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1 Zootaxa 2412: (2010) Copyright 2010 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Thitarodes namnai sp. nov. and T. caligophilus sp. nov. (Lepidoptera: Hepialidae), hosts of the economically important entomopathogenic fungus Ophiocordyceps sinensis in Bhutan NORBERT MACZEY 1, KUENZANG DHENDUP 2, PAUL CANNON 1,4, NIGEL HYWEL-JONES 3, & TEK BAHADUR RAI 2 1 CABI Europe-UK, Bakeham Lane, Egham, Surrey TW20 9TY, United Kingdom. n.maczey@cabi.org 2 Renewable Natural Resources Research Centre, Yusipang, Bhutan 3 Mycology Laboratory, National Center for Genetic engineering and Biotechnology, Science Park, Pathum Thani Thailand 4 Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW93AB, United Kingdom Abstract Two new species of Thitarodes Viette (Lepidoptera: Hepialidae) are described from specimens collected in alpine grassland habitats of north-west Bhutan. The new species are compared with their closest relatives from Nepal and neighbouring parts of China and Japan. Male and female adults and genitalia are illustrated. Key words: Cordyceps, Ghost moth, grassland, Himalaya, new species Introduction Cordyceps ( yartsa guenbub in Bhutan or chongchao in China) is a valuable commodity, collected intensively throughout alpine grasslands of the Eastern Himalayas and parts of the Tibetan Plateau. This component of traditional Chinese medicine consists of dried fruiting bodies of the entomopathogenic fungus Ophiocordyceps sinensis (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora and cuticular remains of its insect host, which still preserve the distinct shape of the caterpillar (Cannon et al. 2009). As the only known hosts for Ophiocordyceps sinensis, members of the ghost moth genus Thitarodes Viette play a significant role in the ecological system at the heart of harvest and trade of yartsa guenbub. Caterpillars of this genus are subterranean root-feeders living polyphagously on alpine forbs and sedges (Yang et al. 1996). Larvae take 2 6 years to reach maturity depending on species and environmental conditions (Chen et al. 2002, Yin et al. 2004). It is assumed that caterpillars of this genus are infected by spores of the Ophiocordyceps fungus during early life stages when they stay dormant for a considerable time. Death occurs in the final larval instar prior to pupation with the host being consumed by growing mycelium, turning it into a solid mummy. An elongate fruiting body emerges after snow melt in spring from behind the head capsule of the former caterpillar and the combined feature is intensively collected at this time of the year. Every year, from Bhutan alone, millions of these chinese caterpillars are exported as part of a worldwide multimilliondollar trade in this commodity. Despite being of such high ecological and economic importance, for large parts of the collecting area, the taxonomic identity of the host moths remains unclear (Cannon et al. 2009). The type species of Thitarodes was first described as Hepialus armoricanus Oberthür in Viette (1968) proposed the genus Thitarodes whilst describing three new species from Nepal. The genus seems to be restricted to eastern parts of Asia with a geographical range stretching from Nepal in the west, southwards into Myanmar, and towards the east into Taiwan, Japan, and northeastern parts of Russia. The most recent inventory of the Hepialoidea (Nielsen et al. 2000) lists 51 species belonging to Thitarodes, the majority of which have only relatively recently been described from China, particularly 42 Accepted by J. Rota: 10 Feb. 2010; published: 29 Mar. 2010

2 Yunnan and Tibet. A more recent publication by Zhu et al. (2004) reviewing the Chinese Hepialidae lists several additional species, while not referring to several other species originally described from China. In a number of cases the existing descriptions and relative inaccessibility of type material do not allow these taxa to be properly compared with each other and to possible new species within this genus. Consequently, there is a high possibility that a substantial part of the existing descriptions refer to synonyms as has been already pointed out by Nielsen et al. (2000). Matters are further complicated by the fact that in Chinese publications the genus Thitarodes is not adopted and all relevant taxa are still referred to as Hepialus. In their revision of Chinese hepialids, Chu and Wang (1985) regarded the differences in male genitalia as described by Viette as too insignificant for a generic character. A revision of Thitarodes is urgently needed, particularly considering the great economic importance of some of the species belonging to this genus. Specimens of the new species described here were collected in the alpine region of northwest Bhutan at a grassland site well known locally for