated with plants; sequence available ( Crous et al. 2021).

Circumfusicillium Trovão et al. 2022 , type species: Circumfusicillium cavernae J. Trovão, F. Soares, D.S. Paiva and A. Portugal; one species; known its asexual morph; inhabiting in limestone or cave-like environment; sequence available ( Trovão et al. 2022).

Clibanites (P. Karst.) P. Karst. 1871 , type species: Clibanites paradoxa (P. Karst.) P.Karst. ; one species; represented by its sexual morph (stromatic, cylintrical ascospores); saprobic on wood; sequences unavailable ( Rossman et al. 1999; Perera et al. 2023).

Clonostachys Corda 1839 , type species: Clonostachys araucaria Corda ; ca. 130 species; it is an asexual morphtypified genus, and the name was recommended for conservation over Bionectria ( Rossman et al. 2013) ; including soil-borne species, mycoparasites, endophytes, epiphytes or saprotrophs; sequences available ( Schroers 2001; Perera et al. 2023; Zhao et al. 2023).

Dimerosporiella Speg. 1908 , type species: Dimerosporiella paulistana Speg. ; eight species; holomorphic, fungicolous or found on living leaves of Asterina, Meliola, Schiffnerula or related species ( Rossman et al. 1999; Perera et al. 2023); Bermúdez-Cova et al. (2023) provided the first sequence for this genus ( D. cephalosporii MB 139, OQ787065), which was excluded in our phylogenetic analysis because of its rather short sequence yield unstable phylogenetic position.

Emericellopsis J.F.H. Beyma 1940 , type species: Emericellopsis terricola J.F.H. Beyma ; 33 species; cleistothecial, with conspicuous acremonium-like conidial morphs; soilborne species; sequences available ( Hou et al. 2023).

Fusariella Sacc. 1884 , type species: Fusariella atrovirens (Berk.) Sacc. ; 22 species; represented by asexual morph; saprobic on plant material, also isolated from soil, plant pathogenic; sequences available ( Hou et al. 2023; Perera et al. 2023).

Geonectria Lechat & J. Fourn. 2018 , type species: Geonectria subalpina Lechat, J. Fourn., M. Vega & Priou ; two species; represented by sexual morph; superficially on soil; sequences available ( Lechat et al. 2018b; Perera et al. 2023).

Geosmithia Pitt 1979 , type species: Geosmithia lavendula (Raper & Fennell) Pitt ; ca. 37 species; known its asexual morph; sequences available ( Hou et al. 2023).

Gliomastix Guég. 1905 , type species: Gliomastix chartarum (Cooke) Guég ; ca. 23 species; known its asexual morph and could be connected to a sphaeriaceous sexual morph (e.g., Hydropisphaera spinulosa ); commonly distributed in soil, and may also be associated with plants or isolated as airborne fungi; sequences available ( Hou et al. 2023).

Gossypinidium L. W. Hou, L. Cai & Crous 2023, type species: Gossypinidium sporodochiale L. W. Hou, L. Cai & Crous ; one species; known its asexual morph (abundant cottony sporodochia with abundant conidia arranged in chains); sequences available ( Hou et al. 2023).

Halonectria E.B.G. Jones 1965 , type species: Halonectria milfordensis E.B.G. Jones ; one species; represented by sexual morph; on submerged wood in marine habitat; sequence unavailable ( Jones 1965; Perera et al. 2023).

Hapsidospora Malloch & Cain 1970 , type species: Hapsidospora irregularis Malloch & Cain ; ca. seven species; holomorphic; isolated from agricultural soil, dung of cow; sequences available and was considered congeneric with Nigrosabulum and Mycoarachis ( Hou et al. 2023) .

Hydropisphaera Dumort. 1822 , type species: Hydropisphaera peziza (Tode) Dumort. ; Hydropisphaera s. lat. is highly polyphyletic and Hydropisphaera s. str. is restrict to four nectria-like species and one cleistothecial genus Heleococcum ; more discussion of this Hydropisphaera could be found in Hou et al. (2023).

Laniatria Döbbeler & P.G. Davison 2017 , type species: Laniatria myxostoma Döbbeler & P.G. Davison ; one species; morphology ( Döbbeler and Davison 2018); sequence unavailable ( Perera et al. 2023).

Lasionectria (Sacc.) Cooke 1884 , type species: Lasionectria mantuana (Sacc.) Cooke ; ca. 30 species; the sexual morph is nectria-like, asexual morph is acremonium-like; sequences available, but most of species lack molecular data ( Hou et al. 2023).

Lasionectriella Lechat & J. Fourn. 2016 , type species: Lasionectriella rubioi Lechat C. & Fournier J. ; four species; morphology see Lechat and Fournier (2016); sequence available ( Hou et al. 2023).

Lasionectriopsis Lechat & P.-A. Moreau 2019, type species: Lasionectriopsis germanica Lechat, P. -A. Moreau & H. Bender; four species; morphology see Lechat et al. (2019); sequence available ( Hou et al. 2023).

