Reassessment of type specimens of Cordyceps and its allies, described by Dr. Yosio Kobayasi and preserved in the mycological herbarium of the National Museum of Nature and Science. Part 4. Cordyceps s. l. on Lepidoptera

Abstract

Dr. Kobayasi and Mr. Shimizu described 31 species of Cordyceps infecting Lepidoptera. Holotype specimens of 14 species and two authentic specimens of one of the 31 species were rediscovered from a herbarium of the National Museum of Nature and Science (TNS). Registration numbers (TNS-F-number) were given to these 16 specimens, and one was lectotypified as follows. Holotypes: Metarhizium indigoticum TNS-F-230337; Yosiokobayasia kusanagiensis TNS-F-197994 (Clavicipitaceae); Beauveria hepialidicola (Kobayasi & Shimizu) Hirok. Sato, S. Ban & Hosoya, comb. nov. TNS-F-197986; Cordyceps ampullacea TNS-F-197981, Cordyceps militaris f. alba TNS-F-230340, Cordyceps ochraceostromata TNS-F-195471, and Cordyceps rosea TNS-F-197972 (Cordyceps sensu stricto, Cordycipitaceae); Ophiocordyceps aurantia TNS-F-195485, Ophiocordyceps cochlidiicola TNS-F-195470, and Ophiocordyceps hiugensis TNS-F-197978 (Ophiocordy-cipitaceae); and Cordyceps changpaishanensis TNS-F-195501, Cordyceps ootakiensis TNS-F-197976, Cordyceps shimizui TNS-F-197995, and Cordyceps sulfurea TNS-F-197974 (Cordyceps sensu lato). Lectotype: Cordyceps bulolensis TNS-F-230327 (Cordyceps sensu lato). A new combination Beauveria hepialidicola comb. nov., is proposed for Cordyceps hepialidicola based on morphological observations.

Cordyceps is the most known genus among insect pathogenic fungi. More than 160 species of this genus and allied species have been described by Dr. Yosio Kobayasi and Mr. Daisuke Shimizu (Kobayasi 1982; Index Fungorum, http://www.indexfungorum.org/Names/Names.asp, as of October 2020). Unfortunately, no registration numbers were provided when they were described, making it difficult for researchers to specify and re-examine the specimens. Nevertheless, most specimens housed in the herbarium of the National Museum of Nature and Science (TNS) now have registration numbers. Furthermore, we have been assessing the rediscovery and specifying the registration number of the type specimens of Cordyceps in TNS (Sato, Ban, Masuya, & Hosoya, 2010a, 2010b, 2012).

Fortunately, additional specimens have been rediscovered in the herbarium of the University Museum of the University of Tokyo (Sato, Awano, & Yago, 2014). From both collections in TNS and the University of Tokyo, 47 species of holotypes, lectotypes, and authentic Kobayasi's specimens were listed (Sato, 2017), including Cordyceps, Isaria, Synnematium, and Torrubiella.

Kobayasi and Shimizu described 31 taxa (including five formae in C. tuberculata (Lebert) Maire) from Lepidoptera (Kobayasi, 1941; 1981a, 1981b; 1982; 1983; Kobayasi & Shimizu, 1976; 1978; 1980; 1982a, 1982b; 1983b). The scientific names of five formae under C. tuberculata were given by referring to species that have been described originally out of Japan. Type specimens of C. bokyoensis Kobayasi have been kept in the herbarium of the National Taiwan University (Kobayasi, 1983), and C. cotopaxiana Kobayasi has been kept in the herbarium of the Department of Botany, Swedish Museum of Natural History, Stockholm (Kobayasi, 1981b). These seven taxa were excluded from this study. Cordyceps nikkoensis Kobayasi was not included in the 31 taxa due to the revision of the host insect from Lepidoptera in the original description (Kobayasi, 1941) to Coleoptera (Kobayasi & Shimizu, 1983b).

