(C) 2011 Xiao-Liang Wang. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.
Ophiocordyceps sinensis (≡ Cordyceps sinensis) is one of the most valued medicinal fungi in China, used for its invigorating effects in strengthening the body and restoring energy. The fungus parasitizes larvae of moths and converts them into sclerotia from which the fungus fruiting body grows. Since the late 1950s, considerable effort has been devoted to the study of host insects related to the fungus. In the present paper, the research history of insect species associated with Ophiocordyceps sinensis is briefly reviewed and an extensive literature survey is presented. Ninety-one insect names, spanning 13 genera, related to host insects of Ophiocordyceps sinensis are investigated. The relationships between the reported insect species and Ophiocordyceps sinensis are analyzed. Fifty-seven of these are considered as recognizable potential host species of the fungus distributed throughout the Tibetan Plateau, whilst eight are considered as indeterminate hosts and 26 as non-hosts. Among the names of recognizable potential host insects, three are invalid (nomen nudum) and require further study. This work provides basic information for management of the insect resources and for the conservation and sustainable use of Ophiocordyceps sinensis.
Cordyceps, Fungi, Hepialidae, host insects, Ophiocordyceps
Ophiocordyceps sinensis
(Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora is an
ascomycete fungus, which is also known as the Chinese Caterpillar
Fungus or “Dong Chong Xia Cao” (winter worm, summer grass) in Chinese,
or “Hia Tsao Tong Tchong” and “Hea Tsaon Tsong Chung” in early English
translations (
The natural product of Ophiocordyceps sinensis
for medicinal use is actually a combination of the fungus and an insect
larva. The fungus parasitizes underground dwelling larvae of moths and
converts them into sclerotia, from which the fruiting body of the
fungus grows (
Insect host species of Ophiocordyceps sinensis belong to the family Hepialidae (Lepidoptera) (
Recently, a global inventory of the suborder Exoporia, comprising Mnesarchaeoidea and Hepialoidea, was presented by
Based upon an exhaustive literature search, a total of 4793 publications related to Cordyceps/Ophiocordyceps and Hepialus/Thitarodes, in either English or Chinese, were gathered. Those publications relevant to host insects of Ophiocordyceps sinensis,
including reports on taxonomy, checklists, fauna, biological
characteristics, ecology and geographical distribution were examined
for information about these insects. All the insect names associated
with Ophiocordyceps sinensis
were assessed based on the following criteria to determine their
relationship with the fungus. Taxa which met both of the following
requirements were considered as recognizable potential insect host
species of Ophiocordyceps sinensis:(1) The distribution areas of the insect overlapped that of Ophiocordyceps sinensis,
which was determined on the basis of field collections made by this
research group during the years 2000−2010, examination of herbarium
specimens, and another exhaustive literature analysis carried out in
this laboratory (
A total of 91 names in 13 genera of Hepialidae were found in the literature search. They are listed in alphabetical order in Table 1, together with geographic distribution, altitude, main references and the relationship with Ophiocordyceps sinensis as determined by this study. Insect names used in the references, if different from that in
Fifty-seven species are considered here as recognizable potential host insects of Ophiocordyceps sinensis, whilst eight as indeterminate hosts and 26 as non-hosts. The recorded altitude ranges of the recognized potential host insects were found to vary from 2800 to 5100 m. The distribution areas of these species covered 26 provinces in China and more than 12 other countries. Three of the recognizable potential host names are invalid (nomen nudum).
Table 1. Potential insect hosts of Ophiocordyceps sinensis.