its abundance of Ophiocordyceps sinensis. Material and methods Collecting of Thitarodes specimens was carried out in Bhutan during June 2005, June/July 2006, July 2007 and June The work was part of a project on sustainable use of Cordyceps funded through the Darwin Initiative of DEFRA (Department for Environment, Food and Rural Affairs), UK. Due to the inaccessibility of the field site, equipment and supplies had to be carried by horses and mules on a five-day trek from the nearest road-head. The field site (Fig. 36), known as one of the best Cordyceps sites in the north-west of Bhutan, is a north-west facing slope ranging in altitude from approximately 4,400 m to 4,900 m and is locally known as Namna. The short treeless vegetation at the site is characterized by a small scale mosaic of alpine Kobresia grassland, dominated by Kobresia schoenoides (C.A. Meyer) Steudel (Cyperaceae), with dwarf rhododendrons (Rhododendron setosum D. Don & R. nivale Hook. f.; Ericaceae). The chosen method of collecting was light-trapping using a gas lamp in all four sampling periods (Fig. 37). In 2008 a battery powered torch using light emitting diodes (LED) was used as an additional light source. Due to the inaccessibility of type material deposited in China, only specimens from the Natural History Museum London, UK (T. danieli Viette [paratypes], T. dierli Viette [paratypes], T. eberti Viette [paratypes], T. kingdonwardi Ueda [holotype], T. maculatum Ueda [holotype], T. nebulosus Alpheraky) and the Museum Alexander König Bonn, Germany (T. amoricanus) could be directly examined for this study. Currently legislation in Bhutan does not allow the export of any biological specimens. All specimens are deposited at the National Biodiversity Centre (NBC) in Serbithang, Bhutan; holotype collection-nr ; paratypes collection-nr Description Thitarodes namnai Maczey, sp. nov. Figs Etymology. The name refers to the location, where both holotype and paratypes were sampled, locally known as Namna. Male. Description based on holotype (Figs. 1 3, 24). SIZE: Wingspan: 37.5 mm; forewing length: 18.2 mm; hindwing length: 15.0 mm; body length: 14.8 mm. ANTENNAE: Rust brown with segments 4 to 9 dorsally darkened to blackish brown. BODY: Head and thorax blackish brown covered densely with rust coloured hairs turning ochraceous posteriorly; prothorax and mesothorax interspersed with dark brown hairs; abdomen brown covered in ochraceous yellow hairs. THITARODES OF BHUTAN Zootaxa Magnolia Press 43

3 FOREWINGS: Mainly rust coloured with extensive grey-white and black markings; narrow sub-terminal fascia grey-white extending from R 3 to Cu 2, interrupted between M 2 and Cu 2, basal accompanied by slightly wider, wavelike, black fascia extending basal towards centres of cells in places interrupted by rust coloured scales; reniform stigma, posterior half black with small grey-white discal spots, anterior half grey-white with black borders basal and distal; sub-basal patch of grey-white and black spots; large black spot filling cell between vein A and dorsum reaching distal 1/4 of wing; second large black spot filling cell between Cu 2 and A in distal half; costa accompanied by series of small grey-white and black spots; fringe from R 1 to Cu 2 ochraceous yellow and blackish brown; dorsum edged grey-white. Costa straight; Sc unbranched; R 2 and R 3 stalked, length of stalk about 3/4 length from base of R 2+3 to margin; R 4 and R 5 stalked, length of stalk about 1/ 6 of length from base R 4+5 to margin; cross vein R 5 -M 1 very weak but discernible, reaching R 5 distad from furcation of R 4 and R 5 ; two cross veins between A and CuP; one crossvein between Cup and CuA; A reaching margin. FIGURES 1 5. Thitarodes namnai: 1. holotype, male, label 10005; 2. & 3. holotype genitalia in caudal (2) and laterocaudal (3) view; 4. paratype, female, label 10010; 5. female paratype genitalia. HINDWINGS: Uniformly grey, slightly translucent; fringe between R 1 and Cu 2 ochraceous yellow and blackish brown; wing venation similar to forewing but CuP reaching margin; A weak, not reaching margin. LEGS: All legs densely setose with ochraceous and dark brown hairs, lacking spurs; hind tibia slightly broadened and dorsally densely covered with ochraceous 5 to 6 mm long scent-brushes. GENITAL STRUCTURES: In caudal view tegumen triangular elongated towards the inner process. Dorsal process of tegumen obtusely lobed and only weakly sclerotised. Caudal margin of tegumen curved outwards in basal half, but almost straight dorsoventral in inner half (in caudal view). Margin of tegumen curved caudally along inner half displaying a row of coarse, sclerotised lobes (in lateral view). The caudal margin sclerotised dark brown to almost black; the inner half, particularly including the inner process, fully 44 Zootaxa Magnolia Press MACZEY ET AL.

