Akanthomyces Lebert, Z. Wiss. Zool. 9: 449 (1858).
Type: Akanthomyces aculeatus Lebert, Z. Wiss. Zool. 9: 449 (1858).
The genus Akanthomyces as proposed by Lebert (1858), including the type A. aculeatus, primarily infects Lepidoptera and forms a clade distinct from Beauveria and Cordyceps. It includes the moth pathogen Cordyceps tuberculata, which is linked to an asexual morph described as A. pistillariiformis (Samson & Evans 1974). Other fungi in this clade include C. coccidioperitheciata and C. confragosa, pathogens of spiders and scales insects, respectively, which produce torrubielloid perithecia (Kobayasi & Shimizu 1982). The production of superficial perithecia on a stipe distinguishes C. coccidioperitheciata from other sexual forms infecting spiders in Cordycipitaceae, which either lack a stipe or lack superficial perithecia if a stipe is present. Cordyceps confragosa was described by Mains (1949) in Torrubiella, and while the morphology is torrubielloid, Akanthomyces has taxonomic priority by date over Torrubiella (Boudier 1885). In addition, the sexual morph C. confragosa is linked to Lecanicillium lecanii, the type species of Lecanicillium, now considered a synonym of Akanthomyces, which has priority over Lecanicillium (Gams & Zare 2001). Chiriví-Salomón et al. (2015) also showed that L. lecanii (as C. confragosa) as well as some other species of Lecanicillium, namely L. attenuatum, L. muscarium, and L. sabanense, fall within Akanthomyces. nthomyces also includes asexually typified species names previously assigned to the genus Isaria, but not the type species, I. farinosa, which belongs in Cordyceps. In general, the host range for asexual and sexual forms of Akanthomyces are similar, although L. attenuatum (CBS 402.78) was cultured from leaf litter with no host reported. The morphological characters associated with Akanthomyces are also found in a clade of spider-pathogenic species sister to the Gibellula clade (see Hevansia below).
The type species of Torrubiella, T. aranicida, known from a spider in France, was not available for inclusion in molecular phylogenetic analyses. However, several morphological characteristics of T. aranicida suggest that it may belong in Akanthomyces. These include the superficial and separated arrangement of the perithecia and the lack of a subiculum in the type specimen as shown in Johnson et al. (2009). A number of sexual morphs now placed in Akanthomyces have torrubielloid ascomata, specifically A. coccidioperitheciatus on spiders, A. lecanii on scale insects, and A. tuberculata on moths.
Akanthomyces attenuatus (Zare & W. Gams) Spatafora, Kepler & B. Shrestha, comb. nov.
MycoBank MB820860
Basionym: Lecanicillium attenuatum Zare & W. Gams, Nova Hedwigia 73: 19 (2001).
Akanthomyces coccidioperitheciatus (Kobayasi & Shimizu) Spatafora, Kepler & B. Shrestha, comb. nov.
MycoBank MB820880
Basionym: Cordyceps coccidioperitheciata Kobayasi & Shimizu, Bull. Natl. Sci. Mus. Tokyo, B 8: 79 (1982).
Akanthomyces dipterigenus (Petch) Spatafora, Kepler, Zare & B. Shrestha, comb. nov.
MycoBank MB823235
Basionym: Cephalosporium dipterigenum Petch, Naturalist (Hull) 56: 102 (1931).
Synonyms: Cephalosporium longisporum Petch, Trans. Brit. Mycol. Soc. 10: 166 (1925).
Lecanicillium longisporum (Petch) Zare & W. Gams, Nova Hedwigia 73: 16 (2001).
?Acrostalagmus aphidum Oudem., Nederl. Kruidk. Arch. 3(2): 759 (1902) [no type collection found in L, only a drawing which does not allow a definite conclusion].
Non Verticillium longisporum (Stark) Karapappa et al., Mycol. Res. 101: 1293 (1997).
Non Akanthomyces longisporus B. Huang et al., Mycosystema 19: 172 (2000).
Akanthomyces lecanii (Zimm.) Spatafora, Kepler & B. Shrestha, comb. nov.
MycoBank MB820881
Basionym: Cephalosporium lecanii Zimm., Teysmania 9: 241 (1899).
Synonyms: Verticillium lecanii (Zimm.) Viégas, Revista Inst. Café Estado São Paulo 14: 754 (1939).
Lecanicillium lecanii (Zimm.) Zare & W. Gams, Nova Hedwigia 73: 10 (2001).
Torrubiella confragosa Mains, Mycologia 41: 305 (1949).
Cordyceps confragosa (Mains) G.H. Sung et al., Stud. Mycol. 57: 49 (2007).
Hirsutella confragosa Mains, Mycologia 41: 303 (1949).
For further synonyms see Zare & Gams (2001).
Akanthomyces muscarius (Petch) Spatafora, Kepler & B. Shrestha, comb. nov.