Insect name | Geographic distribution | Altitude (m) | Main references | Status of host insect† | Different name in the main references |
---|---|---|---|---|---|
Bipectilus yunnanensis Chu & Wang, 1985 | Yunnan Province: Lijiang County‡ | 3200 |
|
P | |
Bipectilus zhejiangensis Wang, 2001§ | Zhejiang Province: Anji County‡; Fujian Province | — |
|
N | |
Endoclita anhuiensis (Chu & Wang, 1985)| | Anhui Province: Yuexi County‡ | — |
|
N | Phassus anhuiensis Chu & Wang, 1985 |
Endoclita davidi (Poujade, 1886) | Sichuan Province: Baoxing and Danba Counties; Fujian and Guangxi Provinces | 3600 |
|
P | Hepialus davidi Poujade, 1886;Phassus giganodus Chu & Wang, 1985 |
Endoclita excrescens (Butler, 1877)| | Sichuan Province: Yingjing County; Anhui, Hebei, Heilongjiang, He’nan, Jilin, Liaoning, Shandong and Shanxi Provinces; Inner Mongolia Autonomous Region; Japan | — |
|
N | Phassus excrescens (Butler, 1877) ;Phassus camphorae Sasaki, 1908 |
Endoclita fijianodus (Chu & Wang, 1985)| | Fujian Province‡ | — |
|
N | Phassus fujianodus Chu & Wang, 1985 |
Endoclita jingdongensis (Chu & Wang, 1985)| | Yunnan Province: Jingdong County‡, Xishuangbanna Prefecture‡ | — |
|
N | Phassus jingdongensis Chu & Wang, 1985 |
Endoclita nodus (Chu & Wang, 1985)| | Anhui Province: Yuexi County‡; Guangxi, Guizhou, Hainan, Hu’nan, Jiangxi and Zhejiang Provinces | — |
|
N | Phassus nodus Chu & Wang, 1985 |
Endoclita signifer (Walker, 1856)| | Hu’nan Province | — |
|
N | Phassus hunanensis Chu & Wang, 1985 |
Endoclita sinensis (Moore, 1877)| | Fujian, Guangdong, Guangxi, Hainan, Hebei, He’nan, Hubei, Hu’nan, Jiangxi, Shandong, Shanxi, Sichuan, Yunnan and Zhejiang Provinces; Shanghai Municipality; D.P.R. Korea; India; Japan; Sri Lanka | — |
|
N | Phassus sinensis Moore, 1877;Phassus herzi Fixsen, 1887 |
Endoclita xizangensis (Chu & Wang, 1985)| | Tibet Autonomous Region: Nyalam County‡ | — |
|
N | Phassus xizangensis Chu & Wang, 1985 |
Endoclita yunnanensis (Chu & Wang, 1985)| | Yunnan Province: Jinghong Municipality‡; Guangdong and Hainan Provinces | — |
|
N | Phassus yunnanensis Chu & Wang, 1985 |
Gazoryctra ganna (Hübner, [1808]) | Qinghai Province: Zadoi County; Heilongjiang Province; Inner Mongolia Autonomous Region; Northern Europe; Russia | 3900¶ |
|
P | Hepialus ganna (Hübner, [1808]) |
Gazoryctra macilentus (Eversmann, 1851) | Hebei and Heilongjiang Provinces; Inner Mongolia Autonomous Region; Eastern Siberia; Mongolia | 340–1300 |
|
N | Hepialus macilentus Eversmann, 1851 |
Hepialiscus jiangbeiensis Chu & Wang, 2004§ | Chongqing Municipality‡ | — |
|
N | |
Hepialiscus ledongensis Chu & Wang, 2004§ | Hainan Province: Ledong County‡ | — |
|
N | |
Hepialiscus nepalensis (Walker, 1856) | Tibet Autonomous Region: Nyalam County; India; Nepal; Sikkim | — |
|
I | Hepialiscus flavus Chu & Wang, 1985 |
Hepialus bibelteus Shen & Zhou, 1997§ | Yunnan Province: Deqên County‡ | 4500 |
|
P | |
Hepialus biruensis Fu, 2002§ | Tibet Autonomous Region: Biru County‡ | 4400–4700 |
|
P | |
Hepialus dinggyeensis Chu & Wang, 2004§ | Tibet Autonomous Region: Dinggyê County‡ | — |
|
I | |
Hepialus gangcaensis Chu & Wang, 2004§ | Qinghai Province: Gangca County‡ | 3195¶ |