4 sclerotised. Base of vinculum with an oval shaped dilation. Saccus with a median indentation. Valve densely hairy, base sclerotised dark brown with single acute spine diverging from valve at 1/4 of the length of valve. Juxta slightly trapezoidal widening at base, posterior edge rounded with a median indentation. Aedeagus not sclerotized. FIGURES Thitarodes namnai: paratypes males; 6. label 10001; 7. label 10006; 8. label 10007; 9. label 10008; 10. label 10009; paratypes females; 11. label 10002; 12. label 10003; 13. label Female. Description based on paratype (Figs. 4 5). Similar to male with the exceptions described below. SIZE: Wingspan: 41.5 mm; forewing length: 19.0 mm; hindwing length: 17.0 mm; body length: 16.0 mm. ANTENNAE: Uniformly red-brown. BODY: Hairs on thorax yellow caudally. FOREWINGS: Silver-grey coloured with extensive grey-white, rust coloured and black markings; most black spots fringed by a thin rust-brown line; sub-terminal fascia grey-white extending uninterrupted from R 2 to A, basally accompanied by a series of black oval to reniform spots; mosaic of black, rust-brown and silvergrey patches at middle of wing reaching base of wing; costa accompanied by series of small grey-white and rust coloured spots; fringe from R 1 to Cu 2 beige and brown; dorsum edged grey. Wing venation as in male, but crossvein between R 5 and M not reaching M 1. HINDWINGS: Fringe between R 1 and Cu 2 beige and brown; wing venation as in male. LEGS: All legs densely setose with ochraceous hairs, lacking spurs. GENITAL STRUCTURES: Lateral arms of prevulvar sclerite semicircular; preanal sclerite of oval shape with acute, sclerotised apex and blunt dilatation in middle of interior edge, densely setose; intergenital lobe bluntly bilobed dorsally and more acutely bilobed ventrally, densely setose. Holotype. Male, labelled: Bhutan, Namna, N E ; Altitude 4,750 m a.s.l., 16.vii.2008, 19:10, N. Maczey leg., Paratypes (same data as holotype except collection date and label nr.). female, 16.vii.2008, 10010; male, 03.vii.2007, 19:10, 10001; female, 06.vii.2007, 19:30, 10002; female 01.vii.2007, 19:20, 10003; female, 02.vii.2007, 19:35, 10004; male, 16.vii.2008, 19:15, 10006; male, 13.vii.2008, 19:10, 10007; male, 15.vii.2008, 19:15, 10008; male, 16.vii.2008, 19:15, Diagnosis. Thitarodes namnai is a typical medium-sized and slightly dimorphic representative of its genus. Males are on average smaller than females and their forewings tend to be predominantly rust brown THITARODES OF BHUTAN Zootaxa Magnolia Press 45

5 whereas females appear more often predominantly silvery-grey (Figs. 6 13). Male genitalia show the diagnostic features as designated in the original description of the genus (Figs ). Within the Thitarodes species described so far this species can readily be identified in the field by its characteristic two large spots in the forewing bordering vein A in a straight line. This gives both males and females a typical four-spotted appearance when adults are in a resting position. Equally, shape and position of these four spots differ significantly when compared with museum specimens of closely related species prepared according to standards. FIGURES Thitarodes namnai, male genitalia, paratypes: caudal view; latero-caudal view; 14. & 19. label 10001; 15. & 20. label 10006; 16. & 21. label 10007; 17. & 22. label 10008; 18. & 23. label Discussion. RELATED SPECIES FROM NEPAL. Size and colour scheme are similar to Thitarodes danieli and T. eberti, both described from lower altitudes than T. namnai. However, the two contrasting