Phylogenetic analyses

The reciprocal comparisons of 70 % bootstrap trees from individual data sets of the 162-taxon 5-gene dataset did not reveal any significantly supported contradictory nodes (data not shown). These results were interpreted as indicating that no strong incongruence existed among the individual data sets that would be indicative of different phylogenetic gene histories (e.g., lineage sorting or horizontal gene transfer). As a result, all five individual data sets were combined in simultaneous analyses.

MP analyses of the 162-taxon 5-gene data set resulted in 156 equally parsimonious trees. These trees were 21,323 steps with a consistency index (CI) of 0.1598 and a retention index (RI) of 0.6131. One of 156 equally parsimonious trees is shown in Fig. 1. Nodes that collapse in the strict consensus tree are denoted with asterisks. ML analyses of the 162-taxon 5-gene data set resulted in a tree with a log-likelihood (–ln) of 92019.95. In the Bayesian analyses, the five-million generation analysis converged on the log-likelihood (harmonic mean = –ln 9951.22) at approximately around 250,000 generations. The results from five of two-million generation analyses also showed a convergence on the log-likelihood at approximately 2 0,000 generations and the topologies were identical. As a result, the 3,000 trees from the first 300,000 generations were deleted from the five million generation analysis to generate a 50 % majority-rule consensus tree.

A 50 % majority consensus tree (Fig. 2) was generated from the million generation analysis. Since the topology of ML analyses (tree not shown) was nearly identical to that of the Bayesian consensus tree of Fig. 2, the bootstrap proportions of ML analyses are provided above the corresponding nodes in Fig. 2. Previous studies have shown that in interpreting the supports of the phylogenetic estimates of relationships, the posterior probability tends to overestimate the phylogenetic confidence (Doaudy et al. 2003, Lutzoni et al. 2004, Reeb et al. 2004). As a result, the posterior probabilities were used as a supplementary indicator to bootstrap proportions. In this study, nodes were considered strongly supported when supported by both bootstrap proportions (BP ≥ 70 %) and posterior probabilities (PP ≥ 0.95) (Lutzoni et al. 2004).

Species in Clavicipitaceae clade A

Clavicipitaceae clade A comprises five well-supported subclades (Fig. 4). All known species of Cordyceps in the clade are included in the C. taii clade. Species of Cordyceps in the clade possess partially or completely immersed perithecia on clavate to cylindrical fertile parts of the stromata (Zang et al. 1982, Liang et al. 1991, Zare et al. 2001). They produce ascospores that either disarticulate or remain intact at maturity and include species that possess ordinal and obliquely embedded perithecia. In the current classification, clade A includes species of Cordyceps that were formerly classified in three subgenera of Cordyceps. Cordyceps liangshanensis M. Zang, D. Liu & R. Hu forms ordinal perithecia and possess disarticulating ascospores, consistent with C. subg. Cordyceps (Kobayasi 1982, Zang et al. 1982). Cordyceps chlamydosporia H.C. Evans possesses nondisarticulating ascospores, consistent with C. subg. Ophiocordyceps (Zare et al. 2001). Cordyceps taii Z.Q. Liang & A.Y. Liu, a known teleomorph species linked to the anamorph genus Metarhizium Sorokin, produces disarticulating ascospores and obliquely embedded perithecia in the stromata, a trait used to recognize C. subg. Neocordyceps (Liang et al. 1991). Importantly, Cordyceps sp. EFCC 2131 and 2135 (described below as Metacordyceps yongmunensis) produce non-disarticulating ascospores and obliquely embedded perithecia in the stromata, characters inconsistent with any of the subgenera in the current classification.

These results suggest that ascospore morphology and arrangement of perithecia are not phylogenetically informative in recognizing either the C. taii clade, or higher clades of clavicipitaceous fungi. Rather, they are more useful at species level classification. For example, our phylogenetic analyses revealed that C. taii is closely related to C. brittlebankisoides Z. Y. Liu, Z.Q. Liang, Whalley, Y.J. Yao & A.Y. Liu, the purported teleomorph of M. flavoviride (Huang et al. 2005). Although these species are similar to each other in macromorphology (e.g., greenish clavate stromata), they differ in the arrangement of the perithecia. C. brittlebankisoides possesses perithecia that are ordinally placed in the stromata, whereas C. taii has obliquely embedded perithecia. These results therefore suggest that arrangement of the perithecia in the stromata is useful in delimiting these closely related species in the C. taii clade (Fig. 4).

Species in Clavicipitaceae clade B

Species of Cordyceps in Clavicipitaceae clade B possess disarticulating or non-disarticulating ascospores and produce superficial to completely immersed perithecia that are ordinally or obliquely inserted in the stromata. As with the Cordyceps species of clade A, this clade also includes members of the former C. subg. Cordyceps (e.g., C. ophioglossoides (Ehrh.) Link and C. variabilis Petch), C. subg. Ophiocordyceps (e.g., C. acicularis and C. unilateralis (Tul. & C. Tul.) Sacc.), and C. subg. Neocordyceps (e.g., C. nutans Pat. and C. sphecocephala). The majority of Cordyceps species in this clade produce wiry to pliant or fibrous stromata that typically are completely or partially darkly pigmented and parasitize subterranean or wood-inhabiting hosts, which are buried in soil or embedded in decaying wood. Exceptions to this morphology and ecology do exist, however; for example, C. melolonthae (Tul. & C. Tul.) Sacc. is pigmented bright yellow but stains darkly upon handling, and members of the C. sphecocephala clade parasitize adult insects.

Clade B consists of five subclades. All subclades include either species of Cordyceps or anamorphs with potential links to Cordyceps (e.g., Nomuraea atypicola (Yasuda) Samson linked to C. cylindrica Petch) (Fig. 6, Evans & Samson 1987). The well-resolved tree in the present study (Fig. 6) provides the basis to characterize three of the five subclades of clade B. Due to insufficient taxon sampling, it is not possible to characterize the members of the Cordyceps species in the C. gunnii and Pa. lilacinus subclades. In the light of this, we focus on the remaining three subclades that include sufficient numbers of Cordyceps species.

The C. ophioglossoides subclade primarily consists of Cordyceps species that parasitize species of the genus Elaphomyces (e.g., C. ophioglossoides and C. capitata (Holmsk.) Link) and the nymphs of cicadas (e.g., C. inegoënsis Kobayasi and C. paradoxa Kobayasi) buried in soil (Kobayasi 1939, Mains 1957, Kobayasi & Shimizu 1960, 1963). Species in this subclade produce partially or completely immersed perithecia, in clavate to capitate fertile parts of stromata that are darkly pigmented with olivaceous tints (Kobayasi & Shimizu 1960, 1963). Because they produce disarticulating ascospores and ordinal perithecia, all known species of this clade are classified in C. subg. Cordyceps.

Cordyceps subsessilis Petch is unique to the C. ophioglossoides subclade (Fig. 6). It produces perithecia on white or pallid reduced stromata, arising from a rhizomorph-like structure from scarabaeid beetle larvae (Hodge et al. 1996). It is the only member of the subclade that parasitizes beetles embedded in decaying wood (Hodge et al. 1996). Therefore, C. subsessilis differs greatly in ecology and morphology of its stromata from most other taxa in the C. ophioglossoides clade, but it possesses several characters shared by its close relative, C. ophioglossoides (Kobayasi & Shimizu 1960, Hodge et al. 1996). Both species grow axenically on simple media, produce verticilliate anamorphs (C. subsessilis has a Tolypocladium anamorph, whereas C. ophioglossoides has verticillium-like conidiophores), possess nearly identical part-spore morphologies, and produce stromata that are connected to their hosts via rhizomorph-like structures. In contrast, C. capitata/C. longisegmentis have not successfully been grown in culture, they are attached directly to the host, and an anamorph is unknown.

The C. ophioglossoides subclade (Fig. 6) also includes parasites of subterranean cicada nymphs (e.g., C. inegoënsis and C. paradoxa), which are grouped with their close relatives (e.g., C. jezoënsis S. Imai and C. ophioglossoides) that parasitize subterranean truffles of Elaphomyces. Despite low support of inter-species relationships within the C. ophioglossoides subclade due to short branch lengths, C. paradoxa and C. inegoënsis are morphologically more similar to C. jezoënsis and C. ophioglossoides than to any other members of the clade. These taxa produce clavate fertile parts of the stromata rather than capitate stromata like other members of the clade (e.g., C. capitata and C. fracta Mains). Many of these species (e.g., C. jezoënsis and C. paradoxa) are also known to connect to their hosts via rhizomorph-like structures (Kobayasi & Shimizu 1960, 1963), supporting a close phylogenetic relationship.

The C. unilateralis subclade includes the most morphologically diverse assemblages of Cordyceps species (Fig. 6). Most of the species in the clade parasitize larval, pupal or nymph stages of arthropods (Kobayasi 1941, Mains 1958). Species of this clade produce superficial to completely immersed perithecia on the stromata with morphologies ranging from capitate to clavate to filiform (Kobayasi 1941, Mains 1958). They typically possess tough, pliant, or fibrous stromata that are entirely or partially darkly pigmented, although some exceptions (e.g., C. melolonthae and C. variabilis) do exist, which produce brightly pigmented stromata (Mains 1958). Many species in the clade (e.g., C. brunneipunctata Hywel-Jones, C. stylophora Berk. & Broome, and C. unilateralis) are also differentiated by aperithecial stromatal apices while the production of perithecia occurs in subterminal regions of the stroma.

Similar to Cordyceps species in clade A, the C. unilateralis subclade includes species that produce disarticulating or non-disarticulating (intact) ascospores. For example, some species in the C. unilateralis subclade (e.g., C. sinensis (Berk.) Sacc. and C. unilateralis) were formerly classified in C. subg. Ophiocordyceps. But these species are interspersed among other species (e.g., C. agriotidis A. Kawam. and C. robertsii (Hook.) Berk.) that are classified in C. subg. Cordyceps. This indicates that, while ascospore morphology is useful in delimiting closely related Cordyceps species and uniting others in species complexes, it is not diagnostic of the C. unilateralis subclade itself (Fig. 6).

Most members of C. subg. Neocordyceps, as classically treated by Kobayasi (1941, 1982) and others (e.g., Artjariyasripong et al. 2001, Stensrud et al. 2005), form a monophyletic group labelled as the C. sphecocephala subclade within the C. unilateralis group (Fig. 6). The majority of species in the C. sphecocephala subclade produce long, thin, pliant, brightly coloured (or dark marasmioid in a few species) stromata, which terminate in clavate to elongated fertile parts, and possess ascospores that disarticulate into sixty-four part-spores (Kobayasi 1941, 1982, Hywel-Jones 2002). Species in this clade produce perithecia, which are partially or completely immersed in the stromata at strongly oblique angles (Kobayasi 1941, 1982, Mains 1958, Hywel-Jones 1996). This clade is one of the best characterized by its morphology (obliquely embedded perithecia in a well-defined clava) and its ecology of parasitizing adult stages of insects.

Species in Clavicipitaceae clade C

Clavicipitaceae clade C includes C. militaris, the type species of the genus Cordyceps (Fig. 8). Most Cordyceps species in this clade are currently classified in C. subg. Cordyceps (Kobayasi 1941, 1982). This clade also contains the members of the former C. subg. Ophiocordyceps and C. subg. Bolacordyceps, resulting in C. subg. Cordyceps being paraphyletic within clade C (Eriksson 1982, Hywel-Jones 1994, Sung & Spatafora 2004). Species of Cordyceps in this clade produce three ascospore types, including disarticulating ascospores (e.g., C. militaris), intact ascospores (e.g., C. cardinalis and C. pseudomilitaris Hywel-Jones & Sivichai), and bola-ascospores (e.g., C. bifusispora O.E. Erikss.). Of particular note, this clade includes Phytocordyceps ninchukispora C.H. Su & H.-H. Wang in the unispecific genus Phytocordyceps C.H. Su & H.-H. Wang. The genus Phytocordyceps was originally described based on bola-ascospores and its host affiliation as a pathogen of Beilschmiedia erythrophloia Hayata (Lauraceae) plant seeds (Su & Wang 198). Morphologically, this species is most similar to C. bifusispora in that it produces bola-ascospores typical of C. subg. Bolacordyceps. However, the phylogenetic analyses in this study reveal that species producing bola-ascospores (e.g., C. bifusispora and P. ninchukispora) do not form a monophyletic group (Fig. 8). Rather, they are interspersed among other Cordyceps species possessing disarticulating ascospores, most notably C. militaris.