|
P | |
Hepialus guidera Yan, 2001§ | Qinghai Province: Guide County | 3400–3600 |
|
P, IN | Hepialus guidera Yan, 2001 |
Hepialus hainanensis Chu & Wang, 2004§ | Hainan Province: Ledong County‡ | — |
|
N | |
Hepialus humuli (Linnaeus, 1758) | Heilongjiang Province; Europe and Siberia | — |
|
N | |
Hepialus lagii Yan, 2001§ | Qinghai Province: Guide County | 3400–3600 |
|
P, IN | |
Hepialus latitegumenus Shen & Zhou, 1997§ | Yunnan Province: Deqên County‡ | 4500 |
|
P | |
Hepialus maquensis Chu & Wang, 2004§ | Gansu Province: Maqu County‡ | 3300¶ |
|
P | |
Hepialus namensis Chu & Wang, 2004§ | Tibet Autonomous Region: Damxung County‡ | 4200¶ |
|
P | |
Hepialus namlinensis Chu & Wang, 2004§ | Tibet Autonomous Region: Namling County‡ | 3704¶ |
|
P | |
Hepialus pui Zhang, Gu & Liu, 2007§ | Tibet Autonomous Region: Nyingchi County‡ | 4100–5000 |
|
P | |
Hepialus xiaojinensis Tu, Ma & Zhang 2009§ | Sichuan Province: Xiaojin‡ and Jinchuan County‡ | 3500–4800 |
|
P | |
Hepialus xingazeensis Chu & Wang, 2004§ | Tibet Autonomous Region: Xigazê Prefecture‡ | — |
|
I | |
Hepialus yadongensis Chu & Wang, 2004§ | Tibet Autonomous Region: Yadong County‡ | — |
|
I | |
Hepialus yongshengensis Chu & Wang, 2004§ | Yunnan Province: Yongsheng County‡ | — |
|
I | |
Hepialus zadoiensis Chu & Wang, 2004§ | Qinghai Province: Zadoi County‡ | 3900¶ |
|
P | |
Magnificus jiuzhiensis Yan, 2000§ | Qinghai Province: Jigzhi County‡ | 3800–3900 |
|
P | |
Magnificus zhiduoensis Yan, 2000§ | Qinghai Province: Zhidoi County‡ | 4400–4600 |
|
P | |
Napialus chenzhouensis Chu & Wang, 2004§ | Hu’nan Province: Chenzhou City‡; Shanghai Municipality | — |
|
N | |
Napialus chongqingensis Wu, 1992 | Chongqing Municipality‡ | — |
|
N | |
Napialus hunanensis Chu & Wang, 1985 | Hu’nan Provinces: Changsha City‡; Guangdong, Guangxi, Hainan, and Jiangxi Provinces | — |
|
N | |
Napialus jiangxiensis Chu & Wang, 2004§ | Jiangxi Province: Taihe County‡ | — |
|
N | |
Palpifer sexnotatus (Moore, 1879)| | Sichuan and Taiwan Provinces; Kashmir; India; Sri Lanka; Japan | — |
|
N | |
Parahepialiscus borneensis (Pfitzner in Pfitzner & Gaede, 1933) | Hu’nan Province; Malaysia | — |
|
N | Hepialiscus borneensis Pfitzner, 1933 |
Pharmacis carna ([Denis & Schiffermüller], 1775) | Sichuan Province: Luhuo County; Europe | 3050¶ |
|
P | Hepialus carna ([Denis & Schiffermüller], 1775) |
Pharmacis fusconebulosa (De Geer, 1778) | Sichuan Province: Kangding District; Europe; Russia | 3500¶ |
|
P | Hepialus fusconebulosa (De Geer, 1778);Hepialus gallicus Lederer, 1852 |
Pharmacis pyrenaicus (Donzel, 1838) | Sichuan Province: Dêgê County; Southwest Europe | 3880¶ |
|
P | Hepialus alticola Oberthür, 1881 |
Sthenopis regius (Staudinger, 1896)| | — | — |
|
N | Phassus regius (Staudinger, 1896) |
Sthenopis roseus (Oberthür, 1911)| | Hubei Province | — |
|
N | Phassus miniatus Chu & Wang, 1985 |
Thitarodes albipictus (Yang, 1993) | Yunnan Province: Deqên County‡ | 4500–4800 |
|
P | Hepialus albipictus Yang, 1993 |
Thitarodes altaicola (Wang, 1990) | Xinjiang Uygur Autonomous Region‡ | 1300–l800 |
|
N | Hepialus altaicola Wang, 1990 |