MycoBank MB820861
Basionym: Cephalosporium muscarium Petch, Naturalist (Hull) 56: 102 (1931).
Synonyms: Lecanicillium muscarium (Petch) Zare & W. Gams, Nova Hedwigia 73: 13 (2001).
Cephalosporium aphidicola Petch, Trans. Brit. Mycol. Soc. 16: 71 (1931).
Verticillium hemileiae Bouriquet, Encycl. Mycol. 12: 155 (1946).
For further synonyms see Zare & Gams (2001).
Akanthomyces sabanensis (J.S. Chirivi-Salomón et al.) J.S. Chirivi-Salomón, T. Sanjuan & S. Restrepo, comb. nov.
MycoBank MB820862
Basionym: Lecanicillium sabanense J.S. Chiriví-Salomón et al., Phytotaxa 234: 68 (2015).
Akanthomyces tuberculatus (Lebert) Spatafora, Kepler & B. Shrestha, comb. nov.
MycoBank MB820863
Basionym: Akrophyton tuberculatum Lebert, Z. Wiss. Zool. 9: 448 (1858).
Synonyms: Cordyceps tuberculata (Lebert) Maire, Bull. Soc. Hist. Nat. Afrique N. 8: 165 (1917).
Isaria pistillariiformis Pat., Bull. Soc. Mycol. Fr. 9: 163 (1893); as “pistillariaeformis”.
Insecticola pistillariiformis (Pat.) Mains, Mycologia 42: 579 (1950); as “pistillariaeformis”.
Akanthomyces pistillariiformis (Pat.) Samson & H.C. Evans, Acta Bot. Neerl. 23: 29 (1974).
Ascopolyporus Möller, Bot. Mitt. Tropen 9: 300 (1901).
Type: Ascopolyporus polychrous Möller, Bot. Mitt. Tropen 9: 300 (1901).
Ascopolyporus is a genus containing seven species, represented in this study by the type A. polychrous and A. villosus. These two species are strongly supported as monophyletic, however their relationship to other taxa in Cordycipitaceae remains poorly resolved. Sexual or asexual morphologies have been observed in individual collections of Ascopolyporus, and they co-occur in some species (Bischoff et al. 2005). In the sexual form perithecia are produced in a dense hyphal mat directly on top of the scale insect host, and the appearance is similar to that of species in Hypocrella or Moelleriella in Clavicipitaceae. Ascopolyporus species produce multiseptate conidia, a feature also found in Hyperdermium. Ascopolyporus shares another characteristic with some species in Hypocrella in the apparent utilization of plant resources via the scale insect cadaver to attain sizes greatly in excess of the original host (Hywel-Jones & Samuels 1998, Bischoff et al. 2005, Chaverri et al. 2008).
Beauveria Vuill., Bull. Soc. Bot. France 59: 40 (1912).
Type: Beauveria bassiana (Bals.-Criv.) Vuill., Bull. Soc. Bot. France 59: 40 (1912).
The recognition of Beauveria as a genus separate from Cordyceps is a significant change for Cordycipitaceae; their respective type species are not congeneric. The morphological features that unite species of Beauveria have proved remarkably durable over time and no isolates described from other asexually typified genera are known in this clade. Direct links between species of Beauveria and cordyceps-like sexual morphs are well established from molecular data and culture-based experiments, including B. bassiana (Li et al. 2001, Huang et al. 2002), B. brongniartii (Shimizu et al. 1988), and B. sungii (Shrestha et al. 2014a). The host range for the asexual morphs is extensive, infecting many insect species across multiple orders (de Faria & Wraight 2007) as well as being isolated from soil and as foliar endophytes (Vega et al. 2009). The sexual morphs are known from Coleoptera, Lepidoptera, Orthoptera, and Phasmatodea, and here we also describe a new species from Blattodea, expanding the known host range of sexual morphs of Beauveria.
Beauveria acridophila (T. Sanjuan & Franco-Mol.) T. Sanjuan, B. Shrestha, Kepler & Spatafora, comb. nov.
MycoBank MB820883
Basionym: Cordyceps acridophila T. Sanjuan & Franco-Mol., Mycologia 106: 268 (2014).
Beauveria bassiana (Bals.-Criv.) Vuill., Bull. Soc. Bot. Fr. 59: 40 (1912).
Basionym: Botrytis bassiana Bals.-Criv., Linnaea 10: 611 (1835).
Synonyms: Spicaria bassiana (Bals.-Criv.) Vuill., Bull. Soc. Sc. Nancy, ser. 3, 10: 153 (1910).
Penicillium bassianum (Bals.-Criv.) Biourge, Cellule 33: 101 (1923).
Cordyceps bassiana Z.Z. Li et al., Chin. Sci. Bull. 9: 751 (2001).