Species of Cordyceps in clade C produce superficial to partially immersed perithecia on fleshy stromata that are pallid to brightly pigmented. This is in contrast to Cordyceps species in clade B, which produce darkly pigmented, wiry to pliant or fibrous stromata. This suggests that pigmentation and texture of stromata may be phylogenetically informative at a higher level of classification. It should be noted, however, that some Cordyceps species in clade C are morphologically similar to distantly related Cordyceps species (e.g., C. melolonthae and C. variabilis) in stromatal pigmentation. Although these characters are useful in recognizing Cordyceps species of clade C, the utility of these characters for any future infrageneric classification is probably limited (Fig. 8). For example, C. militaris is macroscopically similar to C. cardinalis and C. pseudomilitaris. All three species produce orangish to red-coloured and fleshy stromata; however, these species differ in ascospore and anamorph morphology (Sung & Spatafora 2004). Furthermore, C. militaris is known as exhibiting considerable variability in stroma morphology (Sung & Spatafora 2004). Potentially conspecific species, such as C. roseostromata Kobayasi & Shimizu and C. kyusyuënsis A. Kawam., differ in stroma morphology, but are closely related to C. militaris and possess identical ascospore and ascus morphologies (Fig. 8, Hywel-Jones 1994, Sung & Spatafora 2004).

The variation in ascospore morphology of Clavicipitaceae clade C combined with old descriptions and unavailable type material complicates species identification for many taxa, as is the case for much of Cordyceps. For example, this study reveals a close relationship between the anamorphic species, Mariannaea pruinosa Z.Q. Liang from China, C. cf. pruinosa from Korea and Thailand, and Phytocordyceps ninchukispora from Taiwan (Fig. 8). The teleomorph of M. pruinosa is C. pruinosa Petch, which was originally described as producing disarticulating ascospores and reddish orange stromata, parasitizing lepidopteran cocoons (Petch 1924, Kobayasi 1941, Liang 1991). Although the isolate of M. pruinosa was obtained from ascospores (Liang 1991), the morphology of the ascospores was not well characterized. The species was identified primarily based on its host affiliation and macroscopic characters. In our study, C. cf. pruinosa EFCC 5197 and N.H.J. 10627 were collected from the same host family (Lepidoptera, Limacodidae) in Korea and Thailand. They are also closely related and produce reddish orange stromata (Fig. 8) and bola-ascospores and not the typical C. subg. Cordyceps part-spores. It should be noted, however, that Petch did not provide any drawings or images of ascospores and it is possible that the terminal cells of bola-ascospores could easily be interpreted as part-spores. Thus, at this time we use the name C. pruinosa for the Chinese, Korean and Thai collections and, if further attempts fail to locate type material for C. pruinosa, one of these may have to be designated a neotype. The C. pruinosa collections are closely related to and morphologically indistinguishable from P. ninchukispora with the exception of host affiliation, suggesting the possibility of host misidentification in the original description of P. ninchukispora. Because the tree topology of the C. pruinosa/P. ninchukispora complex is indicative of greater phylogenetic species diversity, i.e., the Korean, Thai, and Taiwanese material may represent unique phylogenetic species (Fig. 8), we retain the use of both names until more detailed sampling and analyses have been conducted.

Clavicipitaceae clade C not only includes members of Cordyceps but also species of the genus Torrubiella, which generally parasitize spiders and scale insects (Kobayasi & Shimizu 1982). The genus Torrubiella is morphologically characterized by the production of superficial perithecia on a mycelial subiculum that partially or completely surrounds the host (Kobayasi & Shimizu 1982, Humber & Rombach 1987). Species of Torrubiella also produce disarticulating (e.g., T. ratticaudata Humber & Rombach) and intact (e.g., T. wallacei) ascospores. Among species of Cordyceps, C. tuberculata (Lebert) Maire, a pathogen of adult Lepidoptera, has been considered an intermediate species between Torrubiella and Cordyceps (Humber & Rombach 1987, Kobayasi 1941, Mains 1958). Phylogenetic analyses in this study indicate that the members of Torrubiella do not form a monophyletic group within clade C and are interspersed among species of Cordyceps. This suggests that the stipitate stromata of Cordyceps have been gained or lost several times during the evolution of these fungi. Currently, more than 50 species of Torrubiella have been described and the members of genus Torrubiella are clearly undersampled in this study (Kobayasi & Shimizu 1982).

In summary, the characters of ascospore morphology and the arrangement of perithecia used in the current classification of the genus Cordyceps are not congruent with the three higher clades inferred in these analyses. These characters are likely to prove useful, however, in lower level classifications, such as the delimitation of closely related species and species complexes. The characters most congruent with the three higher clades of clavicipitaceous fungi are texture, pigmentation and morphology of the stromata, but with exceptions. Although we have divided Cordyceps species into three major clades, it is difficult to characterize Cordyceps species within the Clavicipitaceae clade A due to the relatively few teleomorph species that are part of this clade (see key on p. 54). They tend to produce green to white stromata, often with lilac tints, but additional sampling is needed to more definitively characterize the teleomorphs of these species. However, the majority of species within clades B and C are morphologically and/or ecologically distinct (Figs ​(Figs1, 1, ​,22).

The majority of Cordyceps species in clade B are characterized by darkly pigmented, wiry, pliant or fibrous stromata. The dominant form of parasitism exhibited by these species is on subterranean or wood-inhabiting hosts, buried in soil or embedded in decaying wood, such as larval and pupal stages of arthropods. In contrast, Cordyceps species of clade C have brightly pigmented and fleshy stromata and parasitize their hosts in relatively more accessible environments, such as leaf litter, moss, or the uppermost soil layer. Exceptions to these morphological and ecological traits are found in some Cordyceps species in clade B. Cordyceps melolonthae, for example, produces brightly-coloured stromata, although it bruises dark upon handling and its hosts are the larvae of cockchafers or June beetles buried in soil (Mains 1958). Cordyceps unilateralis parasitizes adult ants, but is darkly pigmented with a wiry stroma and subterminal production of perithecia, and members of the C. sphecocephala clade are at least partially brightly pigmented and are restricted to adult stages of insects. These findings suggest that the traits described above are not universally informative, but collectively useful in characterizing Cordyceps species within clade B. That is, there have been gains, losses, and diversifications of most if not all traits during the evolution of these fungi, but general trends in character state evolution are evident.

Clavicipitaceae Clade A

Clavicipitaceae clade A is a well-supported monophyletic group that represents the Clavicipitaceae s. s. (MP-BP = 98 %, ML-BP = 99 %, PP = 1.00 in Figs ​Figs1, 1, ​,2, 2, ​,10).10). The name Clavicipitaceae was first used in 1901 by Earle for the former Hypocreaceae subfam. Clavicipiteae Lindau (Earle 1901). However, Earle (1901) used it without description and without reference to its basionym. The name was then invalidly used by subsequent workers, such as Nannfeldt (1932) and Diehl (1950), until it was validated by Rogerson (1970) as confirmed by Eriksson & Hawksworth (1985). Although Clavicipitaceae is well characterized by cylindrical asci, thickened ascus apices, and filiform ascospores that tend to disarticulate at maturity as in the original description, we restrict the application of Clavicipitaceae s. s. to the members of Clavicipitaceae clade A because of the non-monophyly of Clavicipitaceae s. l. (Fig. 10). These findings suggest that the character states of cylindrical asci and filiform ascospores that disarticulate at maturity are plesiomorphic for the Clavicipitaceae s. l. / Hypocreaceae clade. Importantly, the Hypocreaceae also possesses cylindrical asci and while its ascospores are subglobose to fusiform and easily distinguished from those of Clavicipitaceae s. l., they show a similarly high frequency of disarticulation (Rogerson 1970, Rossman et al. 1999).

The family Clavicipitaceae s. s. includes the grass-associated genera Balansia Speg., Claviceps, Epichloë (Fr.) Tul. & C. Tul., and Myriogenospora G.F. Atk., which were classified in Clavicipitaceae subfam. Clavicipitoideae sensu Diehl 1950 (Fig. 10). Recent molecular studies show that Aciculosporium I. Miyake, Atkinsonella Diehl, Heteroëpichloë E. Tanaka, C. Tanaka, Gafur & Tsuda, Neoclaviceps J.F. White, Bills, S.C. Alderman & Spatafora, and Parepichloë J.F. White & P.V. Reddy are also members of this clade, thus supporting their classification in the Clavicipitaceae s. s. (White & Reddy 1998, Sullivan et al. 2001, Tanaka et al. 2002). Clavicipitaceae s. s. also includes the plant-associated Shimizuomyces paradoxus Kobayasi, which occurs on seeds of Smilax (Smilacaceae). In addition to plant-associated fungi, Clavicipitaceae s. s. contains four arthropod-associated lineages. Three of the four arthropod-associated lineages are characterized as pathogens of scale insects, including Hypocrella (pathogens of scale insects and white flies; Hywel-Jones & Evans 1993, Hywel-Jones & Samuels 1998), Regiocrella P. Chaverri & K.T. Hodge (pathogen of scale insects; Chaverri et al. 2006), and Torrubiella luteorostrata Zimm. (pathogen of scale insects; Hywel-Jones 1993). The fourth lineage is described here as Metacordyceps; it comprises former species of Cordyceps and their related anamorphs and as a genus displays relatively broad arthropod host associations.

CLAVICIPITACEAE (Lindau) Earle ex Rogerson, Mycologia 62: 900. 1970, emend. G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora

Stromata or subiculum darkly or brightly coloured, fleshy or tough. Perithecia superficial to completely immersed, ordinal or oblique in arrangement. Asci cylindrical with thickened ascus apex. Ascospores usually cylindrical and multiseptate, disarticulating into part-spores or non-disarticulating.

Type: Claviceps Tul., Ann. Sci. Nat. Bot., Sér. 3, 20: 43. 1853.

Teleomorphic genera: Aciculosporium, Atkinsonella, Balansia, Claviceps, Epichloë, Heteroepichloë, Hypocrella, Metacordyceps gen. nov., Myriogenospora, Neoclaviceps, Parepichloë, Regiocrella, Shimizuomyces.

Anamorphic genera: Aschersonia, Ephelis Fr., Metarhizium, Neotyphodium A.E. Glenn, C.W. Bacon & Hanlin, Nomuraea, paecilomyces-like, Pochonia, Sphacelia Lév., verticillium-like.

METACORDYCEPS G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, gen. nov. MycoBank MB504182.

Stromata solitaria vel nonnulla aggregata, simplicia vel ramosa. Stipes carnosus vel tenax, albidus, viridi-luteus vel viridulus, cylindricus vel sursum dilatatus. Pars fertilis cylindrica vel clavata. Perithecia partim vel omnino in stromate immersa, perpendicularia vel oblique inserta. Asci cylindrici, apice inspissato. Ascosporae cylindricae, multiseptatae, in cellulas diffrangentes vel maturae integrae remanentes.

Stromata solitary or several, simple or branched. Stipe fleshy or tough, whitish, greenish yellow to greenish, cylindrical to enlarging in fertile part. Fertile part cylindrical to clavate. Perithecia partially or completely immersed in stromata, ordinal or oblique in arrangement. Asci cylindrical with thickened ascus apex. Ascospores cylindrical, multiseptate, disarticulating into part-spores or remaining intact at maturity.

Type: Cordyceps taii Z.Q. Liang & A.Y. Liu

Etymology: Greek meta = behind, a genus close to Cordyceps (and suggesting relationship to Metarhizium).

Anamorphic genera: Metarhizium, Nomuraea, paecilomyces-like, Pochonia.

Commentary: The genus Metacordyceps is proposed for species of Cordyceps s. l. in the Clavicipitaceae s. s. based on the phylogenetic placement of C. taii (Figs ​(Figs1, 1, ​,2, 2, ​,10).10). The genus is applied to the C. taii clade, which is strongly supported (MP-BP = 73 %, ML-BP = 78 %, PP = 1.00 in Figs ​Figs1, 1, ​,2, 2, ​,10).10). Among the members of the clade, the best-known taxon is the anamorphic genus Metarhizium, because of its importance in biological control (Samson et al. 1988, Evans 2003). Currently, three species of Cordyceps (viz., C. brittlebankisoides, C. campsosterni, and C. taii) are known as teleomorphs of Metarhizium (Liang et al. 1991, Liu et al. 2001, Zhang et al. 2004). The genus name Metacordyceps is here used to emphasize that the clade includes the species of Cordyceps s. l. that produce Metarhizium anamorphs although other species of Cordyceps (e.g., C. chlamydosporia) in the clade are not connected to Metarhizium anamorphs.

Metacordyceps yongmunensis G.H. Sung, J.M. Sung & Spatafora, sp. nov. MycoBank MB504183. Figs 5B, 5F-K, 11A-G.

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Fig. 5.

A–E. Representative species of Cordyceps and its allies in Clavicipitaceae clade A. F-K. Morphology of Cordyceps sp. (described here as Metacordyceps yongmunensis sp. nov. below). A. C. liangshanensis on lepidopteran larva, EFCC 1452. B. Cordyceps sp. on lepidopteran pupa, EFCC 1228. C. Hypocrella schizostachyi on scale insect (Hemiptera). D. Shimizuomyces paradoxus on seed of plant (Smilax sieboldii: Smilacaceae). E. Metarhizium sp. on adult of cicada. F. Section of perithecium, EFCC 2131. G. Asci and fascicle, EFCC 2131. H. Asci showing prominent ascus cap, EFCC 2131. I. Asci showing ascus foot, EFCC 2131. J. Ascospores showing indistinct septation, EFCC 2131. K. Discharged intact ascospores on SDAY agar, EFCC 2131. Scale bars: A–E = 10 mm, F = 200 μm, G = 100 μm, H–J = 10 μm, K = 100 μm.