Monohydropisphaera L. W. Hou, L. Cai & Crous 2023, type species: Monohydropisphaera fusigera (Berk. & Broome) L.W. Hou, L. Cai & Crous ; one species; holomorphic and sequence available, more discussion of this genus could be found in Hou et al. (2023).

Musananaesporium L.W. Hou, L. Cai & Crous 2023 , type species: Musananaesporium tectonae (R.F. Castañeda) L.W. Hou, L. Cai & Crous ; known its asexual morph; monotypic and is established to accommodate Acremonium tectonae ( Hou et al. 2023) .

Mycocitrus Möller 1901 , type species: Mycocitrus aurantium Möller ; seven species; the sexual morph characterized by its large fleshy orange stromata that clasp and surround bamboo stems, and perithecial ascomata partially to fully immersed in the upper region of the stromata, and the asexual morph is acremonium-like; occurred on culms of living bamboo ( Guadua ) and on Microstachys sp. ( Euphorbiaceae ); sequences available ( Hou et al. 2023).

Nectriella Nitschke ex Fuckel 1870 , type species: Nectriella fuckelii Nitschke ex Fuckel ; sequence unavailable; morphology see Rossman et al. (1999).

Nectriopsis Maire 1911 , type species: Nectriopsis violacea (J.C. Schmidt ex Fr.) Maire ; known its sexual morph, which is distinguished from Nectria s. str. in its thin perithecial wall with textura epidermoidea at the surface, combined with yellow pigmentation; fungicolous and also found on bamboo; sequence available ( Rossman et al. 1999; Hou et al. 2023).

Ochronectria Rossman & Samuels 1999 , type species: Ochronectria calami (Henn. & E. Nyman) Rossman & Samuels ; three species; discussion of its morphology and available sequences could be found in Hou et al. (2023).

Ovicillium Zare & W. Gams 2016 , type species: Ovicillium attenuatum Zare & W. Gams ; six species; morphology see Zare & Gams (2016); sequence available ( Hou et al. 2023).

Ovicuculispora Etayo 2010 , type species: Ovicuculispora parmeliae (Berk. & M.A. Curtis) Etayo ; two species; lichenicolous; morphology and available sequences could be fund in Tadome & Ohmura (2021) and Perera et al. (2023).

Paracylindrocarpon Crous, Roets & L. Lombard 2016 , type species: Paracylindrocarpon aloicola Crous, Roets & L. Lombard ; mostly represented by sexual morph, and the asexual morph reported in Tibpromma et al. (2018); associated with plants; sequences available ( Hou et al. 2023).

Paragliomastix L. W. Hou, L. Cai & Crous 2023, type species: Paragliomastix luzulae (Fuckel) L.W. Hou, L. Cai & Crous ; four species; this genus resembles Gliomastix by having darkly pigmented ameroconidia; sequence available ( Hou et al. 2023).

Periantria Döbbeler & P.G. Davison 2017 , type species: Periantria frullaniae (Racov.) Döbbeler & P.G. Davison ; two species; morphology see Döbbeler & Davison (2018), sequence unavailable ( Perera et al. 2023).

Proliferophialis L. W. Hou, L. Cai & Crous 2023, type species: Proliferophialis apiculata L.W. Hou, L. Cai & Crous ; one species; sequences available, see more details in Hou et al. (2023).

Pronectria Clem. 1931 , type species: Pronectria lichenicola (Ces.) Clem. ; sequence unavailable; herbarium specimen has been illustrated in Perera et al. (2023).

Protocreopsis Yoshim. Doi 1977 , type species: Protocreopsis zingibericola Yoshim. Doi ; the sexual morph is nectria-like, and the asexual morph is acremonium-like; usually occurred on monocotyledonous hosts (commonly on leaves of palms); sequence available ( Hou et al. 2023; Perera et al. 2023).

Proxiovicillium L. W. Hou, L. Cai & Crous 2023, type species: Proxiovicillium blochii (Matr.) L.W. Hou, L. Cai & Crous ; two species; represented by asexual morph; sequence available ( Hou et al. 2023).

Pseudoacremonium Crous 2014 , type species: Pseudoacremonium sacchari Crous ; one species; represented by acremonium-like asexual morph; more details could be found in Crous et al. (2014), Hou et al. (2023).

Pseudobionectria S.N. Zhang, K.D. Hyde & Jian K. Liu 2024 , type species: Pseudobionectria phoenicis S.N. Zhang, K.D. Hyde & Jian K. Liu ; one species; represented by sexual morph, this study.

Pseudosynnemellisia M. Li, Raza & L. Cai 2023 , type species: Pseudosynnemellisia favida M. Li, M. Raza & L. Cai ; one species; represented by asexual morph; isolated from mangrove sediments; sequence available ( Li et al. 2023).

Ramosiphorum L. W. Hou, L. Cai & Crous 2023, type species: Ramosiphorum polyporicola L.W. Hou, L. Cai & Crous ; three species; represented by asexual morph; sequence available ( Hou et al. 2023).

Roumegueriella Speg. 1880 , type species: Roumegueriella muricospora Speg. ; three species; the sexual morph has cleistothecial ascomata with globose ascospores; sequence available and more discussion of this genus could be found in Hou et al. (2023).