Beauveria blattidicola M. Chen, Aime, T.W. Henkel & Spatafora, sp. nov. (Fig. 2)
MycoBank MB821050
Etymology: The species epithet refers to the fungus’ occurrence on the host insect family Blattidae.
Diagnosis: Similar in host association to Ophiocordyceps blattarioides but differs in the yellow-orange fleshy stromata, long, sinuous stipe, and cylindrical to narrowly clavate fertile region with partially immersed perithecia.
Type: Guyana: Region 8, Potaro-Siparuni: Pakaraima Mountains, Upper Potaro River Basin, within a 4 km radius of Potaro base camp at 5°18′04.8″N, 59°54′40.4″W, 710–750 m elev.; on adult cockroach, 31 May, 2001, M.C. Aime MCA 1727 (BRG — holotype).
Description: Stromata solitary or paired, unbranched, arising from head or thorax of infected adult of cockroach, with fleshy texture, total length 50–60(–90) mm long; stalk 0.8–1.5 mm broad, light yellow (4A3–4A4); fertile area apical, cylindrical to narrowly clavate, 6–8 × 1.2–1.4 mm, yellowish orange (4A7–4A8). Perithecia partially immersed, darker concolorous, interspersed with white mycelial wefts, presented at right angle to the surface of stroma, in longitudinal section oval to ovoid, 250–370 × 110–250 µm. Asci hyaline, cylindrical, 170–300 × 3–4 µm, with a prominent apical cap 3.0–4.0 µm diam. Ascospores filiform, nearly as long as the asci, smooth, hyaline, distinct irregularly multiseptate, not easily breaking into part-spores. Part-spores 6.0–23.0 × 1.0 µm, cylindrical with truncate ends.
Known distribution: Guyana.
Additional specimens examined: Guyana: Region 8, Potaro-Siparuni: Pakaraima Mountains, Upper Potaro River Basin, within a 4 km radius of Potaro base camp at 5°18′04.8″N, 59°54′40.4″W, 710–750 m elev., on adult cockroach adhered to leaf in litter, 12 June 2000, M.C. Aime MCA 1203 (PUL); on adult cockroach in leaf litter 14 July 2000, T.W. Henkel TH 7645 (HSC); on adult cockroach, 25 May 2001, M.C. Aime MCA 1628 (PUL); on adult cockroach, 7 June 2001, M.C. Aime MCA 1814 (PUL); on adult cockroach in leaf litter 24 July 2003, T.W. Henkel TH 8607 (HSC); on adult cockroach on mineral soil below leaf litter, 17 July 2009, T.W. Henkel TH 9049 (OSC); on adult cockroach, partially buried in litter, 27 May 2010, M.C. Aime & L. Williams MCA 4043 (PUL); on adult cockroach, 9 June 2012, M.C. Aime MCA 4883 (PUL).
GenBank: MCA1727 MF416593, MF416539, MF416483, MF416640; MCA1814 MF416594, MF416540, MF416484, MF416641
Commentary: Species of Cordyceps s. lat. infecting cockroaches have rarely been collected in nature. Only two cockroach-associated species are recorded in the literature: Ophiocordyceps blattarioides (Sanjuan et al. 2015) and O. blattae (Petch 1924, 1931), both classified in Ophiocordycipitaceae. The Neotropical O. blattarioides is associated with adult Blattodea, and is closely related to the morphologically similar adult Orthoptera-associated O. amazonica (Sanjuan et al. 2015). Ophiocordyceps blattae was described originally from Sri Lanka (as Cordyceps blattae), but the species remains poorly known, as sequence data are lacking, and current taxonomic concepts are based entirely on the original description and drawings of Petch (1924). Based on the possession of ophio-ascospores and clavate asci with reduced apices, O. blattae is likely closely related to O. unilateralis (Petch 1931) and is the type of the genus Ophiocordyceps (Sung et al. 2007). Beauveria blattidicola is easily distinguished from both O. blattarioides and O. blattae by the combination of yellow to yellowish orange fleshy stromata, the long and sinuous stalk, cylindrical to narrowly clavate fertile region, and partially immersed perithecia typical of other sexual morphs of Beauveria. Molecular data also strongly support the placement of B. blattidicola in Beauveria (Fig. 1). While B. blattidicola cultures are currently unavailable, its phylogenetic placement predicts a Beauveria-type asexual morph.
Beauveria brongniartii (Sacc.) Petch, Trans. Brit. Mycol. Soc. 10: 249 (1926).
Basionym: Botrytis brongniartii Sacc., Syll. Fung. 10: 540 (1892).
Synonym: Cordyceps brongniartii Shimazu, Trans. Mycol. Soc. Japan 29: 328 (1989).
Beauveria diapheromeriphila (T. Sanjuan & S. Restrepo) T. Sanjuan, B. Shrestha, Kepler & Spatafora, comb. nov.