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Fig. 11.

A–B. Line drawings of morphology of Metacordyceps yongmunensis. C–G. Line drawings of pochonia-like anamorph of M. yongmunensis. A. Non-disarticulating ascospore and ascus. B. Oblique arrangement of perithecia in stroma. C. Conidia and phialides. D. Developing conidia germinated from ascospore. E. Chlamydospores submerged in SDAY agar. F. Developing chlamydospores submerged in SDAY agar. G. Intercalary swollen hyphae. Scale bars: A, C–G = 10 μm, B = 200 μm.

Anamorph: pochonia-like.

Stromata nonnulla vel raro singula, clavata, simplicia vel saepius ramosa, in chrysalidibus Lepidopterarum. Pars fertilis alba vel dilute lutea, a stipite haud distincta. Perithecia sparsa vel dense aggregata, partim immersa, brunneo-lutea, dilute brunnea vel aurantiobrunnea, oblique inserta, fusiformia vel clavata, 550–800 × 450–500 μm. Asci 8-spori, hyalini, cylindrici, 205–360 × 5–7 μm, apice conspicue inspissato. Ascosporae filiformes, hyalinae, inconspicue multiseptatae, haud fragmentatae, 180–345 × 1 μm. Anamorphe Pochoniae similis.

Stromata several or rarely solitary, clavate, simple or more usually branched, on pupa of Lepidoptera. Fertile area white to pale yellow, not differentiated from stipe. Perithecia scattered or crowded, loosely immersed, brownish yellow, pale brown to orangish brown, oblique in arrangement, fusiform to clavate, 550–800 × 450–500 μm. Asci 8-spored, hyaline, cylindrical, 205– 360 × 5–7 μm, possessing a prominent apical cap. Ascospores filiform, hyaline, multiseptate with indistinct septation, not fragmenting into part-spores, 180–345 × 1 μm. Conidiophores erect, produced in prostrate aerial hyphae. Phialides hyaline, solitary, awl-shaped, 20–28 × 2–2.2 μm. Conidia hyaline, elliptical to oblong, in slimy heads, 2–3.5 × 1.5–2.4 μm. Chlamydospores present.

Etymology: Yongmunensis in reference to the known locality of the first record of the species being Mt. Yongmun, Republic of Korea.

Known distribution: Republic of Korea.

Specimens examined: Mt. Yongmun, Gyunggi Province, Republic of Korea: 13 June 1998, EFCC 2131 (holotype); 13 June 1998, EFCC 2134; 13 June 1998, EFCC 2135; 30 June 1999, EFCC 3379; 30 June 1999, EFCC 3380; 29 Aug. 1999, EFCC 4342; 8 Aug. 1999, EFCC 4343; 8 June 2000, EFCC 49 1; 30 June 2004, EFCC 12287; 30 June 2004, EFCC 12288; 30 June 2004, EFCC 12291; 8 Aug. 2004, EFCC 12467. Mt. Chiak, Kangwon Province, Republic of Korea: 8 Aug. 2000, EFCC 5750. Bukbang-myun, Kangwon Province, Republic of Korea: 21 June 2002, EFCC 8808. Living culture in EFCC.

Commentary: Most specimens of M. yongmunensis possess several stromata (up to 10), on a large pupa of Lepidoptera deeply buried in soil (Fig. 5B). Stroma of the species is typically branched in a dichotomous way at its basal or upper regions (Fig. 5B). Perithecia are usually obliquely inserted in the stromata with a few exceptions that are ordinally arranged, i.e. at right angles to the surface of the stromata (Fig. 5B). While some perithecia are characterized by an acute narrowing of the perithecium at the ostiole, producing a narrow terminal end (Fig. 5F), others are not significantly narrowed (Fig. 11B). In the asci the ascospores are arranged parallel for their entire length and almost reach the ascus foot, suggesting that ascospores are of approximately the same length as the asci (Figs ​(Figs5I, 5I, 11A). Unlike the distinct septation of ascospores as seen in C. militaris (Fig. 9O), the septa of the ascospores are indistinct and discharged ascospores do not disarticulate into part-spores (Figs ​(Figs5K, 5K, 11A).

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Fig. 9.

A–M. Representative species of Cordyceps and its allies in Clavicipitaceae clade C. N–S. Perithecia, asci, and ascospores. A. C. militaris on lepidopteran pupa, EFCC 5192. B. C. kyusyuënsis on lepidopteran larva, EFCC 10985. C. C. chichibuënsis on coleopteran pupa, EFCC 422. D. C. cf. ochraceostromata on lepidopteran larva, EFCC 11846. E. C. scarabaeicola on scarabaeid beetle (Coleoptera), EFCC 5014. F. C. staphylinidicola on coleopteran larva, EFCC 783. G. C. bifusispora on lepidopteran pupa, EFCC 2626. H. C. cf. pruinosa on lepidopteran pupa (Limacodidae), EFCC 11756. I. C. cardinalis on lepidopteran larva, EFCC 12212. J. C. tuberculata on adult of moth (Lepidoptera), EFCC 2067. K. Torrubiella sp. on spider (Arachnida), EFCC 10882. L. Beauveria sp. on adult of beetle (Coleoptera), EFCC 1357. M. Isaria tenuipes on lepidopteran pupa, EFCC 1497. N. C. cardinalis, section of perithecia in stroma, OSC 93609. O. C. militaris, ascus with disarticulating ascospores, OSC 93623. P. C. cardinalis, ascus with nondisarticulating ascospores, OSC 93609. Q. C. cf. pruinosa, fusiform terminal parts of ascospores in ascus, EFCC 7481. R. C. militaris, multiseptated ascospores in ascus, OSC 93623. S. C. cf. pruinosa, thread-like structures connecting fusiform terminal parts of ascospores, EFCC 7481. Scale bars: A–M = 10 mm, N = 100 μm, O–S = 5 μm.

In the anamorph of M. yongmunensis, cultures derived from ascospores are moderately fast growing in SDAY (Sabouraud-dextrose-yeast extract agar) and the colonies reach 25–35 mm diam at 25 °C in 10 d. Colonies are slightly cottony without zonation and white with a green margin, remaining greenish brown at the reverse side of the cultures. Conidiophores are erect and produced in prostrate aerial hyphae. Phialides are solitary, not in whorls, broader at the base and tapering towards the end, measuring 20–28 × 2.0–2.2 μm (Fig. 11C). Conidia are in slimy heads (with usually 2 or 3 conidia) and ellipsoidal to oblong, measuring 2–3.5 × 1.5–2.4 μm (Fig. 11C). In submerged areas of the cultures, chlamydospores are developed in chains or reduced to intercalary swollen structures (Figs 11E-G). The anamorph of M. yongmunensis is best classified as pochonia-like because of its subulate phialides and production of chlamydospores, although verticillium-like whorls of phialides were not observed (Zare et al. 2001). In Metacordyceps, M. yongmunensis is most similar to M. chlamydosporia (= C. chlamydosporia) in the shape of perithecia and its anamorph. Both species produce brownish perithecia that possess long terminal ends in white or pale yellow stromata (Zare et al. 2001). The anamorph of M. chlamydosporia is identical with the type of Pochonia. Thus the production of chlamydospores can be informative for recognizing some species of Metacordyceps.

Accepted names and new combinations for Metacordyceps

The following taxa are accepted species of Metacordyceps based on their inclusion in molecular phylogenies presented herein¹ (see Table 1) or morphological descriptions matching the characters described above². The known anamorph connection is provided for the species of Metacordyceps.

• ²Metacordyceps brittlebankisoides (ZuoY. Liu, Z.Q. Liang, Whalley, Y.-J. Yao & A.Y. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504184.

  • • ≡ Cordyceps brittlebankisoides ZuoY. Liu, Z.Q. Liang, Whalley, Y.-J. Yao & A.Y. Liu, J. Invert. Pathol. 78: 179. 2001.

  • Anamorph: Metarhizium

• ²Metacordyceps campsosterni (W.M. Zhang & T.H. Li) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504185

  • • ≡ Cordyceps campsosterni W.M. Zhang & T.H. Li, Fungal Diversity 17: 240. 2004. [as C. `campsosterna'].

  • Anamorph: Metarhizium

• ¹Metacordyceps chlamydosporia (H.C. Evans) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504186.

  • • ≡ Cordyceps chlamydosporia H.C. Evans, in Zare et al., Nova Hedwigia 73: 59. 2001.

  • Anamorph: Pochonia chlamydosporia (Goddard) Zare & W. Gams

• ¹Metacordyceps liangshanensis (M. Zang, D. Liu & R. Hu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504187.

  • ≡ Cordyceps liangshanensis M. Zang, D. Liu & R. Hu, Acta Bot. Yunnanica 4: 174. 1982.

• ¹Metacordyceps taii (Z.Q. Liang & A.Y. Liu) G.H. Sung, J.M. Sung, Hywel-Jones, Spatafora, comb. nov. MycoBank MB504188.

  • • ≡ Cordyceps taii Z.Q. Liang & A.Y. Liu, Acta Mycol. Sin. 10: 257. 1991.

  • Anamorph: Metarhizium anisopliae var. anisopliae (Metschn.) Sorokin

• ¹Metacordyceps yongmunensis G.H. Sung, J.M. Sung, Spatafora, sp. nov., see p. 27.

  • Anamorph: pochonia-like

New combinations for anamorphs associated with Metacordyceps

T. parasiticum is transferred to the genus Pochonia based on molecular phylogenies presented herein¹.

• ¹Pochonia parasitica (G.L. Barron) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504189.

  • ≡ Tolypocladium parasiticum G.L. Barron, Canad. J. Bot. 58: 439. 1980.

CLAVICIPITACEAE Clade B

Clavicipitaceae clade B is strongly supported (MP-BP = 93 %, ML-BP = 98 %, PP = 1.00 in Figs ​Figs1, 1, ​,2, 2, ​,10)10) and the family Ophiocordycipitaceae is proposed for it with the type genus Ophiocordyceps Petch. Most species of the Ophiocordycipitaceae produce darkly pigmented stromata that are pliant to wiry, or fibrous to tough in texture. Ecologically, many species of the family are known as pathogens of subterranean or wood-inhabiting hosts, buried in soil or embedded in decaying wood. Notable exceptions do exist to these traits with brightly coloured species that may or may not attack adult stages of hosts and occur in exposed habitats.

OPHIOCORDYCIPITACEAE G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, fam. nov. MycoBank MB504190.

Stromata vel subiculum fusca vel raro laete colorata, tenacia, fibrosa vel flexibilia, raro carnosa, saepe ostiolis peritheciorum prominentibus, summa saepe peritheciis carentia. Perithecia superficialia vel omnino immersa, perpendicularia ad superficiem vel oblique inserta. Asci cylindrici, apice inspissato. Ascosporae cylindricae, multiseptatae, maturae in cellulas diffrangentes vel integrae remanentes.

Stromata or subiculum darkly pigmented or rarely brightly coloured, tough, fibrous to pliant, rarely fleshy, often with aperithecial apices or lateral pads. Perithecia superficial to completely immersed, ordinal or oblique in arrangement. Asci usually cylindrical with thickened ascus apex. Ascospores usually cylindrical, multiseptate, disarticulating into part-spores or nondisarticulating.

Type: Ophiocordyceps Petch, Trans. Brit. Mycol. Soc. 16: 74. 1931.

Teleomorphic genera: Elaphocordyceps, Ophiocordyceps

Anamorphic genera: Haptocillium, Harposporium Lohde, Hirsutella, Hymenostilbe, paecilomyces-like, Paraisaria, Syngliocladium, Tolypocladium, verticillium-like.

ELAPHOCORDYCEPS G.H. Sung & Spatafora, gen. nov. MycoBank MB504191.

Stromata singula vel nonnulla aggregata, simplicia vel ramosa. Stipes fibrosus vel tenax, raro carnosus, obscure brunneus vel olivaceo-viridulus, raro albidus, cylindricus vel sursum dilatatus. Stromata hospite insidentia vel rhizomorphis eo conjuncta. Pars fertilis clavata vel capitata, raro indistincta. Perithecia partim wel omnino in stromate immersa, perpendicularia ad superficiem. Asci cyindrici, apice inspissato. Ascosporae cylindricae, multiseptatae, maturae in cellulas diffrangentes. Anamorphe Verticillii similis vel absens.