In the present assessment, we recognized 15 species of 16 authentic specimens of Cordyceps infecting lepidopteran insects. Moreover, as necessary, we registered the rediscovered specimens in TNS, following the same procedure of Sato et al. (2010a).

We checked the data between each label on the specimens and original papers (e.g., place and date) and compared the specimens' morphology to the figures in the papers. The specimen was judged as a holotype when data were congruent on one specimen. Interpretations were added when data were not congruent or when two specimens occurred with the same data. The latest scientific names revised by a molecular analysis were applied (Sung et al., 2007; Kepler, Humber, Bischoff, & Rehner, 2014; Kepler et al., 2017; Mongkolsamrit et al., 2020).

The summary of the results on the type determination is shown in Table 1. Nomenclatural treatments are discussed in the following text.

Table 1. List of the relocated type specimens originally described as Cordyceps spp. on Lepidoptera by Dr. Kobayasi and Mr. Shimizu in TNS

Fig.	Current name Japanese name	Consistency of				Determined status	Registration number (TNS-F-)	Preserved status	Comments	References
		Place	Collector	Date	Original collection number
1	Metarhizium indigoticum Midori-tosakatake	○	○	○	-	Holotype	230337	Formalin	Difficult to judge if the illustrations in Kobayasi & Shimizu (1983a, PL40-1) & Shimizu (1994, PL 99) originated from the type specimen.	Kobayasi & Shimizu (1978), Kepler et al. (2012), (2014)
2	Yosiokobayasia kusanagiensis Kusanagi-himetanpotake	○	○	○	○	Holotype	197994	Formalin	One of the two specimens is illustrated in Kobayasi & Shimizu (1983a, PL30-2), Shimizu (1994, PL 98).	Kobayasi & Shimizu (1983b), Kepler et al. (2014), Mongkolsamrit et al. (2020)
3	Beauveria hepialidicola Kusagi-mushitake	-	×	○	○	Holotype	197986	Formalin	Collected not by Shimizu but Suzuki. Infected by the fungal parasite Syspastospora sp. on a stroma. Color illustration in Kobayasi & Shimizu (1983a PL 26), Shimizu (1994, PL89) b.	Kobayasi & Shimizu (1983b), Sung et al. (2007)
4	Cordyceps ampullacea Iragano-benisanagitake	○	-	○	○	Holotype	197981	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 1-1), Shimizu (1994, PL 95).	Kobayasi & Shimizu (1982b), Sung et al. (2007)
5	C. militaris f. alba Shiro-sanagitake	○	-	○	○	Holotype	230340	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 19-1), Shimizu (1994, PL 77).	Kobayasi & Shimizu (1978), Yao et al. (1995)
6	C. ochraceostromata Imomushi- hanayasuritake	○	-	○	○	Holotype	195471	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 38-1), Shimizu (1994, PL 91-1).	Kobayasi & Shimizu (1980), Sung et al. (2007)
7	C. rosea Usuaka- shakutorimushitake	○	○	○	○	Holotype	197972	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL3-1), Shimizu (1994 PL 90).	Kobayasi & Shimizu (1982b), Sung et al. (2007)
8	Ophiocordyceps aurantia Sanchuusuge-haritake	○	○	○	○	Holotype	195485	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 7-1), Shimizu (1994, PL 101).	Kobayasi & Shimizu (1980), Sung et al. (2007)
9	O. cochlidiicola Iraga-haritake, Iraga-tsubutake	○	-	○	○	Holotype	195470	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 16-1), Shimizu (1994, PL 102).	Kobayasi & Shimizu (1980), Sung et al. (2007)
10	O. hiugensis Kotsubu-sanagiharitake	○	○	○	○	Holotype	197978	Formalin	Without host insect.	Kobayasi & Shimizu (1983b), Sung et al. (2007)
11	C. bulolensis a No Japanese name	○	○	○	○	Lectotype	230327	Formalin	Congruent morphology with Fig. 17A in Kobayasi & Shimizu (1976).	Kobayasi & Shimizu (1976)
12	C. changpaishanensis a Chohaku-mushitake	○	-	○	-	Holotype	195501	Formalin	Host insects were broken into three pieces.	Kobayasi (1981a)
13	C. ootakiensis a Beni-imomushitake	○	×	○	-	Holotype	197976	Formalin	Collected by not Shimizu but Yamanaka. Color illustration in Kobayasi & Shimizu (1983a, PL 25-2), Shimizu (1994, PL 84).	Kobayasi & Shimizu (1983b)
14	C. shimizuia Oomino-sanagitake	○	-	○	○	Holotype	197995	Formalin	Color illustration in Kobayasi & Shimizu (1983a, PL 8), Shimizu (1994, PL 78).	Kobayasi & Shimizu (1983b), Yao et al. (1995)
15	C. sulfureaa Kogane-imomushitake	○	○	○	○	Holotype	197974	Dried	Color illustration in Kobayasi & Shimizu (1983a, PL 21-1), Shimizu (1994, PL 82).	Kobayasi & Shimizu (1983b)