colours of wing fringes are more evenly alternating within T. namnai whereas in T. danieli and T. eberti the dark colour is significantly reduced in both fore- and hindwings. The sub-terminal grey fascia is not accompanied by a continuous dark line or line of spots at the basal side in T. eberti and T. danieli; both of these species show here only a few irregular dark spots. A large black spot in the cell between veins Cu 2 and A has a more or less straight to concave margin along vein A, but is widely curved convexly along Cu 2 in T. namnai. In T. eberti and T. danieli the margin of the same black marking is broadly curved and convex along vein A but straight along Cu 2. There are distinctive differences in the male genital structure between all three species described from Nepal. The main differences from T. eberti, the species with the greatest similarity to T. namnai, are a narrower tegumen, a curved spine on the valve, the rounded median dilatation of vinculum and a missing median indentation of the saccus of the vinculum in T. eberti. Main differences to T. danieli and T. dierli are the shape of the tegumen, which is deeply bilobed in T. danieli and much wider with a heavily sclerotized and finely serrated edge in T. dierli, and a different position of the spine on the valve in both species. RELATED SPECIES FROM CHINA. Thitarodes namnai seems to be closely related to a group of species recently described from Tibet and Sichuan including T. gonggaensis Fu & Huang, T. jialangensis Yang, Hepialus namliensis Zhu & Wang, H. namensis Zhu & Wang, and H. hainanensis Zhu & Wang (Fu et al. 1991, Yang 1994, Zhu et al. 2004). However, all differ distinctly in the shape of the male genitalia as well 46 Zootaxa Magnolia Press MACZEY ET AL.

6 as the shape and pattern of forewings and wing venation from T. namnai. Wing venation in T. jialangensis and H. namensis are similar to T. namnai but the shape of the sacculus and the position of the spine on the valve differ. The species probably closest to T. namnai is T. gonggaensis. This species has been described from Sichuan and is reputedly at the heart of recently reported successful artificial rearing methods for Ophiocordyceps sinensis in China. Differences from T. namnai can be summarized as follows. Wing shape and pattern: The apex of forewings is much more rounded in T. namnai in comparison with T. gonggaensis. The large black spot filling the cell between vein A and the dorsum, present in all specimens recorded from Namna, is missing in T. gonggaensis. The black spot filling the cell between Cu 2 and A in the exterior half is more rudimentary in T. gonggaensis and of a more reniform shape with a rounded posterior edge. A greywhite colouring of the stigma is absent in T. gonggaensis. Male genitalia: The single spine on the valve is inserted at a distance of 1/5 of the total length of the valve from the base in T. namnai, but much closer to the base in T. gonggaensis. This spine of the valve is considerably straighter and longer in T. namnai. The u- shaped median indentation in the saccus of the vinculum of T. namnai is missing in T. gonggaensis. The valve is not hairy in T. gonggaensis but is intensely so in T. namnai. Wing venation: The terminal branch of CuP in T. gonggaensis is not reaching A as in T. namnai. FIGURE 24. Thitarodes namnai: underside holotype, showing wing venation, label It has to be pointed out that morphological features, including the male genitalia, can vary significantly between specimens of a single Thitarodes species. Based on incomplete knowledge