MycoBank MB820882
Basionym: Cordyceps diapheromeriphila T. Sanjuan & S. Restrepo, Mycologia 106: 270 (2014).
Beauveria locustiphila (Henn.) B. Shrestha, Kepler & Spatafora, comb. nov.
MycoBank MB820884
Basionym: Cordyceps locustiphila Henn., Hedwigia 43: 246 (1904).
Beauveria scarabaeidicola (Kobayasi) S.A. Rehner & Kepler, comb. nov.
MycoBank MB820891
Basionym: Cordyceps scarabaeidicola Kobayasi, Bull. Natl. Sci. Mus. Tokyo, B 2: 137 (1976); as “scarabaeicola”.
Synonym: Beauveria sungii S.A. Rehner & R.A. Humber, Mycologia 103: 1070 (2011).
Beauveria staphylinidicola (Kobayasi & Shimizu) B. Shrestha, Kepler & Spatafora, comb. nov.
MycoBank MB820895
Basionym: Cordyceps staphylinidicola Kobayasi & Shimizu, Bull. Natl. Sci. Mus. Tokyo, B 8: 88 (1982); as “staphylinidaecola”.
Blackwellomyces Spatafora & Luangsa-ard, gen. nov.
MycoBank MB820864
Etymology: This genus is named for Meredith Blackwell and honours her invaluable contributions to our knowledge of insect-associated fungi.
Diagnosis: Blackwellomyces is the least inclusive genus-level clade that includes the species B. cardinalis and B. pseudomilitaris. Blackwellomyces is diagnosed by the unique characters of the ascospore, which have irregularly spaced septa and do not disarticulate into part-spores at maturity.
Type: Blackwellomyces cardinalis (G.H. Sung & Spatafora) Spatafora & Luangsa-ard 2017.
Description: Sexual morph: Stromata solitary or multiple, simple or branched. Stipe fleshy, orange to red, cylindrical to enlarging apically, 4–50 × 0.5–3.0 mm. Fertile area terminal, cylindrical, fusiform to clavate to irregularly shaped, 2–9 × 1–4 mm. Perithecia crowded, loosely embedded, ordinal in orientation, elliptical to fusiform to obclavate. Asci 8-spored, hyaline, cylindrical, possessing a prominent apex. Ascospores smooth, filiform, hyaline, irregularly multiseptate, not fragmenting into part-spores.
Asexual morph: Cultures moderately fast growing in PDA and may turn the media red. Aerial mycelium is whitish to whitish yellow and the reverse side of cultures is red or cream. Conidiogenous cells phialides, solitary or in whorls of 2 or 3, swollen at the base or slightly flask-shaped, wider near the base and tapering at the apex. Conidia hyaline, aseptate, ellipsoidal to elliptical, in some species produced in sympodially imbricate chains. Asexual morphs have been described as similar to species in Clonostachys, Hirsutella, Isaria, and Mariannaea.
Hosts: On larva of Lepidoptera.
Distribution: Southeastern USA, eastern China, Japan, Korea, and Thailand.
Commentary: The species placed here are supported as a distinct clade and separate from other genera of Cordycipitaceae based on the placement of their type species. We describe these taxa as Blackwellomyces on the basis of their phylogenetic novelty and irregularly septate ascospores that do not disarticulate into part-spores. This contrasts with other members of the family in which septation and disarticulation is common.
Blackwellomyces cardinalis (G.H. Sung & Spatafora) Spatafora & Luangsa-ard, comb. nov. MycoBank MB820865
Basionym: Cordyceps cardinalis G.H. Sung & Spatafora, Mycologia 96: 660 (2004).
Blackwellomyces pseudomilitaris (Hywel-Jones & Sivichai) Spatafora & Luangsa-ard, comb. nov.
MycoBank MB820866
Basionym: Cordyceps pseudomilitaris Hywel-Jones & Sivichai, Mycol. Res. 98: 940 (1994).
Cordyceps Fr., Observ. Mycol. 2: 316 [cancellans] (1818), nom. cons.
Type: Cordyceps militaris (L.) Fr., Observ. Mycol. 2: 317 [cancellans] (1818).