○: Information between the labels and the original papers matched. ×: Information did not match. -: No information on the label, or not cited in the paper, although the number was on the label.

a: Treated as Cordyceps s.l., after Sung et al. (2007). b: Completely congruent illustrations were used in both books (Kobayasi & Shimizu 1983a, Shimizu 1994).

Clavicipitaceae

1. Metarhizium indigoticum (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber, in Kepler et al., Mycologia 106: 823. 2014. MB 806083

≡Metacordyceps indigotica (Kobayasi & Shimizu) Kepler, G.-H. Sung & Spatafora, in Kepler et al. Mycologia 104: 185. 2012. MB 519857

≡Cordyceps indigotica Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 4: 53. 1978. MB 311796

Fig. 1. (Holotype TNS-F-230337)

Three host larvae were preserved in one bottle and fruiting bodies had been decomposed into several parts, making it impossible to reconstruct the original fruiting bodies. The formalin solution turned black due to chemical degradation. Whether the color illustration in Kobayasi & Shimizu (1983a) and Shimizu (1994) originated from the type specimen or not was difficult to evaluate.

2. Yosiokobayasia kusanagiensis (Kobayasi & Shimizu) Samson, Luangsa-ard & Thanakitp. in Mongkolsamrit et al., Studies in Mycology 95: 243. 2020. MB 834912

≡Metarhizium kusanagiense (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber, in Kepler et al., Mycologia 106: 823. 2014. MB 806085

≡Metacordyceps kusanagiensis (Kobayasi & Shimizu) Kepler, G.H. Sung & Spatafora, in Kepler et al. Mycologia 104: 185. 2012. MB 519859

≡Cordyceps kusanagiensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, B 9: 7. 1983. MB 114246

Fig. 2. (Holotype TNS-F-197994)

Figs. 1-2 - Specimens determined as holotypes of the family Clavicipitaceae. 1. Metarhizium indigoticum TNS-F-230337. 2. Yosiokobayasia kusanagiensis TNS-F-197994. Bars: 1-2 10 mm.

Cordycipitaceae

3. Beauveria hepialidicola (Kobayasi & Shimizu) Hirok. Sato, S. Ban & Hosoya, comb. nov. MycoBank no.: MB 843664

Fig. 3. (Holotype TNS-F-197986)

Figs. 3A-D - Beauveria hepialidicola. TNS-F-197986. 3A. Holotype. 3B. Black perithecia of Syspastospora sp. parasitizing on the stroma of B. hepialidicola. 3C. Sympodial conidiogenous cell of B. hepialidicola. 3D. Conidia of B. hepialidicola stained by cotton blue. Bars: 3A 10 mm; 3B 1 mm; 3C, 3D 5 μm.