of Thitarodes, it is possible that T. namnai may actually represent a subscpecies of or geographic variation within T. gonggaensis. RELATED SPECIES FROM JAPAN. So far only one species (T. nipponensis Ueda) of Thitarodes has been described from Japan (Ueda 1996). This species has wing venation similar to that of T. namnai, but it differs significantly in the shape of the male genitalia, particularly the valves. Thitarodes caligophilus Maczey, sp. nov. Figs Etymology. Latin for the fog and gloom loving one ; the name refers to weather conditions prevalent during the flight period of this species. THITARODES OF BHUTAN Zootaxa Magnolia Press 47

7 FIGURES Thitarodes caligophilus: 25. holotype, male, label 10027; 26. paratype, female, label 10028; 27. paratype, male, label 10025; 28. paratype, male, label Male. Description based on holotype (Figs. 25, 29, 32). SIZE: Wingspan: 44.0 mm; forewing length: 20.0 mm; hindwing length: 17.0 mm; body length: 14.5 mm. ANTENNAE: Uniformly rust brown. BODY: Head and thorax red-brown covered densely with ochraceous hairs; prothorax interspersed with some dark brown hairs; abdomen brown, covered in ochraceous yellow hairs. FOREWINGS: Mainly different shades of grey derived from a mixture of silvery, golden and dark brown scales, mottled with dark brown markings; between C and R uniformly golden grey; narrow sub-terminal fascia grey-white extending from R 3 to Cu 2, interrupted between M 2 and Cu 2 by dark spots, basally accompanied by wavelike black fascia of varying width, extending basad towards centres of cells; oval dark brown stigma with small silvery-white discal spot; sub-basal patch of grey-white and black spots; large black spot bordered by a lining of golden-grey scales filling cell between vein A and dorsum reaching distal ¼ of wing length; second large black spot filling the centre of cell between Cu 2 and A; borders of spot lobed; fringe dark brown in regular intervals interspersed with yellow scales. Costa straight; Sc unbranched; R 2 and R 3 stalked, length of stalk about ¾ length from base of R 2+3 to margin; R 4 and R 5 stalked, length of stalk about 1/6 of length from base R 4+5 to margin; cross vein R 5 -M 1 well discernible, reaching R 5 distad from furcation of R 4 and R 5 ; two cross veins between A and CuP; one crossvein between Cup and CuA; A reaching margin. HINDWINGS: Uniformly dark grey, covered with light brown hairs, slightly translucent; fringe uniformly brown; wing venation similar to forewing but CuP reaching margin; vein A weak, not reaching margin. LEGS: All legs densely setose with ochraceous hairs, lacking spurs; hind tibia slightly broadened and dorsally densely covered with ochraceous 6 mm long scent-brushes. GENITAL STRUCTURES: In caudal view tegumen triangular, elongated towards the inner process. Dorsal process of tegumen obtusely lobed and only weakly sclerotised. Caudal margin of tegumen curved outwards in basal half, but almost straight dorso-ventral in inner half (in caudal view). Margin of tegumen curved caudal along inner half, margin smooth. The caudal margin is sclerotised dark brown to almost black; the inner half particularly including the inner process is fully sclerotised. Base of vinculum with evenly curved-shaped dilation. Saccus evenly curved without any indentation. Valve densely hairy, base sclerotised dark brown with acute, spine-like apex; single acute spine diverging from valve at 1/3 of its length. Juxta almost quadratical, posterior end with rounded edges. Aedeagus not sclerotized. 48 Zootaxa Magnolia Press MACZEY ET AL.