In this analysis, many species of Cordyceps, including the type, are resolved as a well-supported clade interspersed with genera described originally for asexual morphs, including Evlachovaea, Isaria, and Microhilum (Fig. 1). Additionally, our analysis indicates this core Cordyceps is not monophyletic with C. cardinalis and C. pseudomilitaris, a result consistent with Sung et al. (2007); the latter two species are proposed in the new genus Blackwellomyces here (see above). Within the core Cordyceps clade, internal relationships are generally well-supported, giving rise to a phylogenetic structure that roughly corresponds to stromatal colour (red to orange vs. white to yellow). A thorough review of the taxonomic history of Cordyceps was provided by Shrestha et al. (2014b), who concluded that Cordyceps is the oldest accepted generic name in this clade and is typified by a sexual morph. Based on the cylindrical shape of the stroma, pre-Linnaean literature of the 17th and early 18th century had recorded C. militaris, the type species of Cordyceps, under the old but obsolete generic names Fungus and Fungoides (Shrestha et al. 2014b). The species was transferred to Clavaria by Linnaeus (1753). Clavaria militaris was then transferred to the ascomycete genus Sphaeria (now rejected in favour of Hypoxylon), a classification that was followed until the early 19th century (Shrestha et al. 2014b). The genus Cordyceps was established (Fries 1818, Link 1833) and over the years was circumscribed to include pathogens of more than 12 insect orders and the fungal genera Elaphomyces and Claviceps (Kobayasi 1941, Mains 1958, Sung et al. 2007, Kepler et al. 2012, Araújo & Hughes 2016, Shrestha et al. 2016). This generic concept of Cordyceps stood for approximately 200 years until the polyphyletic nature of Cordyceps as it had been understood by Kobayasi and Mains was revealed (Sung et al. (2007).
Three monotypic generic names are now considered to be synonyms of Cordyceps. Our data confirmed the findings of Humber et al. (2013) who demonstrated that the type species of the monotypic Evlachovaea, E. kintrischica, is a synonym of Isaria (see below). Similarly, the type species of Microhilum, M. oncoperae, known to have a Cordyceps sexual morph, was nested within Cordyceps, as is the type species of Phytocordyceps, P. ninchukispora.
The generic name Isaria is the oldest available name for the entire group of taxa considered here, including Cordyceps. However, the concept of Isaria has a long and convoluted history, with many changes of status and differences of opinion in how the name should be applied (e.g. Hodge et al. 2005, Gams et al. 2005). Petch (1934) concluded the name was too confusing to use and suggested it be applied to a subgenus of Spicaria. However, Hodge et al. (2005) lectotypified Isaria sensu Fries using an illustration of I. farinosa that appeared in the original description of this species as Ramaria farinosa (Holmskjold 1781). Gams et al. (2005) proposed the use of Isaria for Paecilomyces sect. Isarioidea, now also regarded as a synonym of Cordyceps. Entomogenous species morphologically similar to Isaria can be found distributed throughout Hypocreales (Luangsa-ard et al. 2004), and here are shown to be polyphyletic within Cordycipitaceae. The ex-epitype isolate of I. farinosa (CBS 111113, Gams et al. 2005) is here determined to belong within Cordyceps. We therefore propose the rejection of Isaria in favour of Cordyceps owing to the confusion surrounding the application of Isaria. Additionally, rejecting Cordyceps would be disruptive to a large user community while the name Isaria is not as widely used. Species of Isaria are herein integrated into the monophyletic application of Cordyceps. The diversity of species infecting cicada nymphs complicates the transfer of I. cicadae, and will be addressed in a subsequent paper focused on the group.
Cordyceps amoene-rosea (Henn.) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820975
Basionym: Isaria amoene-rosea Henn., Hedwigia 41: 66 (1902).
Synonym: Paecilomyces amoeneroseus (Henn.) Samson, Stud. Mycol. 6: 37 (1974).
Cordyceps cateniannulata (Z.Q. Liang) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820976
Basionym: Paecilomyces cateniannulatus Z.Q. Liang, Acta Phytopathol. Sin. 11: 10 (1981).
Synonym: Isaria cateniannulata (Z.Q. Liang) Samson & Hywel-Jones, Mycol. Res. 109: 588 (2005).
Cordyceps cateniobliqua (Z.Q. Liang) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820977
Basionym: Paecilomyces cateniobliquus Z.Q. Liang, Acta Phytopathol. Sin. 11: 9 (1981).
Synonym: Isaria cateniobliqua (Z.Q. Liang) Samson & Hywel-Jones, Mycol. Res. 109: 588 (2005).
Cordyceps coleopterorum (Samson & H.C. Evans) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820978
Basionym: Paecilomyces coleopterorum Samson & H.C. Evans, Stud. Mycol. 6: 47 (1974).
Synonym: Isaria coleopterorum (Samson & H.C. Evans) Samson & Hywel-Jones, Mycol. Res. 109: 588 (2005); as “coleopterora”.
Cordyceps farinosa (Holmsk.) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820979
Basionym: Ramaria farinosa Holmsk., K. Danske Vidensk. Selsks. Skr., Nye Samling 1: 279 (1781).
Synonyms: Clavaria farinosa (Holmsk.) Dicks., Fasc. Pl. Crypt. Brit. 2: 25 (1790).
Isaria farinosa (Holmsk.) Fr., Syst. Mycol. 3: 271 (1832); nom. sanct.
Corynoides farinosa (Holmsk.) Gray, Nat. Arr. Brit. PI. 1: 654 (1821).
Spicaria farinosa (Holmsk.) Vuill., Bull. Soc. Mycol. France 27: 76 (1911).