Thitarodes anomopterus (Yang, 1994) | Yunnan Province: Jianchuan‡ and Lijiang Counties‡ | 2800–3100 |
|
P | Hepialus anomopterus Yang, 1994 |
Thitarodes armoricanus (Oberthür, 1909) | Gansu, Qinghai, Sichuan and Yunnan Provinces; Tibet Autonomous Region; Xinjiang Uygur Autonomous Region | 3600–5000 |
|
P | Hepialus armoricanus Oberthür, 1909 |
Thitarodes baimaensis (Liang in |
Yunnan Province: Deqên County‡ | 4500–4900 |
|
P | Hepialus baimaensis Liang, 1988 |
Thitarodes baqingensis (Yang & Jiang, 1995) | Tibet Autonomous Region: Baqên County‡ | 4600–4800 |
|
P | Hepialus baqingensis Yang & Jiang, 1995 |
Thitarodes callinivalis (Liang, 1995) | Tibet Autonomous Region; Yunnan Province : Deqên County‡ | 4300–4600 |
|
P | Hepialus callinivalis Liang, 1995 |
Thitarodes cingulatus (Yang & Zhang in |
Gansu Province: Wenxian County‡ | 3200–3800 |
|
P | Hepialus cingulatus Yang & Zhang, 1995 |
Thitarodes damxungensis (Yang in |
Tibet Autonomous Region: Damxung County‡ | 4500–4680 |
|
P | Hepialus damxungensis Yang, 1995 |
Thitarodes deqinensis (Liang in |
Yunnan Province: Deqên County | 4200–4700 |
|
P | Hepialus deqinensis Liang, 1988 |
Thitarodes dongyuensis (Liang in Yang et al. 1992) | Tibet Autonomous Region: Markam County; Yunnan Province: Deqên County | 4000–4700 |
|
P, IN | Hepialus dongyuensis Liang in Yang et al. 1992 |
Thitarodes ferrugineus (Li, Yang & Shen, 1993) | Yunnan Province: Deqên County‡ | 4200–4700 |
|
P | Hepialus ferrugineus Li, Yang & Shen, 1993 |
Thitarodes gonggaensis (Fu & Huang in |
Sichuan Province: Kangding County‡ | 3800–4400 |
|
P | Hepialus gonggaensis Fu & Huang, 1991 |
Thitarodes jialangensis (Yang, 1994) | Tibet Autonomous Region: Zogang County‡ | 4000–4600 |
|
P | Hepialus jialangensis Yang, 1994 |
Thitarodes jianchuanensis (Yang, 1994) | Yunnan Province: Jianchuan County‡ | 2900–3500 |
|
P | Hepialus jianchuanensis Yang, 1994 |
Thitarodes jinshaensis (Yang, 1993) | Yunnan Province: Deqên County‡ | 4600 |
|
P | Hepialus jinshaensis Yang, 1993 |
Thitarodes kangdingensis (Chu & Wang, 1985) | Sichuan Province: Kangding County‡ | 3600–4500 |
|
P | Hepialus kangdingensis Chu & Wang, 1985 |
Thitarodes kangdingroides (Chu & Wang, 1985) | Sichuan Province: Kangding County‡ | 4200 |
|
P | Hepialus kangdingroides Chu & Wang, 1985 |
Thitarodes lijiangensis (Chu & Wang, 1985) | Yunnan Province: Lijiang County‡ | 3500–4400 |
|
P | Hepialus lijiangensis Chu & Wang, 1985 |
Thitarodes litangensis (Liang, 1995) | Sichuan Province: Litang‡ and Batang County; Tibet Autonomous Region | 4300–4700 |
|
P | Hepialus litangensis Liang, 1995 |
Thitarodes luquensis (Yang & Yang in |
Gansu Province: Luqu County‡ | 4276–4300 |
|
P | Hepialus luquensis Yang & Yang, 1995 |
Thitarodes markamensis (Yang, Li & Shen, 1992) | Tibet Autonomous Region: Markam County‡; Yunnan Porvince: Deqên County | 4500–4900 |
|
P | Hepialus markamensis Yang, Li & Shen, 1992 |
Thitarodes meiliensis (Liang in |
Yunnan Province: Deqên County‡ | 3650–4700 |
|
P | Hepialus meiliensis Liang, 1988 |
Thitarodes menyuanicus (Chu & Wang, 1985) | Gansu Province: Jishishan County; Qinghai Province: Hualong, Menyuan‡ and Tongren Counties | — |