8 FIGURES Thitarodes caligophilus, genitalia: caudal, latero-caudal view; 29. & 32. holotype, male, label 10027; 30. & 33. paratype, male, label 10025; 31. & 34. paratype, male, label ; 35. paratype, female, label Female. Description based on paratype (Figs. 26, 35). Similar to male with the exceptions described below. SIZE: Wingspan: 51.5 mm; forewing length: 24.0 mm; hindwing length: 20.0 mm; body length: 15.0 mm. THITARODES OF BHUTAN Zootaxa Magnolia Press 49

9 FIGURE 36. Namna, the habitat of Thitarodes namnai. ANTENNAE: Uniformly rust brown, dorsally slightly darker. BODY: Head and prothorax interspersed with some dark brown hairs; abdomen brown covered in brown hairs. FOREWINGS: Sub-terminal fascia very similar to the one in male, but basally accompanied by wavelike, interrupted black fascia, extending basally towards centres of cells; oval dark brown stigma, bordered by a line of golden scales; large black spot filling centre of cell between Cu 2 and A; in centre with silvery-white discal spot, borders of spot lined with golden-grey scales; wing venation as in male. HINDWINGS: As in male. LEGS: All legs densely setose with ochraceous hairs, lacking spurs. GENITAL STRUCTURES: Preanal sclerite triangular, densely setose; lateral arms of prevulvar sclerite rectangular and elongated, densely setose; intergenital lobe rectangular with slightly dilated and rounded corners dorsally, densely setose; intergenital sclerite as slightly curved rectangle. Holotype. Male, labelled: Bhutan, Namna, N E ; Altitude 4,750 m a.s.l., 20.vii.2007, 19:31, Kuenzang Dhendup leg., Paratypes (same data as holotype except collection date and label nr.). Female, 19.vii.2007, 09:10, 10028, male, 15.vii.2007, 19:25, 10025; male, 20.vii.2007, 19:25, All specimens are deposited at the National Biodiversity Centre (NBC) in Serbithang, Bhutan; holotype collection-nr ; paratypes collection-nr , & Diagnosis. Thitarodes caligophilus is a medium-sized and only slightly dimorphic representative of its genus. With a wingspan ranging from 44.0 to 47.0 mm males are smaller than the only measured female (wingspan 51.5 mm). In comparison to other species of the genus it is characterised by a rather bluntly 50 Zootaxa Magnolia Press MACZEY ET AL.

10 rounded apex of the forewings, which display a characteristic colour pattern of different shades of grey. The species can easily be identified by the shape of the valve of the male genitalia with its combination of a spinelike apex plus a single acute spine diverging from the valve at one third of its length (Figs ). Discussion. RELATED SPECIES FROM NEPAL. Size and colour scheme differ from Thitarodes species described from Nepal so far. Specimens of T. caligophilus are generally larger although females from T. danieli can be of similar size as T. caligophilus. Colour pattern in this species is characterised by a more uniform pattern of different shades of grey not showing as prominent large spots or fasciae as in the species from Nepal. There are distinctive differences in the male genitalia between all three species described from Nepal. The main difference from T. eberti, the species with the greatest similarities to T. caligophilus, is in the shape of the valve with its combination of a spine-like apex plus a single acute spine diverging from the valve at 1/3 of its length. The latter spine is missing in all species described from Nepal. RELATED SPECIES FROM CHINA. On the basis of wing venation Thitarodes caligophilus seems to be closely related to T. jialangensis and T. namensis. However, the position of the spine on the valve differs significantly from the species as it does with all other species described from China. RELATED SPECIES FROM JAPAN. Thitarodes nipponensis is surprisingly similar to T. caligophilus with regards to its wing venation; however, it differs significantly in the shape of the male genitalia, particularly the shape of the valves. FIGURE 37. Light-trapping in the habitat of Thitarodes namnai. Biology and ecology The habitat of both species described above consists of treeless alpine grassland dominated by Kobresia THITARODES OF BHUTAN Zootaxa Magnolia Press 51