Pénicillium farinosum (Holmsk.) Biourge, Cellule 33: 102 (1923).
Paecilomyces farinosus (Holmsk.) A.H.S. Br. & G. Sm., Trans. Brit. Mycol. Soc. 40: 50 (1957).
Cordyceps fumosorosea (Wize) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820980
Basionym: Isaria fumosorosea Wize, Bull. Int. Acad. Sci. Cracovie, Cl. Sci. Math. Nat.: 721 (1905) [“1904”].
Synonyms: Spicaria fumosorosea (Wize) Vassiljevski, Morbi Plant. 18: 146 (1929).
Paecilomyces fumosoroseus (Wize) A.H.S. Br. & G. Sm., Trans. Brit. Mycol. Soc. 40: 67 (1957).
Cordyceps ghanensis (Samson & H.C. Evans) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820981
Basionym: Paecilomyces ghanensis Samson & H.C. Evans, Stud. Mycol. 6: 46 (1974).
Synonym: Isaria ghanensis (Samson & H.C. Evans) Samson & Hywel-Jones, Mycol. Res. 109: 588 (2005).
Cordyceps javanica (Frieder. & Bally) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820982
Basionym: Spicaria javanica Frieder. & Bally, Meded. Koffiebessenboeboek-Fonds 6: 146 (1923).
Synonyms: Paecilomyces javanicus (Frieder. & Bally) A.H.S. Br. & G. Sm., Trans. Brit. Mycol. Soc. 40: 65 (1957).
Isaria javanica (Frieder. & Bally) Samson & Hywel-Jones, Mycol. Res. 109: 588 (2005).
Cordyceps kintrischica (B.A. Borisov & Tarasov) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820983
Basionym: Evlachovaea kintrischica B.A. Borisov & Tarasov, Mikol. Fitopatol. 33: 250 (1999).
Cordyceps locusticola (Z.Q. Liang et al.) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820984
Basionym: Isaria locusticola Z.Q. Liang et ai, Mycotaxon 105: 31 (2008).
Cordyceps oncoperae (H.Y. Yip & A.C. Rath) P.J. Wright, J. Invert. Path. 64: 146 (1994).
MycoBank MB363549
Basionym: Microhilum oncoperae H.Y. Yip & A.C. Rath, J. Invert. Path. 53: 362 (1989).
Cordyceps poprawskii (Cabanillas ef al.) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820985
Basionym: Isaria poprawskii Cabanillas et al., Mycoscience 54: 162 (2013).
Cordyceps tenuipes (Peck) Kepler, B. Shrestha & Spatafora, comb. nov.
MycoBank MB820986
Basionym: Isaria tenuipes Peck, Ann. Rep. N.Y. St. Mus. Nat. Hist. 31: 44 (1879).
Synonyms: Paecilomyces tenuipes (Peck) Samson, Stud. Mycol. 6: 49 (1974).
Cordyceps takaomontana Yakush. & Kumaz., Sci. Rep. Tokyo Bunrika Daig., B 5: 108 (1941).
Engyodontium de Hoog, Persoonia 10: 53 (1978).
Type: Engyodontium parvisporum (Petch) de Hoog, Persoonia 10: 53 (1978).
Engyodontium was erected by de Hoog (1978) to accommodate the type species, E. parvisporum, and E. album, the latter species was formerly classified in Beauveria, and most recently placed in Parengyodontium (see below). Gams et al. (1984) added four more species. The genus is restricted here to species with cobweb-like colonies that produce dense clusters of denticles on elongated rachides. Conidia are hyaline and globose to subglobose. No sexual reproductive morph has been linked to Engyodontium. Molecular phylogenetic analyses based on E. aranearum support the distinction from Beauveria, but additional study of the type species is required. Species are isolated from soil and arthropod cadavers, and as opportunistic cutaneous and subcutaneous infections of animals including humans.
Gibellula Cavara, Atti Ist. Bot. R. Univ. Pavia, 2 ser. 3: 347 (1894).
Type: Gibellula pulchra Cavara, Atti Ist. Bot. Univ. Lab. Crittog. Pavia 3: 347 (1894).
The genus Gibellula is recognized here for spider-pathogenic fungi that produce primarily synnematous, aspergillus-like conidiophores with terminal vesicles, which give rise to phialides produced on metulae. Molecular phytogenies place all sampled Gibellula species in a single clade along with torrubiella-like sexual morphs. Torrubiella has been shown to be polyphyletic, including astipitate taxa throughout Cordycipitaceae and Hypocreales. The status of Torrubiella is complicated further by the uncertain phylogenetic placement of the type species, T. aranicida. The original description indicated an asexual morphology that more closely approximates Lecanicillium or Simplicillium. Further, the production of scattered perithecia directly from the host, rather than aggregated on a subiculate pad, do not suggest inclusion of Gibellula in Torrubiella. Rather, the genus Torrubiella is regarded as a synonym of Akanthomyces here based on the perithecial arrangement and lack of a subiculum in the type specimen (discussed under Akanthomyces). The genus Granulomanus, based on G. aranearum which is linked to the sexually typified T. albolanata, was regarded as a synonym of Gibellula by Humber & Rombach (1987), but no molecular data exist to confirm or refute this suggestion.