Stromata solitary to several, simple or branched. Stipe fibrous to tough, rarely fleshy, dark brownish to greenish with olivaceous tint, rarely whitish, cylindrical to enlarging in the fertile part. Stroma connected directly to the host or indirectly through rhizomorph-like structures. Fertile part clavate to capitate, rarely undifferentiated. Perithecia partially or completely immersed in stromata, ordinal in arrangement. Asci cylindrical with thickened ascus apex. Ascospores cylindrical, multiseptate, disarticulating into part-spores.

Type: Cordyceps ophioglossoides (Ehrh.) Link

Etymology: Greek elaphe = deer, from the host fungus, Elaphomyces.

Commentary: The C. ophioglossoides clade is strongly supported (MP-BP = 71 %, ML-BP = 88 %, PP = 100 in Figs ​Figs1, 1, ​,2, 2, ​,10)10) and includes species of Cordyceps s. l. that parasitize the truffle-like genus Elaphomyces and cicada nymphs (e.g., C. inegoënsis and C. paradoxa) and beetles (e.g., C. subsessilis) (Figs ​(Figs6, 6, ​,10).10). Currently, 22 species are anticipated to be included in the C. ophioglossoides clade, of which more than 18 species are known as parasites of Elaphomyces (Mains 1957, Kobayasi & Shimizu 1960, 1963). The host affiliation of Elaphomyces parasites has long been recognized as a diagnostic character in Cordyceps classification (Massee 1895, Kobayasi 1941, 1982, Mains 1957, 1958). The oldest applicable genus name is Cordylia Fr. 1818 (Massee 1895). However, it cannot be applied to the C. ophioglossoides clade because it is a homonym of Cordylia Pers. 1807 (Mains 1958), which is also homonym of Cordyla Lour. 1790 (Leguminosae). Therefore, the genus Elaphocordyceps is proposed based on the phylogenetic placement of C. ophioglossoides and applied to the well-supported C. ophioglossoides clade. Although C. subsessilis is morphologically and ecologically distinct, the genus is well recognized by its dominant ecology as being pathogens of Elaphomyces and cicadas. The darkly pigmented, fibrous stromata with more or less olivaceous tint are also good diagnostic characters for recognizing the species of Elaphocordyceps.

Anamorphic genera: Tolypocladium, verticillium-like.

Accepted names and new combinations for Elaphocordyceps

The following taxa are accepted species of Elaphocordyceps based on their inclusion in molecular phylogenies presented herein¹ (see Table 1) or morphological descriptions matching the characters described above². Where known we provide anamorph connection for the species of Elaphocordyceps.

• ¹Elaphocordyceps capitata (Holmsk.) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504192.

  • • ≡ Sphaeria capitata Holmsk., Beata Ruris Otia Fungis Danicis 1: 38. 1790 : Fries, Syst. Mycol. 2: 324, 1823.

    • ≡ Torrubia capitata (Holmsk.: Fr.) Tul. & C. Tul., Sel. Fung. Carpol. 3: 22. 1865.

    • ≡ Cordyceps capitata (Holmsk. : Fr.) Link., Handbuch zur Erkennung der nutzbarsten und am häufigsten vorkommenden Gewächse 3: 347. 1833.

  • = Cordyceps canadensis Ellis & Everh., Bull. Torrey Bot. Club 25: 501. 1898.

    • ≡ Cordyceps capitata var. canadensis (Ellis & Everh.) Lloyd, Mycol. Writ. 5: 609. 1916.

  • = Sphaeria agariciformis Bolt., Hist. Fung. Halifax, p. 130. 1789.

    • ≡ Cordyceps agariciformis (Bolt.) Seaver, North Amer. Fl. 3: 33. 1910.

  • = Cordyceps nigriceps Peck, Bull. Torrey Bot. Club 27: 21. 1900.

  • Anamorph unknown, not growing in culture.

• ²Elaphocordyceps delicatistipitata (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504193.

  • ≡ Cordyceps delicatistipitata Kobayasi, Bull. Natn. Sci. Mus. Tokyo 5: 79. 1960 (as C. `delicatostipitata').

• ¹Elaphocordyceps fracta (Mains) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504194.

  • ≡ Cordyceps fracta Mains, Bull. Torrey Bot. Club 84: 250. 1957.

• ¹Elaphocordyceps inegoënsis (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504195.

  • ≡ Cordyceps inegoënsis Kobayasi, Bull. Natn. Sci. Mus. Tokyo 6: 292. 1963.

• ²Elaphocordyceps intermedia (S. Imai) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504196.

  • ≡ Cordyceps intermedia S. Imai, Proc. Imp. Acad. Tokyo 10: 677. 1934.

• ²Elaphocordyceps intermedia f. michinokuënsis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504197.

  • ≡ Cordyceps intermedia f. michinokuënsis Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 116. 1982.

• ¹Elaphocordyceps japonica (Lloyd) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504198.

  • • ≡ Cordyceps japonica Lloyd, Mycol. Writ. 6: 913. 1920.

  • = Cordyceps umemurae S. Imai, Trans. Sapporo Nat. Hist. Soc. 11: 32. 1929 (as C. `umemurai').

• ¹Elaphocordyceps jezoënsis (S. Imai) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504199.

  • ≡ Cordyceps jezoënsis S. Imai, Trans. Sapporo Nat. Hist. Soc. 11: 33. 1929.

• ¹Elaphocordyceps longisegmentis (Ginns) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504200.

  • ≡ Cordyceps longisegmentis Ginns, Mycologia 80: 219. 1988.

• ²Elaphocordyceps minazukiensis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504201.

  • ≡ Cordyceps minazukiensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 117. 1982.

• ²Elaphocordyceps miomoteana (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504202.

  • ≡ Cordyceps miomoteana Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 118. 1982.

• ¹Elaphocordyceps ophioglossoides (Ehrh.: Fr.) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504203.

  • • ≡ Sphaeria ophioglossoides Ehrh., in Pers., Comment de Fung. Clavaef. p. 12. 1797: Fries, Syst. Mycol. 2: 324. 1823.

    • ≡ Torrubia ophioglossoides (Ehrh. : Fr.) Tul., Sel. Fung. Carpol. 3: 20. 1865.

    • ≡ Cordyceps ophioglossoides (Ehrh. : Fr.) Link, Handbuck zur Erkennung der nutzbarsten und am häufigsten vorkommenden Gewächse 3: 347. 1833.

  • Anamorph: verticillium-like

• ²Elaphocordyceps ophioglossoides f. alba (Kobayasi & Shimizu ex Y.J. Yao) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504204.

  • ≡ Cordyceps ophioglossoides f. alba Kobayasi & Shimizu ex Y.J. Yao, in Yao, Li, Pegler & Spooner, Acta. Mycol. Sin. 14: 257. 1995.

• ²Elaphocordyceps ophioglossoides f. cuboides (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504205.

  • ≡ Cordyceps ophioglossoides f. cuboides Kobayasi, Bull. Natn. Sci. Mus. Tokyo 5: 77. 1960.

• ¹Elaphocordyceps paradoxa (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504206.

  • ≡ Cordyceps paradoxa Kobayasi, Bull. Biogeogr. Soc. Japan 9: 156. 1939.

• ²Elaphocordyceps ramosa (Teng) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504207.

  • ≡ Cordyceps ramosa Teng, Sinensia 7: 810. 1936.

• ²Elaphocordyceps rouxii (Cand.) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504208.

  • ≡ Cordyceps rouxii Cand., Mycotaxon 4: 544. 1976.

• ¹Elaphocordyceps subsessilis (Petch) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504209.

  • • ≡ Cordyceps subsessilis Petch, Trans. Brit. Mycol. Soc. 21: 39. 1937.

  • = Cordyceps facis Kobayasi & Shimizu, Trans. Mycol. Soc. Japan 23: 361. 1982.

  • Anamorph: Tolypocladium inflatum W. Gams

• ²Elaphocordyceps szemaoënsis (M. Zang) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504210.

  • ≡ Cordyceps szemaoënsis M. Zang, Acta Bot. Yunnanica 23: 295. 2001.

• ²Elaphocordyceps tenuispora (Mains) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504211.

  • ≡ Cordyceps tenuispora Mains, Bull. Torrey Bot. Club 84: 247. 1957.

• ²Elaphocordyceps toriharamontana (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504212.

  • ≡ Cordyceps toriharamontana Kobayasi, Bull. Natn. Sci. Mus. Tokyo 6: 305. 1963.

• ²Elaphocordyceps valliformis (Mains) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504213.

  • ≡ Cordyceps valliformis Mains, Bull. Torrey Bot. Club 84: 250. 1957.

• ²Elaphocordyceps valvatistipitata (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504214.

  • ≡ Cordyceps valvatistipitata Kobayasi, Bull. Natn. Sci. Mus. Tokyo 5: 81. 1960 (as C. volvatostipitata').

• ²Elaphocordyceps virens (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, comb. nov. MycoBank MB504215

  • ≡ Cordyceps virens Kobayasi, J. Jap. Bot. 58: 222. 1983.

OPHIOCORDYCEPS Petch, Trans. Brit. Mycol. Soc. 16: 73. 1931 emend. G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora

= Cordycepioideus Stifler, Mycologia 33: 83. 1941.

Stromata or subiculum darkly pigmented or rarely brightly coloured, tough, fibrous, pliant to wiry, rarely fleshy, often with aperithecial apices or lateral pads. Perithecia superficial to completely immersed, ordinal or oblique in arrangement. Asci hyaline, cylindrical, usually with thickened ascus apex, rarely fusoid to ellipsoid. Ascospores usually cylindrical, multiseptate, disarticulating into part-spores or non-disarticulating.

Type: Cordyceps blattae Petch, Trans. Brit. Mycol. Soc. 16: 74. 1931.

Anamorphic genera: Hirsutella, Hymenostilbe, Paraisaria, Syngliocladium.

Commentary: The C. unilateralis clade is strongly supported (MP-BP = 88 %, ML-BP = 88 %, PP = 1.00 in Figs ​Figs3, 3, ​,10)10) and includes the species of Ophiocordyceps (e.g., C. acicularis and C. unilateralis) (Petch 1931, 1933). The genus Ophiocordyceps was proposed by Petch (1931, 1933) for species of Cordyceps that produce non-disarticulating ascospores. The genus was not accepted by subsequent workers who reclassified the species of Ophiocordyceps as Cordyceps subg. Ophiocordyceps (Kobayasi 1941) or in multiple subgenera of Cordyceps (Mains 1958). The type of Ophiocordyceps Petch is O. blattae (= C. blattae), but it was not available for this taxonomic treatment. According to the morphological description, it fits in the present generic concept. Because O. unilateralis is a well-known species that was included in the original publication of Ophiocordyceps (Petch 1931) and because additional Ophiocordyceps species (e.g., O. acicularis) are members of this clade, we apply the name Ophiocordyceps based on the placement of O. unilateralis. The genus Ophiocordyceps includes the most morphologically diverse group of the species of Cordyceps s. l. including the members of C. subg. Neocordyceps (Figs ​(Figs6, 6, ​,10).10). For most of the species in Ophiocordyceps, the stromata are fibrous to tough or wiry to pliant in texture and darkly pigmented in at least some part of the stroma. The genus includes many species of Cordyceps s. l. that produce perithecia in subterminal regions of the stromata resulting in aperithecial apices. Of particular note, Ophiocordyceps is characterized by the dominant occurrence of Hirsutella and Hymenostilbe anamorphs (Fig. 6). Although the genus Cordycepioideus possesses thick-walled multiseptate ellipsoid ascospores and its asci lack the thickened ascus tip of most clavicipitaceous fungi (Blackwell & Gilbertson 1984, Ochiel et al. 1997), this study indicates that the genus Cordycepioideus can be merged with Ophiocordyceps according to its placement in molecular analyses and because of the Hirsutella anamorph (Fig. 10, Ochiel et al. 1997, Suh et al. 1998).

Ophiocordyceps communis Hywel-Jones & Samson, sp. nov. MycoBank MB504216. Figs 12A–G.

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Fig. 12.

A–G. Morphology of Ophiocordyceps communis. A. Stromata, bar = 10 mm. B. Arrangement of perithecia. C. Ascus with ascospores. D. Ascus and ascus apex. E. Non-disarticulating ascospores. F. Conidiophores (Hymenostilbe/Hirsutella anamorph). G. Denticles of phialide (Hymenostilbe/Hirsutella anamorph). Scale bars: A, C–G = 10 μm, B = 00 μm.

Anamorph: hirsutella/hymenostilbe-like.