≡Cordyceps hepialidicola Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 9: 11. 1983. MB 124189

Note: The conidial formation of Beauveria was observed on hyphae on a stroma of the type specimen. Sympodulae are hyaline, 2.7 μm wide at the base, and 7.1 μm long, and the neck size is 1.0 × 4.1 μm (Fig. 3C). Conidia are globose to subglobose, hyaline, 1.5-2.2 × 2.0-2.5 (mean: 1.8 × 2.2, n = 16) μm (Fig. 3D), produced by sympodial branching from sympodulae.

DNA from the formalin-impregnated specimen could not be extracted. In the latest taxonomy, almost all Cordyceps s.l. species having Beauveria-state have been transferred to the genus Beauveria s.s. based on phylogenetic systematics (Kepler et al., 2017). Beauveria is defined by the morphological structure of conidio-genous cells that elongate in a sympodial pattern. Here, we propose a new combination since we determined both sexual and asexual structures of the specimen belonging to the genus Beauveria s.s.

4. Cordyceps ampullacea Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 8: 112. 1982. MB 114242

Fig. 4. (Holotype TNS-F-197981)

5. Cordyceps militaris f. alba Kobayasi & Shimizu ex Y.J. Yao [as ‘albina’], in Yao, Li, Pegler & Spooner, Acta Mycol. Sin. 14: 257. 1995. MB 414517

Fig. 5. (Holotype TNS-F-230340)

6. Cordyceps ochraceostromata Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 6: 132. 1980. MB 114359

Fig. 6. (Holotype TNS-F-195471)

7. Cordyceps rosea Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 8: 112. 1982. MB 114258

Fig. 7. (Holotype TNS-F-197972)

Figs. 4-7 - Specimens determined as holotypes of the family Cordycipitaceae. 4. Cordyceps ampullacea. TNS-F-197981. 5. Cordyceps militaris f. alba. TNS-F-230340. 6. Cordyceps ochraceostromata. TNS-F-195471. 7. Cordyceps rosea. TNS-F-197972. Bars: 4-7 10 mm.

Ophiocordycipitaceae

8. Ophiocordyceps aurantia (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, in Sung et al., Stud. Mycol. 57: 40. 2007. MB 504228

≡Cordyceps aurantia Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 6: 125. 1980. MB 114351

Fig. 8. (Holotype TNS-F-195485)

9. Ophiocordyceps cochlidiicola (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, in Sung et al., Stud. Mycol. 57: 41. 2007. MB 504240

≡Cordyceps cochlidiicola Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 6: 128. 1980. MB 114354

Fig. 9. (Holotype TNS-F-195470)

10. Ophiocordyceps hiugensis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, in Sung et al., Stud. Mycol. 57: 43. 2007. MB 504279

≡Cordyceps hiugensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 9: 3. 1983. MB 114245

Fig. 10. (Holotype TNS-F-197978)

Figs. 8-10 - Specimens determined as holotypes of the family Ophiocordycipitaceae. 8. Ophiocordyceps aurantia. TNS-F-195485. 9. Ophiocordyceps cochlidiicola. TNS-F-195470. 10. Ophiocordyceps hiugensis. TNS-F-197978. Bars: 8-10 10 mm.

Only one stroma without a host body has been preserved. Due to the lack of a host, it was impossible to determine if the color illustrations in Kobayasi & Shimizu (1983a) and Shimizu (1994) were drawn based on a type specimen.

Cordyceps s. l.

Sung et al. (2007) proposed a new definition of the genus Cordyceps s.s. and divided the genus Cordyceps s.l. by proposing new genera. Simultaneously, in their study, some Lepidoptera-infecting species were treated as incertae sedis due to the lack of information. Those species were treated as Cordyceps s.l. in this study. One of those species was recently transferred to Metarhizium (M. indigo-ticum) by Kepler et al. (2014).

11. Cordyceps bulolensis Kobayasi, in Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 2: 142. 1976. MB 311791

Fig. 11. (Lectotype TNS-F-230327)

Holotype has not been designated in the original description. However, two specimen vials with the same collection data were found, and both were without a host. TNS-F-230327 was selected as the lectotype because perithecia remained. TNS-F-230332 was also a part of stroma from the same collection but without perithecia.