Gibellula arachnophila (Ditmar) Vuill., Bull. Séanc. Soc. Sci. Nancy, sér. 3 11: 156 (1910).
Basionym: Isaria arachnophila Ditmar, in Sturm, Deutschl. Fl., 3 Abt. (Pilze Deutschi.) 1(4): tab. 55 (1817).
Synonyms: Hymenostilbe arachnophila (Ditmar) Petch, Naturalist (Hull) 56: 249 (1931).
Synsterigmatocystis arachnophila Costantin ex Vuill., Bull. Soc. Mycol. France 27: 81 (1911).
Gibellula arachnophila f. macropus Vuill., Bull. Soc. Mycol. France 36: 41 (1920).
Torrubiella arachnophila f. alba Kobayasi & Shimizu, Kew Bull. 31: 561 (1977).
Cordyceps arachnophila J.R. Johnst., Bull. Puerto Rico Insula Exp. Sta. 10: 23 (1915).
Torrubiella arachnophila (J.R. Johnst.) Mains, Mycologia 42: 316 (1950).
Gibellula aranearum P. Syd., Just’s Bot. Jahresber. 57: 321 (1922).
Synonym: Torrubiella gibellulae Petch, Ann. Mycol 30: 391 (1932).
Gibellula clavata Samson & H.C. Evans, Mycologia 84: 306 (1992).
Synonym: Torrubiella clavata Samson & H.C. Evans, Mycologia 84: 306 (1992).
Gibellula dabieshanensis B. Huang et al., Mycosystema 17: 110 (1998).
Synonym: Torrubiella dabieshanensis B. Huang et al., Mycosystema 17: 110 (1998).
Gibellula dimorpha Tzean et al., Mycol. Res. 102: 1350 (1998).
Synonym: Torrubiella dimorpha Tzean et al., Mycol. Res. 102: 1350 (1998).
Gibellula globosa Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, B 8: 45 (1982).
Synonym: Torrubiella globosa Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, B 8: 45 (1982).
Gibellula globosostipitata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, B 8: 49 (1982).
Basionym: Torrubiella globosostipitata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, B 8: 49 (1982).
Gibellula leiopus (Vuill. ex Maubl.) Mains, Mycologia 42: 313 (1950).
Basionym: Gibellula arachnophila f. leiopus Vuill. ex Maubl., Bull. Soc. Mycol. France 36: 42 (1920).
Torrubiella arachnophila var. leiopus Mains, Mycologia 42: 318 (1950).
Torrubiella leiopus (Mains) Kobayasi & Shimizu, Kew Bull. 31: 564 (1977); as “pleiopus”.
Gibellula pulchra Cavara, Atti Ist. bot. R. Univ. Pavia, 2 sér. 3: 347 (1894).
Synonyms: Torrubiella arachnophila var. pulchra Mains, Mycologia 42: 316 (1950).
Torrubiella pulchra (Mains) Koval, Klavitsipital’nye Griby SSSR: 71 (1984).
Hevansia Luangsa-ard, Hywel-Jones & Spatafora, gen. nov.
MycoBank MB820885
Etymology: This genus is named for Harry C Evans and honours his invaluable contributions to our knowledge of insect associated fungi.
Diagnosis: Hevansia is the least inclusive genus-level clade that includes H. novoguineensis and H. nelumboides. Hevansia is diagnosed by the immersed perithecia, an Akanthomyces-like asexual morph, and parasitism on spiders.
Type: Hevansia novoguineensis (Samson & B.L. Brady) Luangsa-ard et al. 2017 (see below)
Description: Sexual morph: Stroma arising from dorsal abdomen, stipe 1–2 mm, fertile part ca 1 mm, white to cream, or in astipitate perithecial cushions surrounding host abdomen or sides of legs. Perithecia immersed, crowded at stipe apex or in cushions, few (<5) to numerous (30+), narrowly ovoid or sickle-shaped. Asci 8-spored, ascospores filiform, hyaline, whole, or disarticulating into part-spores.
Asexual morph: Stroma white, cream-yellow, brown or grey mycelium completely covering host. Synnemata erect, simple or branched, solitary to numerous, cylindrical to clavate, cream to ash-grey or brownish white. In some species 2–4 prominent synnemata up to 6 mm long interspersed with numerous tiny synnemata scattered over host. Phialides in a monolayer, sparsely scattered or crowded, on a basal cell or arising from lateral cells, usually single, occasionally two or three on lateral basal cell, smooth-walled, cylindrical, globose, obovoid, obpyriform or ellipsoid, terminating in short but distinct neck or tapering into a long neck. Conidia catenate, usually one-celled, smooth-walled, hyaline, clavate, cylindrical, cymbiform, fusiform to narrowly obclavate. Colony on PDA white front, reverse cream, orange to pale red, some species with pale wine-red pigment on the agar.