|
I | Hepialus menyuanicus Chu & Wang, 1985 |
Thitarodes nebulosus (Alpheraky, 1889) | Qinghai Province: Yushu Prefecture; Tibet Autonomous Region: Amdo‡ and Damxung Counties‡, Nagqu Prefecture | 4500 |
|
P | Hepialus nebulosus Alphéraky, 1889 |
Thitarodes oblifurcus (Chu & Wang, 1985) | Qinghai Province: Yushu Prefecture‡; Sichuan Province: Kangding County | 4000–4500 |
|
P | Hepialus oblifurcus Chu & Wang, 1985 |
Thitarodes pratensis (Yang, Li & Shen, 1992) | Yunnan Province: Deqên County‡ | 4350 |
|
P | Hepialus pratensis Yang, Li & Shen, 1992 |
Thitarodes renzhiensis (Yang in Yang et al. 1991) | Yunnan Province: Deqên County‡ | 3880–5100 |
|
P | Hepialus renzhiensis Yang, 1991 |
Thitarodes sichuanus (Chu & Wang, 1985) | Sichuan Province‡: Aba Prefecture, Emei and Kangding Counties; Chongqing Municipality | 3600–3800 |
|
P | Hepialus sichuanus Chu & Wang, 1985 |
Thitarodes varians (Staudinger, 1896) | Sichuan Province: Batang County; Tibet Autonomous Region: Qamdo County | 4500 |
|
P | Hepialus varians Staudinger, 1896 |
Thitarodes xizangensis (Chu & Wang, 1985) | Tibet Autonomous Region: Nyalam County‡ | 2200 |
|
N | Forkalus xizangensis Chu & Wang, 1985 |
Thitarodes xunhuaensis (Yang & Yang in |
Qinghai Province: Xunhua County‡ | 3800 |
|
P | Hepialus xunhuaensis Yang & Yang, 1995 |
Thitarodes yeriensis (Liang, 1995) | Yunnan Province: Deqên County‡ | 4500–4700 |
|
P | Hepialus yeriensis Liang, 1995 |
Thitarodes yulongensis (Liang, 1988) | Yunnan Province: Lijiang County‡ | 4150–4500 |
|
P | Hepialus yulongensis Liang, 1988 |
Thitarodes yunlongensis (Chu & Wang, 1985) | Yunnan Province: Yunlong‡ and Dali Counties; Hainan Province | 3600–4200 |
|
P | Hepialus yunlongensis Chu & Wang, 1985 |
Thitarodes yunnanensis (Yang, Li & Shen, 1992) | Yunnan Province: Jianchuan‡, Lanping‡, Lijiang‡ and Weixi Counties | 3600–4100 |
|
P | Hepialus yunnanensis Yang, Li & Shen, 1992 |
Thitarodes yushuensis (Chu & Wang, 1985) | Qinghai Province: Yushu Prefecture‡, Batang, Chindu and Zadoi Counties; Gansu Province | 4500–4900 |
|
P | Hepialus yushuensis Chu & Wang, 1985 |
Thitarodes zaliensis (Yang, 1994) | Tibet Autonomous Region: Markam County‡ | 4600–4900 |
|
P | Hepialus zaliensis Yang, 1994 |
Thitarodes zhangmoensis (Chu & Wang, 1985) | Tibet Autonomous Region: Nyalam County‡ | 2200 |
|
N | Hepialus zhangmoensis Chu & Wang, 1985 |
Thitarodes zhayuensis (Chu & Wang, 1985) | Tibet Autonomous Region: Zayü‡ and Markam Counties; Yunnan Province: Deqên and Gongshan County | 4200–4400 |
|
P | Hepialus zhayuensis Chu & Wang, 1985 |
Thitarodes zhongzhiensis (Liang, 1995) | Yunnan Province: Deqên County‡ | 4000–4600 |
|
P | Hepialus zhongzhiensis Liang, 1995 |
Triodia nubifer (Lederer, 1853) | Sichuan Province: Kangding Prefecture; Central Asia | — |
|
I | Hepialus nubifer Lederer, 1853 |
Triodia sylvina (Linnaeus, 1761) | Sichuan Province: Kangding County; Central Asia; Central Europe and Northern Europe | — |
|
I | Hepialiscus sylvinus (Linnaeus, 1761) |
† The status of host insect of Ophiocordyceps sinensis determined in this study: I = indeterminate host, N = non-host, IN = invalid name, P = potential host;