Stromata ex duabus (tribus) termitis adultis oriunda, mycelio albo circumdata, filiformia; 50–100 mm sub superficie stramenti oriunda, 300–600 μm lata, albido-grisea, 70–130 mm super stramentum emergentia, 600–1000 μm lata, cuius 30–40 mm pars inferior hyphis sterilibus dematiaceis (luteo-brunneis) tomentosa; pars superior, ca 90 mm longa, fertilis, levis, griseo-brunnea vel grisea, conidia in strato griseo ferens et perithecia dense aggregata. Perithecia superficialia, subterminalia, 285–675 × 195–390 μm. Asci apice conspicue inspissato, 8-spori, filiformes, 215–250 × 15 μm. Ascosporae integrae, crassitunicatae, dilute pigmentatae, (100–)120–150(–180) × 5–6 μm. Cellulae conidiogenae hymenium hyalinum formantes, cylindricae, 10–14 × 2.7–3.3 μm, unum (raro duos) denticulos fertiles apicales ferentes. Blastoconidia hyalina, amygdaliformia, 8–9 × 2.5–3 μm. Anamorphe Hirsutellae vel Hymenostilbe similis.

Hosts two (rarely three) adult termites surrounded by loose, coarse white mycelium. Stromata filiform, 50–100 mm below ground, 300–600 μm wide, whitish-grey; 70–130 mm emerging above leaf litter, 600–1000 μm wide; lower 30–40 mm of above-ground portion usually hirsute with sterile, dematiaceous (yellow-brown) hairs becoming smooth, silver-brown to grey along terminal fertile (anamorph) part of ca 90 mm. Perithecia superficial subterminal; emerging through grey anamorph, tightly packed around the stipe, 285–675 × 195–390 μm. Asci with stout cap, 8-spored, filiform, 215–250 × 15 μm. Ascospores whole, stout, lightly pigmented (100–)120–150(–180) × 5–6 μm. Conidiogenous cells in a palisade, hyaline, cylindrical, 10–14 × 2.7–3.3 μm, solitary (rarely two), prominent, terminal denticle. Conidia hyaline, almond-shaped, 7–9 × 2.5–3 μm.

Etymology: refers to the communal nature of the stromata, i.e. the fact that 600–1000 Cordyceps stromata can be found in a small area (20 × 20 metres).

Type: Holotype: N.H.J. 10673, isotypes: N.H.J. 10674, N.H.J. 10675, N.H.J. 10676, N.H.J. 10677, all on termites; coll. R. Nasit; Khao Yai National Park, Gong Giao Nature Trail; 13 June 2000.

Commentary: Most collections of O. communis were from Khao Yai National Park with the type locality (Gong Giao Nature Trail) regularly having epizootics containing (in any one season) several hundred stromata over a 20 × 20 metre area. A few other collections were from Khao Soi Dao Wildlife Sanctuary (N.H.J. 6422 and N.H.J. 6452) and Sam Lan National Park (N.H.J. 6332). All collections of the species were from adult termites. Although surveys were made over an eighteen-year period from the far north of Thailand to the far south and from sea level to over 2500 metres, O. communis is only known from these three sites in central Thailand below 800 metres elevation.

In any year there appeared to be a single `flush' with O. communis first appearing at the start of the rainy season in May/June. The earliest collections were made in May (10 May 1994: N.H.J. 3687, N.H.J. 3681 and N.H.J. 3683, Heo Sawat Waterfall; 23 May 1996; N.H.J. 6330, Gong Giao Nature Trail). In the first 2–3 weeks after appearance, the stromata appeared slender and acicular with the lower part having a shiny silken appearance and the terminal part dull greyish. The terminal grey region consisted of a palisade of tightly packed conidiogenous cells with typically a stout elongate denticle, giving rise to a single conidium.

This anamorph is intermediate between a typical Hirsutella (e.g., Hi. formicarum, Hi. citriformis, and Hi. saussurei) and a typical Hymenostilbe (e.g., Hy. dipterigena – closer to the latter) (Figs 12F–G). The palisade of crowded conidiogenous cells is indicative of Hymenostilbe rather than Hirsutella, where conidiogenous cells are sparse and mostly immature at any given time (Fig. 12F). The denticulate nature of the conidiogenous cell also is indicative of Hymenostilbe. However, in all specimens examined to date there is no evidence of multiple denticles (five or more) usually associated with Hymenostilbe; only a few conidiogenous cells were seen with two denticles (Figs 12F–G).

The anamorph of O. communis is closest to Hy. ventricosa Hywel-Jones (Hywel-Jones 1995). That species infects cockroaches and is found attached to the under side of leaves. As with the anamorph of O. communis, Hy. ventricosa produces conidiogenous cells with only a single terminal stout denticle. Conidia of Hy. ventricosa have a pronounced point and are not typical of the clavate shape usually associated with Hymenostilbe. Similarly, the conidia of the O. communis anamorph are also fattened naviculate, appearing similar to those of Hy. ventricosa but without the processed tip.

The perithecia erupt through the dull greyish anamorph spike appearing first as longitudinal splits in the palisade of conidiogenous cells at the base of the anamorph spike. Each develops as a superficial perithecium, but they become crowded and give the overall appearance of a brown subterminal fertile region (Kobayasi 1941; Figs 12A–B). The ascus shape and the form of the ascus cap comes close to Kobayasi's Figs 12C–D (Kobayasi 1941) being typical of species in the C. unilateralis clade (with Hirsutella as an anamorph). Mature perithecia eject pigmented, whole ascospores (Fig. 12E) and often the ostiole becomes blocked with these half-emerged ascospores.

Only a few species of Cordyceps sensu Kobayasi and Mains have been reported from termites. Currently accepted species include O. koningsbergeri (= C. koningsbergeri Penz. & Sacc.), which is known only from the type locality (Java, Indonesia) (Kobayasi 1941), and C. termitophila Kobayasi & Shimizu) which is known from Japan and Taiwan (Kobayasi & Shimizu 1976, 1978). Penzig & Saccardo (1904) found O. koningsbergeri to be similar to O. myrmecophila in that it had a terminal, globose head with immersed perithecia. In this feature alone it differs significantly from O. communis with its subterminal and superficial perithecia. However, as with O. communis, Penzig & Saccardo (1904) described whole ascospores of O. koningsbergeri, which were 150 × 1 μm compared with 120–150 × 5–6 μm for O. communis. Cordyceps termitophila differs from O. communis in having a `pale rosy-grey' stroma, much smaller perithecia (280–320 × 175–190 μm for C. termitophila versus 285–675 × 195–390 μm for O. communis) and smaller ascospores (100–125 × 3 μm).

Accepted names and new combinations for Ophiocordyceps

The following taxa are accepted species of Ophiocordyceps based on their inclusion in molecular phylogenies presented herein¹ or morphological descriptions matching the characters described above². Where known, the anamorph connection is provided for the species of Ophiocordyceps.

• ¹Ophiocordyceps acicularis (Ravenel) Petch, Trans. Brit. Mycol. Soc. 18: 60. 1933.

  • • ≡ Cordyceps acicularis Ravenel, J. Linn. Soc. 1: 158. 1857.

  • = Cordyceps caroliniensis Berk. & Ravenel, Fungi Carolina 4: 29. 1855.

  • Anamorph: Hirsutella

• ¹Ophiocordyceps agriotidis (A. Kawam.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504217.

  • • ≡ Cordyceps agriotidis A. Kawam., Icon. Jap. Fungi 8: 837. 1955. [as C. `agriota']

  • Anamorph: Hirsutella

• ²Ophiocordyceps ainictos (A. Möller) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504218.

  • • ≡ Cordyceps ainictos A. Möller, Phycomyceten u. Ascomyceten, p. 226. 1901.

  • Anamorph: Hirsutella

• ²Ophiocordyceps amazonica (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504219.

  • ≡ Cordyceps amazonica Henn., Hedwigia 43: 247. 1904.

• ²Ophiocordyceps amazonica var. neoamazonica (Kobayasi & Hara) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504220.

  • ≡ Cordyceps amazonica var. neoamazonica Kobayasi & Hara, J. Jap. Bot. 57: 17. 1982.

• ¹Ophiocordyceps aphodii (Mathieson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504221.

  • • ≡ Cordyceps aphodii Mathieson, Trans. Brit. Mycol. Soc. 32: 134. 1949.

  • Anamorph: Hirsutella

• ²Ophiocordyceps appendiculata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504222.

  • ≡ Cordyceps appendiculata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 6. 1983.

• ²Ophiocordyceps arachneicola (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504223.

  • ≡ Cordyceps arachneicola Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku, Sect. B,: 123. 1941.

• ²Ophiocordyceps arbuscula (Teng) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504225

  • ≡ Cordyceps arbuscula Teng, Sinensia 7: 812. 1936.

• ²Ophiocordyceps armeniaca (Berk. & M.A. Curtis) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504226.

  • ≡ Cordyceps armeniaca Berk. & M.A. Curtis, J. Linn. Soc. 1: 158. 1857.

• ²Ophiocordyceps asyuënsis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504227.

  • ≡ Cordyceps asyuënsis Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 138. 1980.

• ²Ophiocordyceps aurantia (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504228.

  • ≡ Cordyceps aurantia Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 125. 1980.

• ²Ophiocordyceps australis (Speg.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504229.

  • ≡ Cordyceps unilateralis var. australis Speg., Anales Soc. Ci. Argent. 12: 215. 1881.

  • ≡ Cordyceps australis (Speg.) Speg., Syll. Fung. 2: 571. 1883.

• ²Ophiocordyceps barnesii (Thwaites) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504230.

  • ≡ Cordyceps barnesii Thwaites, J. Linn. Soc. 14: 110. 1875.

  • ≡ Torrubia barnesii (Thwaites) Ces., Atti Accad. Sci. Fis. Mat., Napoli 8: 14. 1879.

• ²Ophiocordyceps bicephala (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504231.

  • ≡ Cordyceps bicephala Berk., J. Bot. (Hooker) 8: 278. 1856.

• ²Ophiocordyceps bispora (Stifler) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504232.

  • • ≡ Cordycepioideus bisporus Stifler, Mycologia 33: 85. 1941.

  • Anamorph: Hirsutella

• ²Ophiocordyceps blattae (Petch) Petch, Trans. Brit. Mycol. Soc. 16: 74. 1931.

  • ≡ Cordyceps blattae Petch, Trans. Brit. Mycol. Soc. 10: 35. 1924.

• ¹Ophiocordyceps brunneipunctata (Hywel-Jones) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504233.

  • • ≡ Cordyceps brunneipunctata Hywel-Jones, Mycol. Res. 99: 1195. 1995. [as C. `brunneapunctata']

  • Anamorph: Hirsutella

• ²Ophiocordyceps caloceroides (Berk. & M.A. Curtis) Petch, Trans. Brit. Mycol. Soc. 18: 63. 1933.

  • • ≡ Cordyceps caloceroides Berk. & M.A. Curtis, J. Linn. Soc. 10: 375. 1868.

  • = Cordyceps wittii Henn., Bot. Jahrb. Syst. 23: 539. 1897.

• ²Ophiocordyceps cantharelloides (Samson & H.C. Evans) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504234.

  • ≡ Cordyceps cantharelloides Samson & H.C. Evans, Proc. Indian Acad. Sci., Pl. Sci. 94: 312. 1985.

• ²Ophiocordyceps carabidicola (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504235.

  • ≡ Cordyceps carabidicola Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 85. 1980. [as C. `carabidiicola']

• ²Ophiocordyceps cicadicola (Teng) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504236.

  • ≡ Cordyceps cicadicola Teng, Sinensia 6: 191. 1935.

• ²Ophiocordyceps clavata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504237.

  • ≡ Cordyceps clavata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 140. 1980.

• ²Ophiocordyceps clavulata (Schwein.) Petch, Trans. Brit. Mycol. Soc. 18: 53. 1933.

  • • ≡ Sphaeria clavulata Schwein., Trans. Amer. Philos. Soc. New Ser. 4, 188. 1832.

    • ≡ Xylaria clavulata (Schwein.) Berk. & M. A. Curtis, J. Linn. Soc. 10: 380. 1868.

    • ≡ Torrubia clavulata (Schwein.) Peck, Ann. Rep. N. Y. State Mus. 28: 70. 1876.

    • ≡ Cordyceps clavulata (Schwein.) Ellis & Everh., North Amer. Pyrenom. p. 61. 1892.

  • = Cordyceps pistillariiformis Berk. & Broome, Ann. Mag. Nat. Hist. Ser. 3, 7: 451. 1861 [as C. `pistillariaeformis'].

    • ≡ Torrubia pistillariiformis (Berk. & Broome) Cooke, Handb. Brit. Fungi 2: 771. 1871.

  • Anamorph: Hymenostilbe lecaniicola (Jaap) Mains

• ¹Ophiocordyceps coccidiicola (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504238.

  • ≡ Cordyceps coccidiicola Kobayasi, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 4: 57. 1978.

• ²Ophiocordyceps coccigena (Tul. & C. Tul.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504239.

  • ≡ Torrubia coccigena Tul. & C. Tul., Sel. Fung. Carpol. 3: 19. 1865.

  • ≡ Cordyceps coccigena (Tul. & C. Tul.) Sacc., Michelia 1: 320. 1879.

• ²Ophiocordyceps cochlidiicola (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504240.

  • ≡ Cordyceps cochlidiicola Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 128. 1980.