12. Cordyceps changpaishanensis Kobayasi, Bull. Natn. Sci. Mus., Tokyo, ser. B 7: 12. 1981. MB 114352

Fig. 12. (Holotype TNS-F-195501)

13. Cordyceps ootakiensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 9: 15. 1983. MB 114253

Fig. 13. (Holotype TNS-F-197976)

14. Cordyceps shimizui Y.J. Yao, in Yao, Li, Pegler & Spooner, Acta Mycol. Sin. 14: 258. 1995. MB 414812

≡Cordyceps chichibuensis Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 9: 8. 1983. MB 114244

Fig. 14. (Holotype TNS-F-197995)

15. Cordyceps sulfurea Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Tokyo, ser. B 9: 15. 1983. MB 114265

Fig. 15. (Holotype TNS-F-197974)

Figs. 11-15 - Specimens determined as holotypes (H) or a lectotype (L) of the genus Cordyceps s. l. 11. Cordyceps bulolensis. TNS-F-230327 (L). 12. Cordyceps changpaishanensis. TNS-F-195501 (H). 13. Cordyceps ootakiensis. TNS-F-197976 (H). 14. Cordyceps shimizui. TNS-F-197995 (H). 15. Cordyceps sulfurea. TNS-F-197974 (H). Bars: 11-15 10 mm.

Although the external morphology of stromata has been changed by the complete vaporization of the alcohol preservative, both the host's external morphology and the appearing position of stromata were identical to the illustration in Kobayasi & Shimizu (1983a) and Shimizu (1994).

Specimens of other species on Lepidoptera

Specimens of the following nine remaining taxa were not found in this survey: C. alba Kobayasi & Shimizu, C. alpicola Kobayasi, C. hokkaidoensis Kobayasi, C. obliqua Kobayasi, C. pallidiolivacea Kobayasi & Shimizu, C. ramosostipitata Kobayasi & Shimizu, C. rubrostromata Kobayasi, C. shimaensis Kobayasi, and O. fasciculati-stromata (Kobayasi & Shimizu) G.H. Sung et al.

Disclosures

The authors declare no conflicts of interest. All experiments in this study complied with the current laws of Japan.

Acknowledgments

We thank Dr. Hayato Masuya for his advice on organizing the information of Dr. Kobayasi's references and species.