11 sedges (K. schoenoides, K. pygmea (C.B. Clarke) C.B. Clarke, K. humilis (C.A. Meyer) Sergievskaja) and dwarf Rhododendron (R. nivale, R. setosum) (Fig. 36). At the site of collection this habitat stretches from the valley bottom at an elevation of 4,300 m to almost 5,000 m where it peters out into steep rocky outcrops often covered with scree and borders the edge of glaciers. Most of the site shows a north-westerly aspect. Winters are characterised by severe frost and snow fall of varying intensity. Soil temperature only reaches thawing point at the end of April/beginning of May, and between the end of May and the beginning of September the site is frequently grazed by domesticated yaks. Rainfall intensifies during the monsoon season with precipitation being highest from June to August. Frost can occur during clear night skies all year round. During the flight periods of both species at dusk temperatures measured varied between 5 and 8 C. Successful light trapping was conducted during three separate flight periods; 1 6 July 2006 (T. namnai only), July 2007 (T. namnai & T. caligophilus) and July 2008 (T. namnai only). With the exception of a single female of T. caligophilus found resting in the vegetation during the early morning hours, all individuals came to light within a very short time window between 7:10 and 7:35 pm. Unsuccessful light trapping during other field site visits earlier in the season (early June 2005, mid June 2007) suggest that the flight period of adult Thitarodes does not start before the last days in June and apparently coincides with the height of the monsoon characterised by frequent rainfall. Although members of the genus Thitarodes are often not easily attracted by light (Zhu et al. 2004), the low number of specimens may suggest that early July still does not represent the height of the flight period at Namna. This might particularly be true for T. caligophilus with specimens of this species never caught before the 15 th of July. Acknowledgments The authors would like to thank Dr. Gaden Robinson (Natural History Museum, London) and Dr. Matthew Cock (CABI Europe-CH, Delemont, Switzerland) for their help and advice with this manuscript. References Cannon, P.F., Hywel-Jones, N.L., Maczey, N., Norbu, L., Tshitila, Samdup, T. & Lhendup, P. (2009) Steps towards sustainable harvest of Ophiocordyceps sinensis in Bhutan. Biodiversity and Conservation, 18, (DOI /s ). Chen, S.J., Yin, D.H., Zhong, G.Y. & Huang, T.F. (2002) Study on the biology of adults parasite of Cordyceps sinensis, Hepialus biruensis. China Journal of Chinese Materia Medica, 27, Chu, H.F. & Wang, L.Y. (1985) Insect-Herb versus Hepialids with descriptions of new genera and new species of Chinese Hepialidae. Sinozoologia 3, Fu, S., Huang, T. & Luo, Q. (1991) A new species of Hepialus (Lepidoptera: Hepialidae). Acta Entomologica Sinica, 34, Nielsen, E.S., Robinson, G.S. & Wagner, D.L. (2000) Ghost-moths of the world: a global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera). Journal of Natural History, 34, Oberthür, C. (1909) Études de Lépidoptérologie comparée, 3. Rennes, 415 pp. Ueda, K. (1996) A new species of Thitarodes Viette (Lepidoptera, Hepialidae) from Japan. Bulletin Kitakyushu Museum Natural History, 15, Viette, P.E.L. (1968) Contribution à l études des Hepialidae (36ème note): Lepidoptera Hepialidae du Népal. Khumbu Himal, 3, Yang, D.R. (1994) Four new species of the genus Hepialus from Yunnan and Xizang, China. Zoological Research, 15, Yang, D.R., Li, C.D., Shu, C. & Yang, Y.X. (1996) Studies on the Chinese species of the genus Hepialus and their geographical distribution. Acta Entomologica Sinica, 39, Yin, D.H., Chen, S.J., Li, L., Za, X., Zhou, X.L., Sun, J.H. & Za, M.C. (2004) Study on the biological character of the host of Cordyceps sinensis, Hepialus biru in Tibet. Special Wild Economic Animal and Plant Research, 2, 1 5. Zhu, H.F., Wang, L.Y. & Han, H.X. (2004) Lepidoptera: Hepialidae, Epiplemidae, Insecta Vol. 38, Fauna Sinica. Science Press, Beijing, China, 29 pp. 52 Zootaxa Magnolia Press MACZEY ET AL.