• ¹Ophiocordyceps communis Hywel-Jones & Samson, sp. nov., see above.

  • Anamorph: hirsutella/hymenostilbe-like

• ²Ophiocordyceps corallomyces (A. Möller) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504241.

  • ≡ Cordyceps corallomyces A. Möller, Phycomyceten u. Ascomyceten, p. 217. 1901.

• ²Ophiocordyceps crassispora (M. Zang, D.R. Yang & C.D. Li) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504242.

  • ≡ Cordyceps crassispora M. Zang, D.R. Yang & C.D. Li, Mycotaxon 37: 58. 1990.

• ²Ophiocordyceps crinalis (Ellis ex Lloyd) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504243.

  • ≡ Cordyceps crinalis Ellis ex Lloyd, Mycol. Writ. 6: 912. 1920.

• ²Ophiocordyceps cucumispora (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504244.

  • ≡ Cordyceps cucumispora H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 442. 1982.

  • Anamorph: Hirsutella ovalispora H.C. Evans & Samson

• ²Ophiocordyceps cucumispora var. dolichoderi (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504245.

  • • ≡ Cordyceps cucumispora var. dolichoderi H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 445. 1982.

  • Anamorph: Hirsutella ovalispora var. dolichoderi H.C. Evans & Samson

• ²Ophiocordyceps curculionum (Tul. & C. Tul.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504246.

  • • ≡ Torrubia curculionum Tul. & C. Tul., Sel. Fung. Carpol. 3: 20. 1865.

    • ≡ Cordyceps curculionum (Tul. & C. Tul.) Sacc., Michelia 1: 320. 1879.

    • ≡ Cordyceps bicephala subsp. curculionum (Tul. & C. Tul.) Moureau, Mém. Inst. Roy. Colon. Belge 7: 50. 1949.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps cylindrostromata (Z.Q. Liang, A.Y. Liu & M.H. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504247.

  • ≡ Cordyceps cylindrostromata Z.Q. Liang, A.Y. Liu & M.H. Liu, Fungal Diversity 14: 97. 2003.

• ²Ophiocordyceps dayiensis (Z.Q. Liang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504248

  • ≡ Cordyceps dayiensis Z.Q. Liang, Fungal Diversity 12: 131. 2003.

• ²Ophiocordyceps dermapterigena (Z.Q. Liang, A.Y. Liu & M.H. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504249.

  • ≡ Cordyceps dermapterigena Z.Q. Liang, A.Y. Liu & M.H. Liu, Fungal Diversity 14: 96. 2003 (as C. `dermapteoigena').

• ²Ophiocordyceps dipterigena (Berk. & Broome) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504250.

  • • ≡ Cordyceps dipterigena Berk. & Broome, J. Linn. Soc. 14: 111. 1875.

  • = Cordyceps muscicola A. Möller, Phycomyceten u. Ascomyceten, p. 221. 1901.

  • = Cordyceps surinamensis Henn., Hedwigia 41: 169 1902.

  • = Cordyceps oumensis Höhn., Sitzungsber. Kaiserl. Akad. Wiss. Wien 118: 309. 1909.

  • = Cordyceps ouwensii Höhn., Sitzungsber. Kaiserl. Akad. Wiss. Wien 118: 309. 1909.

  • = Cordyceps thwaitesii Lloyd, Mycol. Writ. 6: 1060. 1921.

  • = Cordyceps opposita Syd., Bot. Jahrb. Syst. 57: 325. 1922.

  • Anamorph: Hymenostilbe dipterigena Petch

• ²Ophiocordyceps discoideicapitata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504251.

  • ≡ Cordyceps discoideicapitata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 85. 1982 (as C. `discoideocapitata').

• ²Ophiocordyceps ditmarii (Quél.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504252.

  • ≡ Cordyceps ditmarii Quél., Bull. Soc. Bot. France 24: 330 1877. [as C. ditmari]

    • Anamorph: Hymenostilbe

• ²Ophiocordyceps dovei (Rodway) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504253.

  • • ≡ Cordyceps dovei Rodway, Paper Proc. Roy. Soc. Tasmania for 1898–1899, p. 101. 1900.

  • = Cordyceps aemonae Lloyd, Mycol. Notes 2: 932. 1920.

  • Anamorph: hirsutella-like

• ²Ophiocordyceps elateridicola (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504255.

  • ≡ Cordyceps elateridicola Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 4. 1983.

• ²Ophiocordyceps elongata (Petch) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504256.

  • • ≡ Cordyceps elongata Petch, Trans. Brit. Mycol. Soc. 21: 47. 1937.

  • Anamorph: Hirsutella

• ²Ophiocordyceps elongatiperitheciata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504257.

  • ≡ Cordyceps elongatiperitheciata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 126. 1980 (as C. `elongatoperitheciata').

• ²Ophiocordyceps elongatistromata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504258.

  • ≡ Cordyceps elongatistromata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 12. 1983 (as C. `elongatostromata').

• ²Ophiocordyceps emeiensis (A.Y. Liu & Z.Q. Liang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504259.

  • ≡ Cordyceps emeiensis A.Y. Liu & Z.Q. Liang, Mycosystema 16: 139. 1997.

• ²Ophiocordyceps engleriana (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504260.

  • ≡ Cordyceps engleriana Henn., Bot. Jahrb. Syst. 23: 538. 1897.

  • Anamorph: Hymenostilbe

• ¹Ophiocordyceps entomorrhiza (Dicks.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504261.

  • • ≡ Sphaeria entomorrhiza Dicks., Plant. Crypt. Brit., Fasc. 1: 22. 1785.

    • ≡ Cordyceps entomorrhiza (Dicks.) Fr., Obs. Mycol. 2: 317. 1818.

    • ≡ Torrubia entomorrhiza (Dicks.) Tul. & C. Tul, Sel. Fung. Carpol. 3: 13. 1865.

  • = Torrubia cinerea Tul. & C. Tul., Sel. Fung. Carpol. 3: 14. 1865.

    • ≡ Cordyceps cinerea (Tul. & C. Tul.) Sacc., Michelia 1: 320. 1879.

  • = Cordyceps carabi Quél., Comp. Rend. Assoc. Franç. Avancem. Sci. 2: 452. 1898.

  • Anamorph: Hirsutella eleutheratorum (Nees) Petch

• ²Ophiocordyceps evdogeorgiae (Koval) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504262.

  • ≡ Cordyceps evdogeorgiae Koval, Bot. Mater. Otd. Sporov. Rast. 14: 160. 1961.

• ²Ophiocordyceps falcata (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504263.

  • ≡ Cordyceps falcata Berk., J. Bot. (Hooker) 6: 211. 1854 [Decad. Fung. No. 479].

    • Anamorph: Stilbella

• ²Ophiocordyceps falcatoides (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504264.

  • ≡ Cordyceps falcatoides Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 91. 1980.

• ²Ophiocordyceps fasciculatistromata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504265.

  • ≡ Cordyceps fasciculatistromata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 83. 1982 (as C. `fasciculatostromata').

• ²Ophiocordyceps ferruginosa (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504266.

  • ≡ Cordyceps ferruginosa Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 139. 1980.

• ²Ophiocordyceps filiformis (Moureau) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504267.

  • ≡ Cordyceps filiformis Moureau, Mém. Inst. Roy. Colon. Belge 7: 14. 1949.

• ²Ophiocordyceps formicarum (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504268

  • • ≡ Cordyceps formicarum Kobayasi, Bull. Biogeogr. Soc. Japan 9: 28. 1939.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps forquignonii (Quél.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504269.

  • • ≡ Cordyceps forquignonii Quél., 16th Suppl. Champ. Jura et Vosges, p. 6. 1887.

  • Anamorph: Hymenostilbe muscarium Petch

• ²Ophiocordyceps furcicaudata (Z.Q. Liang, A.Y. Liu & M.H. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504270.

  • ≡ Cordyceps furcicaudata Z.Q. Liang, A.Y. Liu & M.H. Liu, Fungal Diversity 14: 95. 2003 (as C. `furcicaodata').

• ²Ophiocordyceps gansuënsis (K. Zhang, C. Wang & M. Yan) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504271.

  • ≡ Cordyceps gansuënsis K. Zhang, C. Wang & M. Yan, Trans. Mycol. Soc. Japan 30: 295. 1989.

• ²Ophiocordyceps geniculata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504272.

  • ≡ Cordyceps geniculata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 85. 1980.

• ²Ophiocordyceps gentilis (Ces.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504273.

  • ≡ Torrubia gentilis Ces., Atti Accad. Sci. Fis. Mat., Napoli, 8: 14. 1879.

  • ≡ Cordyceps gentilis (Ces.) Sacc., Syll. Fung. 2: 569. 1883.

• ²Ophiocordyceps glaziovii (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504274.

  • ≡ Cordyceps glaziovii Henn., Naturw. Wochenschr. 6: 318. 1896.

• ²Ophiocordyceps goniophora (Speg.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504275.

  • ≡ Cordyceps goniophora Speg., Bol. Acad. Nac. Ci. Córdoba 11: 307 1889.

• ²Ophiocordyceps gracilioides (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504276.

  • • ≡ Cordyceps gracilioides Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku, Sect. B, 5: 140. 1941.

  • Anamorph: paecilomyces-like

• ¹Ophiocordyceps gracilis (Grev.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504277.

  • • ≡ Xylaria gracilis Grev., Scot. Crypt. Fl. 2. t. 8. 1824.

    • ≡ Cordyceps gracilis (Grev.) Durieu & Mont., Fl. Algérie Crypt. 1: 449. 1846.

  • = Cordyceps mawleyi Westwood, Gard. Chron. Ser. 3, 9: 553. 1891.

  • Anamorph: Paraisaria dubia (Delacr.) Samson & B.L. Brady

• ²Ophiocordyceps gryllotalpae Petch, Trans. Brit. Mycol. Soc. 25: 255. 1941.

  • ≡ Cordyceps gryllotalpae Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku 5: 70. 1942 [non Lloyd 1924].

  • ≡ Cordyceps koreana Kobayasi, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 7: 8. 1981.

• ¹Ophiocordyceps heteropoda (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504278.

  • ≡ Cordyceps heteropoda Kobayasi, Bull. Biogeogr. Soc. Japan 9: 158. 1939.

• ²Ophiocordyceps hiugensis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504279.

  • ≡ Cordyceps hiugensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 3. 1983.

• ²Ophiocordyceps huberiana (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504280.

  • ≡ Cordyceps huberiana Henn., Hedwigia 48: 105. 1909.

• ²Ophiocordyceps humbertii (C.P. Robin) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504281.

  • • ≡ Cordyceps humbertii C.P. Robin, in Tul. & C. Tul., Sel. Fung. Carpol. 3: 18. 1865 (as C. `humberti').

  • Anamorph: Hirsutella saussurei (Cooke) Speare

• ²Ophiocordyceps insignis (Cooke & Ravenel) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504282.

  • ≡ Cordyceps insignis Cooke & Ravenel, Grevillea 12: 38. 1883.

• ¹Ophiocordyceps irangiensis (Moureau) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504283.

  • • ≡ Cordyceps irangiensis Moureau, Lejeunia, Mém. 15: 33. 1961.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps japonensis (Hara) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504284.

  • ≡ Cordyceps japonensis Hara, Bot. Mag. Tokyo 28: 351. 1914.

• ²Ophiocordyceps jiangxiensis (Z.Q. Liang, A.Y. Liu & Yong C. Jiang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504285.

  • ≡ Cordyceps jiangxiensis Z.Q. Liang, A.Y. Liu & Yong C. Jiang, Mycosystema 20: 30. 2001.

• ²Ophiocordyceps jinggangshanensis (Z.Q. Liang, A.Y. Liu & Yong C. Jiang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504286.

  • ≡ Cordyceps jinggangshanensis Z.Q. Liang, A.Y. Liu & Yong C. Jiang, Mycosystema 20: 307. 2001.

• ²Ophiocordyceps kangdingensis (M. Zang & N. Kinjo) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504287.

  • ≡ Cordyceps kangdingensis M. Zang & N. Kinjo, Mycotaxon 66: 221. 1998.

• ²Ophiocordyceps kniphofioides (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504288.

  • • ≡ Cordyceps kniphofioides H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 434. 1982.

  • Anamorph: Hirsutella stilbelliformis H.C. Evans & Samson

• ²Ophiocordyceps kniphofioides var. dolichoderi (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504289.

  • • ≡ Cordyceps kniphofioides var. dolichoderi H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 437. 1982.

  • Anamorph: Hirsutella stilbelliformis var. dolichoderi H.C. Evans & Samson

• ²Ophiocordyceps kniphofioides var. monacidis (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504290.

  • • ≡ Cordyceps kniphofioides var. monacidis H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 439. 1982.

  • Anamorph: Hirsutella stilbelliformis var. monacidis H.C. Evans & Samson

• ²Ophiocordyceps kniphofioides var. ponerinarum (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504291.