Selinia P. Karst. 1876 , type species: Selinia pulchra (G. Winter) Sacc. ; six species; the sexual morph with fusiform ascospores with sheath and the asexual morph represented by sporodochial hyphomycetes; coprophilous; sequence available ( Rossman et al. 1999; Schroers 2001; Perera et al. 2023).

Septofusidium W. Gams 1971 , type species: Septofusidium elegantulum (Pidopl.) W. Gams ; five species; known its asexual morph; parasitic on foliicolous fungi, and saprobic on plant litter; sequence available but no molecular data for the type species ( Gams 1971; Samson 1974; Perera et al. 2023).

Sesquicillium W. Gams 1968 , type species: Sesquicillium buxi (J.C. Schmidt ex Link) W. Gams ; more than 20 species; represented by both asexual (penicillate conidiophores producing aseptate, ellipsoidal or fusiform conidia) and sexual morphs (perithecial stroma with ellipsoidal to fusiform ascospores); usually superficially on plant tissues; sequence available; Sesquicillium is resurrected to accommodate the former subgenera of Bionectria ( Epiphloea and Uniparietina), and phylogenetically sister to Clonostachys ( Zhao et al. 2023) .

Stanjemonium W. Gams, O'Donnell, Schroers & M. Chr. 1999 , type species: Stanjemonium grisellum W. Gams, Schroers & M. Chr. ; five species; this genus is quite different from Acremonium and is known to have a close relationship with Emericellopsis ; sequence available; more discussion of this genus could be found in Gams et al. (1998), Hou et al. (2023).

Stephanonectria Schroers & Samuels 1999 , type species: Stephanonectria keithii (Berk. & Broome) Schroers & Samuels ; three species; this genus is characterized by a peculiar crown-like structure around the ascomatal ostiole and myrothecium-type asexual morph ( Schroers et al. 1999a), illustration of the herbarium specimen could be found in Perera et al. (2023).

Stilbocrea Pat. 1900 , type species: Stilbocrea dussii Pat. ; nine species; sequence available; illustration of the sexual morph herbarium specimen in Perera et al. (2023).

Synnemellisia N.K. Rao, Manohar. & Goos 1989 , type species: Synnemellisia hyalospora N.K. Rao, Manohar. & Goos ; five species; known its hyphomycetous asexual morph; isolated in vitro from the leaf litter; sequence available ( Perera et al. 2023; Liu et al. 2024).

Verruciconidia L. W. Hou, L. Cai & Crous 2023, type species: Verruciconidia verruculosa ( W. Gams & Veenb. - Rijks) L. W. Hou, L. Cai & Crous ; seven species; represented by acremonium-like asexual morph; sequence available ( Hou et al. 2023).

Verrucostoma Hirooka , Tak. Kobay. & P. Chaverri 2010, type species: Verrucostoma freycinetiae Hirooka , Tak. Kobay. & P. Chaverri; two species; represented by asexual morph ( Hirooka et al. 2010); sequence available ( Crous et al. 2015a, b; Perera et al. 2023).

Waltergamsia L.W. Hou, L. Cai & Crous 2023, type species: Waltergamsia fusidioides (Nicot) L.W. Hou, L. Cai & Crous ; this genus is proposed to accommodate eight Acremonium s. lat. species that form a fully supported clade; isolated from soil, human sputum, and also from leaf rachis of Elaeis guineensis ( Arecaceae ), cocoa leaf, ect. ( Hou et al. 2023).

*Genera that were excluded from Bionectriaceae :

Chrysonectria Lechat & J. Fourn. 2018 , type species Chrysonectria finisterrensis Lechat, J. Fourn. & Priou ; this genus was initially introduced in the family Nectriaceae by Lechat et al. (2018a), however, Hou et al. (2023) replaced the genus with a new family Chrysonectriaceae .

Paranectria Sacc. 1878 , type species Paranectria affinis Sacc. ; Bermúdez-Cova et al. (2023) identified a new species Paranectria longiappendiculata (MB 175, OQ801166) and provided its molecular data. However, it clustered with Acremonium sensu stricto with low support in our multi-gene phylogenetic analysis. Hou et al. (2023) transferred Paranectria to Nectriaceae based on the available ITS sequence of the type species Paranectria affinis (K(M):253675, MZ159749.1), which is close to Paracremonium spp.

*Ambiguous genera that have been placed in Bionectriaceae :

Gracilistilbella Seifert 2000 , type species: Gracilistilbella clavulata (Mont.) Seifert. This genus is the B-anamorph of Stilbocrea and was elevated to the genus by Seifert & Samuels (2000); sequences are unavailable.

Rhopalocladium Schroers, Samuels & W. Gams 1999 , type species: Rhopalocladium myxophilum Schroers, Samuels & W. Gams. A monotypic genus proposed for the asexual morph of Nectriopsis sporangiicola ( Schroers et al. 1999b) . It is mentioned in Hyde et al. (2020), but not listed in more recent studies ( Wijayawardene et al. 2022 and Perera et al. 2023).