Hosts: On spiders on the underside of leaves of forest plants.
Distribution: Primarily in tropical regions globally, although specimens are known from temperate regions.
Commentary: A biphyletic split is observed among the astipitate species previously considered in Torrubiella with asexual morphs in Gibellula forming a clade of species pathogenic on spiders. Another clade includes species on spiders in the tropics, many of which were previously considered members of Akanthomyces. These species constitute the new genus Hevansia described here. Placement of the spider pathogen C. nelumboides in this genus demonstrates the diverse morphology of Hevansia species, as C. nelumboides produces perithecia in a disc sitting atop a well-formed stipe.
Hevansia arachnophila (Petch) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820886
Basionym: Trichosterigma arachnophilum Petch, Trans. Brit. Mycol. Soc. 8: 215 (1923); as “arachnophila”.
Synonyms: Hirsutella arachnophila (Petch) Petch, Trans. Brit. Mycol. Soc. 9: 93 (1923).
Akanthomyces arachnophilus (Petch) Samson & H.C. Evans, Acta Bot. Neerl. 23: 33 (1974).
Torrubiella flava Petch, Trans. Brit. Mycol. Soc. 9: 127 (1923).
Hevansia cinerea (Hywel-Jones) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820887
Basionym: Akanthomyces cinereus Hywel-Jones, Mycol. Res. 100: 1068 (1996).
Hevansia koratensis (Hywel-Jones) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820888
Basionym: Akanthomyces koratensis Hywel-Jones, Mycol. Res. 100: 1067 (1996).
Hevansia longispora (B. Huang et al.) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820889
Basionym: Akanthomyces longisporus B. Huang et al., Mycosystema 19: 172 (2000).
Hevansia nelumboides (Kobayasi & Shimizu) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820890
Basionym: Cordyceps nelumboides Kobayasi & Shimizu, Kew Bull. 31: 557 (1977).
Hevansia novoguineensis (Samson & B.L. Brady) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820892
Basionym: Akanthomyces novoguineensis Samson & B.L. Brady, Trans. Brit. Mycol. Soc. 79: 571 (1982).
Hevansia ovolongata (L.S. Hsieh et al.) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820893
Basionym: Akanthomyces ovalongatus L.S. Hsieh et al., Mycologia 89: 321 (1997).
Hevansia websteri (Hywel-Jones) Luangsa-ard, Hywel-Jones & Spatafora, comb. nov.
MycoBank MB820894
Basionym: Akanthomyces websteri Hywel-Jones, Mycol. Res. 100: 1068 (1996).
Hyperdermium J.F. White et al., Mycologia 92: 910 (2000).
Type: Hyperdermium caulium (Berk. & M.A. Curtis) Chaverri & K.T. Hodge, Stud. Mycol. 60: 63 (2008) [syn. H. bertonii (Speg.) J.F. White et al., Mycologia 92: 910 (2000)].
Hyperdermium was erected by Sullivan et al. (2000) for Epichloë bertonii and H. pulvinatum, which parasitize scale insects in the tropics. Stromata are flattened or pulvinate and vary in colour from white to orange. Perithecia are immersed to sub-immersed with asci and ascospores characteristic of Cordycipitaceae, but are unique in producing multiseptate conidia. Hyperdermium bertonii, the type species of the genus, was found to have an earlier epithet, so the name was corrected to H. caulium by Chaverri (et al. 2008). In these analyses H. caulium, with C. piperis, is placed in an unresolved position.
Parengyodontium C.C. Tsang et al., Med. Mycol. 54: 708 (2016).
Type: Parengyodontium album (Limber) C.C. Tsang et al., Med. Mycol. 54: 709 (2016).
Based on molecular phylogenetic analyses, Parengyodontium was erected by Tsang et al. (2016) to recognize Engyodontium album as a distinct taxon relative to E. parvisporum. To date it is a monotypic genus.
Simplicillium W. Gams & Zare, Nova Hedwigia 73: 38 (2001).
Type: Simplicillium lanosiniveum (J.F.H. Beyma) Zare & W. Gams, Nova Hedwigia 73: 39 (2001).
Simplicillium includes species isolated from other fungi and soil environments (Zare & Gams 2001, Nonaka et al. 2013). Current phylogenetic analyses resolve Simplicillium as the earliest diverging lineage in Cordycipitaceae (Fig. 1). Simplicillium species are morphologically reduced, producing conidia on the tips of long, slender, solitary phialides. No sexual forms have been associated with Simplicillium.