  • • ≡ Cordyceps kniphofioides var. ponerinarum H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 79: 441. 1982.

  • Anamorph: Hirsutella stilbelliformis var. ponerinarum H.C. Evans & Samson

• ²Ophiocordyceps koningsbergeri (Penz. & Sacc.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504292.

  • ≡ Cordyceps koningsbergeri Penz. & Sacc., Malpighia 11: 522. 1897.

• ¹Ophiocordyceps konnoana (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504293.

  • ≡ Cordyceps konnoana Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 84. 1980.

• ²Ophiocordyceps lachnopoda (Penz. & Sacc.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504295.

  • ≡ Cordyceps lachnopoda Penz. & Sacc., Malpighia 11: 521. 1897.

• ²Ophiocordyceps larvarum (Westwood) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504397.

  • • ≡ Sphaeria larvarum Westwood, Proc. Entomol. Soc. Lond. 2: 6. 1836.

    • ≡ Cordyceps larvarum (Westwood) Olliff, Gaz. New South Wales 6: 410. 1895.

  • = Cordyceps huegelii Corda, Icon. Fung. 4: 44. 1840.

• ²Ophiocordyceps lloydii (H.S. Fawc.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504296.

  • • ≡ Cordyceps lloydii H.S. Fawc., Ann. Mag. Nat. Hist., Ser. 5, 18: 317. 1886.

  • = Cordyceps sheeringii Massee, Ann. Bot. 5: 510. 1890.

  • = Cordyceps subdiscoidea Henn., Hedwigia 41: 168. 1902.

  • Anamorph: Hymenostilbe formicarum Petch

• ²Ophiocordyceps lloydii var. binata (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504297.

  • ≡ Cordyceps lloydii var. binata H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 82: 133. 18: 31. 1984.

• ¹Ophiocordyceps longissima (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504298.

  • ≡ Cordyceps longissima Kobayasi, Bull. Natn. Sci. Mus. Tokyo 6: 300. 1963.

• ²Ophiocordyceps lutea (Moureau) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504299.

  • • ≡ Cordyceps lutea Moureau, Mém. Inst. Roy. Colon. Belge 7: 41. 1949.

  • Anamorph: Hymenostilbe sulphurea Samson & H.C. Evans

• ²Ophiocordyceps macularis Mains, Proc. Amer. Philos. Soc. 74: 269. 1934.

  • ≡ Cordyceps macularis (Mains) Mains, Pap. Michigan Acad. Sci. 25: 82. 1940.

• ¹Ophiocordyceps melolonthae (Tul. & C. Tul.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504300.

  • ≡ Torrubia melolonthae Tul. & C. Tul., Sel. Fung. Carpol. 3: 12. 1865.

  • ≡ Cordyceps melolonthae (Tul. & C. Tul.) Sacc., Michelia 1: 320. 1879.

• ²Ophiocordyceps melolonthae var. rickii (Lloyd) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504301.

  • ≡ Cordyceps rickii Lloyd, Mycol. Writ. 6: 914. 1920.

  • ≡ Cordyceps melolonthae var. rickii (Lloyd) Mains, Mycologia 50: 198 1958.

• ²Ophiocordyceps michiganensis (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504302.

  • ≡ Cordyceps michiganensis Mains, Proc. Amer. Philos. Soc. 74: 266. 1934.

• ²Ophiocordyceps minutissima (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504303.

  • ≡ Cordyceps minutissima Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 77. 1980.

• ²Ophiocordyceps monticola (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504304.

  • ≡ Cordyceps monticola Mains, Mycologia 32: 310. 1940.

• ²Ophiocordyceps mrciensis (Aung, J.C. Kang, Z.Q.Liang, Soytong & K.D. Hyde) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504305.

  • ≡ Cordyceps mrciensis Aung, J.C. Kang, Z.Q.Liang, Soytong & K.D. Hyde, Mycotaxon 97: 236. 2006.

• ²Ophiocordyceps multiaxialis (M. Zang & Kinjo) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504306.

  • ≡ Cordyceps multiaxialis M. Zang & Kinjo, Mycotaxon 66: 224. 1998.

• ²Ophiocordyceps myrmecophila (Ces.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504307.

  • • ≡ Cordyceps myrmecophila Ces., Bot. Zeitung 4: 877. 1846.

    • ≡ Torrubia myrmecophila (Ces.) Tul. & C. Tul., Sel. Fung. Carpol. 3: 18. 1865.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps neovolkiana (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504308.

  • ≡ Cordyceps neovolkiana Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku, Sect. B, 5: 169. 1941.

  • Anamorph: Hirsutella neo-volkiana Kobayasi

• ²Ophiocordyceps nepalensis (M. Zang & Kinjo) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504309.

  • ≡ Cordyceps nepalensis M. Zang & Kinjo, Mycotaxon 66: 224. 1998.

• ²Ophiocordyceps nigra (Samson, H.C. Evans & E.S. Hoekstra) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504310.

  • ≡ Cordyceps nigra Samson, H.C. Evans & E.S. Hoekstra, Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 85: 596. 1982.

• ¹Ophiocordyceps nigrella (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504311.

  • ≡ Cordyceps nigrella Kobayasi & Shimizu, Icon. Veg. Wasps and Plant Worms p. 145. 1983.

  • ≡ Cordyceps nigra Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo 9(1): 13. 1983 [non Samson et al. 1982]

• ²Ophiocordyceps nigripes (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504312.

  • ≡ Cordyceps nigripes Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 116. 1982 (as C. `nigripoda').

• ¹Ophiocordyceps nutans (Pat.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504313.

  • • ≡ Cordyceps nutans Pat., Bull. Soc. Mycol. France 3: 127. 1887.

    • ≡ Cordyceps bicephala subsp. nutans (Pat.) Moureau, Mém. Inst. Roy. Colon. Belge 7: 47. 1949.

  • Anamorph: Hymenostilbe nutans Samson & H.C. Evans

• ²Ophiocordyceps obtusa (Penz. & Sacc.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504314.

  • ≡ Cordyceps obtusa Penz. & Sacc., Malpighia 11: 523. 1897.

• ²Ophiocordyceps octospora (M. Blackwell & Gilb.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504315.

  • • ≡ Cordycepioideus octosporus M. Blackwell & Gilb., Mycologia 73: 358. 1981.

  • Anamorph: Hirsutella

• ²Ophiocordyceps odonatae (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504316.

  • • ≡ Cordyceps odonatae Kobayasi, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 7: 6. 1981.

  • Anamorph: Hymenostilbe odonatae Kobayasi

• ²Ophiocordyceps osuzumontana (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504317.

  • ≡ Cordyceps osuzumontana Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 77. 1980.

• ²Ophiocordyceps owariensis (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504318.

  • ≡ Cordyceps owariensis Kobayasi, Bull. Biogeogr. Soc. Japan 9: 166. 1939.

• ²Ophiocordyceps owariensis f. viridescens (Uchiyama & Udagawa) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504319.

  • • ≡ Cordyceps owariensis f. viridescens Uchiyama & Udagawa, Mycoscience 43: 136. 2002.

  • Anamorph: Nomuraea owariensis Uchiyama & Udagawa

• ²Ophiocordyceps oxycephala (Penz. & Sacc.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504320.

  • • ≡ Cordyceps oxycephala Penz. & Sacc., Malpighia 11: 521. 1897.

    • ≡ Cordyceps sphecocephala f. oxycephala (Penz. & Sacc.) Kobayasi, Trans. Mycol. Soc. Japan 23: 361. 1982.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps paludosa Mains, Proc. Amer. Philos. Soc. 74: 269. 1934.

  • • ≡ Cordyceps paludosa (Mains) Mains, Pap. Michigan Acad. Sci. 25: 83. 1940.

  • Anamorph: Polycephalomyces paludosus Mains

• ²Ophiocordyceps pentatomae (Koval) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504321.

  • • ≡ Cordyceps pentatomae Koval, Nov. Sist. Niz. Rast. 1: 166. 1964. (as C. `pentatomi')

  • Anamorph: Hirsutella

• ²Ophiocordyceps petchii (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504322.

  • ≡ Cordyceps petchii Mains, Bull. Torrey Bot. Club 86: 47. 1959.

  • ≡ Cordyceps ramosa Petch, Trans. Brit. Mycol. Soc 21: 42. 1937 [non Teng 1936].

• ²Ophiocordyceps proliferans (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504323.

  • ≡ Cordyceps proliferans Henn., Hedwigia 43: 248. 1904.

• ²Ophiocordyceps pseudolloydii (H.C. Evans & Samson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504324.

  • ≡ Cordyceps pseudolloydii H.C. Evans & Samson, Trans. Brit. Mycol. Soc. 82: 133. 1984.

• ²Ophiocordyceps pseudolongissima (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504325.

  • ≡ Cordyceps pseudolongissima Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 8: 119. 1982.

• ²Ophiocordyceps purpureostromata (Kobayasi) ex G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504326.

  • ≡ Cordyceps purpureostromata Kobayasi, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 136. 1980. Type Shimizu No. 128, preserved in TNS; therefore the basionym was valid from the beginning.

• ²Ophiocordyceps purpureostromata f. recurvata (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504327.

  • ≡ Cordyceps purpureostromata f. recurvata Kobayasi, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 138. 1980.

• ¹Ophiocordyceps ravenelii (Berk. & M.A. Curtis) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504328.

  • ≡ Cordyceps ravenelii Berk. & M.A. Curtis, J. Linn. Soc. 1: 159. 1857.

• ¹Ophiocordyceps rhizoidea (Höhn.) Petch, Trans. Brit. Mycol. Soc. 1: 74. 1931.

  • • ≡ Cordyceps rhizoidea Höhn., Sitzungsber. Kaiserl. Akad. Wiss. Wien 118: 307. 1909.

  • Anamorph: Hirsutella

• ²Ophiocordyceps ridleyi (Massee) Kobayasi, Bull. Biogeogr. Soc. Japan 9: 271. 1939.

  • ≡ Cordyceps ridleyi Massee, Bull. Misc. Inform. Roy. Bot. Gard. Kew, p. 173. 1899.

• ¹Ophiocordyceps robertsii (Hook.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504329.

  • ≡ Sphaeria robertsii Hook. Icon. Plant. 1 pl. 6. 1837.

  • ≡ Cordyceps robertsii (Hook.) Berk., Fl. New Zealand 2: 202. 1855.

• ²Ophiocordyceps rubripunctata (Moureau) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504331.

  • • ≡ Cordyceps rubripunctata Moureau, Mém. Inst. Roy. Colon. Belge 7: 26. 1949.

  • Anamorph: Hirsutella rubripunctata Samson, H.C. Evans & Hoekstra

• ²Ophiocordyceps rubiginosiperitheciata (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504332.

  • ≡ Cordyceps rubiginosiperitheciata Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 14. 1983 (as C. `rubiginosoperitheciata').

• ²Ophiocordyceps ryogamiensis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504333.

  • ≡ Cordyceps ryogamiensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 9: 4. 1983.

• ²Ophiocordyceps salebrosa (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504334.

  • ≡ Cordyceps salebrosa Mains, Mycologia 39: 541. 1947.

• ²Ophiocordyceps scottiana (Olliff) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504335.

  • ≡ Cordyceps scottiana Olliff, Agric. Gaz. New South Wales 6: 407. 1895.

• ²Ophiocordyceps selkirkii (Olliff) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504338.

  • ≡ Cordyceps selkirkii Olliff, Agric. Gaz. New South Wales 6: 411. 1895.

• ²Ophiocordyceps sichuanensis (Z.Q. Liang & B. Wang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504339.

  • ≡ Cordyceps sichuanensis Z.Q. Liang & B. Wang, Fungal Diversity 12: 129. 2003.

• ¹Ophiocordyceps sinensis (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504340.

  • • ≡ Sphaeria sinensis Berk., J. Bot. (Hooker) 2: 207. 1843.

    • ≡ Cordyceps sinensis (Berk.) Sacc., Michelia 1: 320. 1879.

  • Anamorph: Hirsutella sinensis X.J. Liu, Y.L. Guo, Y.X. Yu & W. Zeng

• ²Ophiocordyceps smithii (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504341.

  • ≡ Cordyceps smithii Mains, J. Elisha Mitchell Sci. Soc. 55: 127. 1939.

• ¹Ophiocordyceps sobolifera (Hill ex Watson) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504342.

  • • ≡ Clavaria sobolifera Hill ex Watson, Philos. Trans. Roy. Soc. Lond. 53: 271. 1763.

    • ≡ Sphaeria sobolifera (Hill ex Watson) Berk., J. Bot. (Hooker) 2: 207. 1843.

    • ≡ Torrubia sobolifera (Hill ex Watson) Tul. & C. Tul., Sel. Fung. Carpol. 3: 10. 1865.

    • ≡ Cordyceps sobolifera (Hill ex Watson) Berk. & Broome, J. Linn. Soc. 14: 110. 1875.