References

• Kepler, R. M., Sung, G.-H., Ban, S., Nakagiri, A, Chen, M.-J., Huang, B., Li, Z., & Spatafora, J. W. (2012). New teleomorph combinations in the entomopathogenic genus Metacordyceps. Mycologia, 104, 182-197. https://doi.org/10.3852/11-070
• Kepler, R. M., Humber, R. A., Bischoff, J. F., & Rehner, S. A. (2014). Clarification of generic and species boundaries for Metarhizium and related fungi through multigene phylogenetics. Mycologia, 106, 811-829. https://doi.org/10.3852/13-319
• Kepler, R. M., Luangsa-Ard, J. J., Hywel-Jones, N. L., Quandt, C. A., Sung, G. H., Rehner, S. A., Aime, M. C., Henkel, T. W., Sanjuan, T., Zare, R., Chen, M., Li, Z., Rossman, A. Y., Spatafora, J. W., & Shrestha, B. (2017). A phylogenetically-based nomenclature for Cordycipitaceae (Hypocreales). IMA fungus, 8, 335-353. https://doi.org/10.5598/imafungus.2017.08.02.08
• Kobayasi, Y. (1941). The genus Cordyceps and its allies. Science Reports of the Tokyo Bunrika Daigaku, Section B, 84, 53-260.
• Kobayasi, Y. (1981a). Revision of the genus Cordyceps and its allies 1. Bulletin of the National Science Museum, Tokyo, Series B, 7, 1-13.
• Kobayasi, Y. (1981b). Revision of the genus Cordyceps and its allies 2. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 7, 123-129.
• Kobayasi, Y. (1982). Keys to the taxa of the genera Cordyceps and Torrubiella. Transactions of the Mycological Society of Japan, 23, 329-364.
• Kobayasi, Y. (1983). Miscellaneous notes of fungi (3). The Journal of the Japanese Botany, 58, 221-224.
• Kobayasi, Y., & Shimizu, D. (1976). The genus Cordyceps and its allies from New Guinea. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 2, 133-151.
• Kobayasi, Y., & Shimizu, D. (1978). Cordyceps species from Japan. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 4, 43-63.
• Kobayasi, Y., & Shimizu, D. (1980). Cordyceps species from Japan 3. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 6, 125-145.
• Kobayasi, Y., & Shimizu, D. (1982a). Cordyceps species from Japan 4. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 8, 79-91.
• Kobayasi, Y., & Shimizu, D. (1982b). Cordyceps species from Japan 5. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 8, 111-123.
• Kobayasi, Y., & Shimizu, D. (1983a). Iconography of vegetable wasps and plant worms [in Japanese]. Hoikusha.
• Kobayasi, Y., & Shimizu, D. (1983b). Cordyceps species from Japan 6. Bulletin of the National Science Museum, Tokyo, Series B (Botany), 9, 1-21.
• Mongkolsamrit, S., Khonsanit, A., Thanakitpipattana, D., Tasanathai, K., Noisripoom, W., Lamlertthon, S., Himaman, W., Houbraken, J., Samson, R. A., & Luangsa-ard, J. (2020). Revisiting Metarhizium and the description of new species from Thailand. Studies in Mycology, 95, 171-251. https://doi.org/10.1016/j.simyco.2020.04.001.
• Sato, H. (2017). Situation of type specimens of Cordyceps and allies described by Dr. Kobayasi. Mycology, 8, 303-307. https://doi.org/10.1080/21501203.2017.1394392
• Sato, H., Awano, K., & Yago, M. (2014). Catalogue of the Masayo Kato Fungi Collection, the University Museum, The University of Tokyo. The University Museum, The University of Tokyo Material Reports, 102, 5-69.
• Sato, H., Ban, S., Masuya, H., & Hosoya, T. (2010a). Reassessment of type specimens of Cordyceps and its allies described by Dr. Yosio Kobayasi preserved in the mycological herbarium of the National Museum of Nature and Science (TNS). Part 1: the genus Torrubiella. Mycoscience, 51, 154-161. https://doi.org/10.1007/s10267-009-0015-1
• Sato, H., Ban, S., Masuya, H., & Hosoya, T. (2010b). Reassessment of type specimens of Cordyceps and its allies described by Dr. Yosio Kobayasi, preserved in the National Museum of Nature and Science. Part 2: Cordyceps (Elaphocordyceps) on Elaphomyces. Mycoscience, 51, 387-390. https://doi.org/10.1007/s10267-010-0047-6
• Sato, H., Ban, S., Masuya, H., & Hosoya, T. (2012). Reassessment of type specimens of Cordyceps and its allies, described by Dr. Yosio Kobayasi and preserved in the mycological herbarium of the National Museum of Nature and Science (TNS). Part 3: Cordyceps s. l. on Cicadidae. Mycoscience, 53, 402-408. https://doi.org/10.1007/s10267-011-0170-z
• Shimizu, D. (1994). Color iconography of vegetable wasps and plant worms (pp.1-380) [in Japanese]. Seibundo Shinkosha.
• Sung, G. H., Hywel-Jones, N. L., Sung, J. M., Luangsa-Ard, J. J., Shrestha, B., & Spatafora, J. W. (2007). Phylogenetic classification of Cordyceps and the clavicipitaceous fungi. Studies in Mycology, 57, 5-59. https://doi.org/10.3114/sim.2007.57.01
• Yao, Y. J., Li, Y., Pegler, D. N., & Spooner, B. N. (1995). Nomenclature of some Cordyceps (Clavicipitaceae). Acta Mycologica Sinica, 14, 256-260.