  • Anamorph: Beauveria sobolifera Z.Y. Liu, Z.Q. Liang, Whalley, A.Y. Liu & Y.J. Yao

• ¹Ophiocordyceps sphecocephala (Klotzsch ex Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504343.

  • • ≡ Sphaeria sphecocephala Klotzsch ex Berk., J. Bot. (Hooker) 2: 206. 1843.

    • ≡ Torrubia sphecocephala (Klotzsch ex Berk.) Tul. & C. Tul., Sel. Fung. Carpol. 3: 18. 1865.

    • ≡ Cordyceps sphecocephala (Klotzsch ex Berk.) Berk. & M.A. Curtis, in Berkeley, J. Linn. Soc., Bot. 10: 376. 1868.

  • Anamorph: Hymenostilbe

• ²Ophiocordyceps stipillata (Z.Q. Liang & A.Y. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504344.

  • ≡ Cordyceps stipillata Z.Q. Liang & A.Y. Liu, Mycosystema 21: 11. 2002.

• ¹Ophiocordyceps stylophora (Berk. & Broome) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504345.

  • • ≡ Cordyceps stylophora Berk. & Broome, J. Linn. Soc. 1: 158. 1857.

  • Anamorph: Hirsutella stylophora Mains

• ²Ophiocordyceps subflavida (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504346.

  • ≡ Cordyceps subflavida Mains, Bull. Torrey Bot. Club 86: 47. 1959.

  • ≡ Cordyceps albida Pat. & Gaillard, Bull. Soc. Mycol. France 7: 116. 1888 [non Berk. & M.A. Curtis ex Cooke 1884].

• ²Ophiocordyceps subunilateralis (Henn.) Kobayasi, Bull. Biogeogr. Soc. Japan 9: 271. 1939.

  • ≡ Cordyceps subunilateralis Henn., Hedwigia 41: 168. 1902.

• ¹Ophiocordyceps superficialis (Peck) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504347.

  • ≡ Torrubia superficialis Peck, Rep. N. Y. State Botanist 28: 70. 1876.

  • ≡ Cordyceps superficialis (Peck) Sacc., Syll. Fung. 2: 574. 1883.

• ²Ophiocordyceps superficialis f. crustacea (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504348.

  • ≡ Cordyceps superficialis f. crustacea Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 82. 1980.

• ²Ophiocordyceps takaoënsis (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504349.

  • ≡ Cordyceps sobolifera var. takaoënsis Kobayasi, Bull. Biogeogr. Soc. Japan 9: 165. 1939.

  • ≡ Cordyceps takaoënsis (Kobayasi) Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku, Sect. B, 5: 130. 1941.

• ²Ophiocordyceps taylorii (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504350.

  • ≡ Sphaeria taylorii Berk., J. Bot. (Hooker) 2: 209. 1843 (as S. `taylori').

  • ≡ Cordyceps taylorii (Berk.) Sacc., Michelia 1: 320. 1879.

• ²Ophiocordyceps thyrsoides (A. Möller) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504351.

  • ≡ Cordyceps thyrsoides A. Möller, Phycomyceten u. Ascomyceten, p. 221. 1901.

• ¹Ophiocordyceps tricentri (Yasuda) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504352.

  • • ≡ Cordyceps tricentri Yasuda, in Lloyd, Mycol. Writ. 4: 568. 1915 (as C. `tricentrus').

  • = Cordyceps aphrophorae Yasuda, Bot. Mag. Tokyo 36: 51. 1922.

• ²Ophiocordyceps uchiyamae (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504353.

  • ≡ Cordyceps uchiyamae Kobayasi & Shimizu, Bull. Natn. Sci. Mus. Tokyo, Ser. B, 6: 125. 1980.

• ¹Ophiocordyceps unilateralis (Tul. & C. Tul.) Petch, Trans. Brit. Mycol. Soc. 16: 74. 1931.

  • • ≡ Torrubia unilateralis Tul. & C. Tul., Sel. Fung. Carpol. 3: 18. 1865.

    • ≡ Cordyceps unilateralis (Tul. & C. Tul.) Sacc., Syll. Fung. 2:570. 1883.

  • = Torrubia formicivora Tul. & C. Tul., Sel. Fung. Carpol. 3: 18. 1865.

    • ≡ Cordyceps formicivora (Tul. & C. Tul.) J. Schröt., Krypt.-Fl. Schlesien 3(2) 27. 1894.

  • Anamorph: Hirsutella formicarum Petch

• ²Ophicordyceps unilateralis var. clavata Kobayasi, Bull. Biogeogr. Soc. Japan 9: 272. 1939.

  • ≡ Cordyceps unilateralis var. clavata (Kobayasi) Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku, Sect. B, 5: 78. 1941.

• ¹Ophiocordyceps variabilis (Petch) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504354.

  • • ≡ Cordyceps variabilis Petch, Trans. Brit. Mycol. Soc. 21: 42. 1937.

  • = Cordyceps viperina Mains, Mycologia 29: 74. 1937.

  • Anamorph: Syngliocladium

• ²Ophiocordyceps voeltzkowii (Henn.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504355.

  • ≡ Cordyceps voeltzkowii Henn., in Voeltzkow, Reise Ostafrica 3: 29. 1908.

• ²Ophiocordyceps volkiana (A. Möller) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504356.

  • • ≡ Cordyceps volkiana A. Möller, Phycomyceten u. Ascomyceten, p. 233. 1901.

  • Anamorph: Hirsutella

• ²Ophiocordyceps wuyishanensis (Z.Q. Liang, A.Y. Liu & J.Z. Huang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504357.

  • • ≡ Cordyceps wuyishanensis Z.Q. Liang, A.Y. Liu & J.Z. Huang, Mycosystema 21: 162. 2002.

  • Anamorph: paecilomyces-like

• ¹Ophiocordyceps yakusimensis (Kobayasi) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504358.

  • ≡ Cordyceps yakusimensis Kobayasi, Bull. Natn. Sci. Mus. Tokyo 6: 302. 1963.

• ²Ophiocordyceps zhangjiajiensis (Z.Q. Liang & A.Y. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, comb. nov. MycoBank MB504359.

  • • ≡ Cordyceps zhangjiajiensis Z.Q. Liang & A.Y. Liu, Mycosystema 21: 163. 2002.

  • Anamorph: Hirsutella zhangjiajiensis Z.Q. Liang & A.Y. Liu

CLAVICIPITACEAE Clade C

Clavicipitaceae clade C is a strongly supported group that includes the type species, C. militaris, of Cordyceps (MP-BP = 100 %, ML-BP = 100 %, PP = 1.00 in Figs ​Figs1, 1, ​,2).2). Because of the non-monophyly of Cordyceps, we reintroduce the preexisting family name Cordycipitaceae for Clavicipitaceae clade C. This family name was not validly published and it is validated herein based on the type genus Cordyceps. Most of the species in the family parasitize hosts in leaf litter, moss, or upper soil layers and produce superficial to partially immersed to completely immersed perithecia on a fleshy stroma or subiculum that is pallid or brightly coloured. The family contains species of Cordyceps and Torrubiella (Figs ​(Figs5, 5, ​,7).7). The unispecific genus Phytocordyceps is also recognized as a member of this family and transferred to Cordyceps (Fig. 6). In addition, the recent molecular study shows that species of the genera Ascopolyporus A. Möller and Hyperdermium J. White, R. Sullivan, G. Bills & N. Hywel-Jones 2000 [non Link], both pathogens of scale insects, are also inferred to be members of the family (Sullivan et al. 2000, Bischoff et al. 2005).

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Fig. 7.

A–S. Representative species of Cordyceps and its allies in Clavicipitaceae clade B. T–X. Ascus and ascospore of Cordyceps species in this clade. A. C. ophioglossoides on truffle (Elaphomyces sp.: Eurotiomycetes). B. C. japonica on truffle (Elaphomyces muricatus: Eurotiomycetes), OSC 110991. C. C. subsessilis on scarabaeid beetle in decaying wood (Coleoptera), OSC 128581. D. C. gracilis on lepidopteran larva, EFCC 10121. E. C. heteropoda on nymph of cicada (Hemiptera), EFCC 1012. F. C. nigrella on coleopteran larva, EFCC 3438. G. C. sobolifera on nymph of cicada (Hemiptera), EFCC 7768. H. C. longissima on nymph of cicada (Hemiptera), EFCC 8576. I. C. unilateralis on ant (Hymenoptera). J. C. cochlidiicola on lepidopteran larva, EFCC 377. K. C. agriotidis on coleopteran larva, EFCC 5274. L. C. sinensis on larva of Hepialus sp. (Lepidoptera), EFCC 3248. M. C. brunneipunctata on coleopteran larva. N. C. sphecocephala on wasp (Hymenoptera). _O. C. nutans on stink bug (Hemiptera). _P. C. tricentri on adult of Tricentrus sp. (Hemiptera), EFCC 1001; bar = 10 mm. Q. Hymenostilbe odonatae on adult of dragonfly (Odonata), EFCC 12459; bar = 10 mm. R. Hirsutella sp. on wasp (Hymenoptera). S. Paecilomyces lilacinus. T. C. robertsii, ascus with disarticulating ascospores, MICH 2874. U. C. acicularis, ascus and nondisarticulating ascospores, OSC 110987. V. C. paludosa, non-disarticulating ascospores, MICH 14366. W. C. variabilis, disarticulated part-spores in ascus, and X. Part-spores, OSC 128581. Scale bars: A–B = 10 mm, C = 1 mm, D–H = 10 mm, I = 5 mm, J–S = 10 mm, T–X = 10 μm.

CORDYCIPITACEAE Kreisel 1969 ex G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, fam. nov. MycoBank MB504360.

Cordycipitaceae Kreisel, Grundz. Natürl. Syst. Pilze: 112. 1969 [nom. inval., Art. 36].

Stromata vel subiculum pallida vel laete colorata, carnosa. Perithecia superficialia vel omnino immersa, perpendicularia ad superficiem. Asci cylindrici, apice inspissato. Ascosporae cylindricae, multiseptatae, maturae diffrangentes vel integrae remanentes.

Stromata or subiculum pallid or brightly pigmented, fleshy. Perithecia superficial to completely immersed, oriented at right angles to the surface of the stroma. Asci cylindrical with thickened ascus apex. Ascospores usually cylindrical, multiseptate, disarticulating into part-spores or remaining intact at maturity.

Type: Cordyceps Fr.

Teleomorphic genera: Ascopolyporus, Cordyceps, Hyperdermium, Torrubiella.

Anamorphic genera: Beauveria, Engyodontium, Isaria, Lecanicillium, mariannaea-like, Microhilum, Simplicillium.

CORDYCEPS Fr., Observ. Mycol. 2 (revis.): 316. 1818 emend. G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora

= Phytocordyceps C.H. Su & H.-H. Wang, Mycotaxon 2: 338. 1986.

Stromata or subiculum pallid or brightly pigmented, fleshy. Perithecia superficial to completely immersed, ordinal in arrangement. Asci hyaline, cylindrical with thickened ascus apex. Ascospores hyaline, cylindrical, multiseptate, disarticulating into part-spores or nondisarticulating, rarely possessing a thread-like structure connecting the fusiform ends.

Type: Cordyceps militaris (L.: Fr.) Fr., Observ. Mycol. 2(revis.): 317. 1818.

Anamorphic genera: Beauveria, Isaria, Lecanicillium, mariannaea-like, Microhilum, Simplicillium.

Commentary: Species of Cordyceps s. s. are characterized by possessing fleshy stromata that are pallid or brightly coloured. Because species of Torrubiella are interspersed among Cordyceps species in the basal part of the Cordycipitaceae, its ultimate application to a monophyletic taxon within the Cordycipitaceae is not clear, however (Fig. 10). The genus Torrubiella was erected in 1885 by Boudier with the type species T. aranicida Boud. (Kobayasi & Shimizu 1982). Our sampling included several species of Torrubiella that were interspersed amongst species of Cordycipitaceae, but we could not get hold of T. aranicida. Thus, Cordyceps s. s. is narrowly applied to the strongly supported clade (MP-BP = 98 %, ML-BP = 98 %, PP = 1.00 in Figs ​Figs1, 1, ​,2, 2, ​,10)10) that includes Cordyceps species closely related to C. militaris. Cordyceps species that are placed outside of the Cordyceps s. s. node, but within the Cordycipitaceae, are provisionally retained within Cordyceps s. l. Torrubiella species that are part of the Cordyceps s. s. are transferred accordingly. The full extent to which the names Cordyceps and Torrubiella will ultimately be applied awaits additional sampling of Torrubiella, especially that of T. aranicida with the possibility that Torrubiella will need to be synonymized with Cordyceps. Although Phytocordyceps is characterized by its possession of bola-ascospores, it is also synonymized with Cordyceps because of its phylogenetic placement (Figs ​(Figs8, 8, ​,1010).