‡ Type-locality;
§ Names not included in
| Stem-borers;
¶ The lowest altitude of the reported locality in China.
DiscussionThrough an extensive literature survey, all the Hepialidae species reported from China were listed and analyzed using detailed information on their geographic distribution, altitude and nomenclature. The relationships between the insect species and Ophiocordyceps sinensis were clarified based on available information. The data provided here serve as a foundation for further investigations on the conservation biology of this endangered fungal species and its insect hosts.
Species in different genera of Chinese hepialids can be
divided into two categories according to the feeding strategy of the
larvae (
Eight species, including 1 Hepialiscus, 4 Hepialus, 1 Thitarodes and 2 Triodia species (Table 1), are considered as indeterminate hosts of Ophiocordyceps sinensis. While the distribution ranges of these species are within that of Ophiocordyceps sinensis, they lack an altitude record and require further confirmation before being considered as potential hosts of Ophiocordyceps sinensis, e.g., Hepialus yadongensis, Triodia sylvina, etc.
Fifty-seven taxa are recognized as potential hosts of Ophiocordyceps sinensis, including 1 Bipectilus, 1 Endoclita, 1 Gazoryctra, 12 Hepialus, 2 Magnificus, 3 Pharmacis and 37 Thitarodes species (Table 1). The distribution ranges of these insects overlap that of Ophiocordyceps sinensis.
Altitude information for these insects was reported in three ways in
the literature: (1) The altitude range of the insect was reported
unambiguously above 3000 m, e.g., Thitarodes baimaensis, Thitarodes meiliensis, etc. (37 species). Among these, the lowest altitude of 3200 m was reported for Thitarodes cingulatus (
Three names of the recognizable potential host insects are invalid (nomen nudum)
because no full description of the species was published in the
literature, although the names appeared several times in various
publications (Table 1). Among them, Thitarodes dongyuensis was described by Yang (1992) as ‘Hepialus dongyuensis’ and deemed as a nomen nudum in
Species of Hepialus and Phassus described from China after 1984 have been transferred to Thitarodes and Endoclita respectively by
Natural production of Ophiocordyceps sinensis has been declining significantly over the last few decades while the market demands on the fungus have increased sharply in recent years. Clarification of the host insects of Ophiocordyceps sinensis will provide basic information for management of the insect resources and for the conservation and sustainable use of the fungus. This work has gathered the available information on the host insects of Ophiocordyceps sinensis and will lay a foundation for further studies of the relationship between the fungus and its hosts, especially their co-evolution (an ongoing research project based on DNA sequence analyses in this laboratory), and also for the cultivation of this valuable fungus for massive production.
This work is supported by the National Science and Technology Supporting Projects operated by the Ministry of Science and Technology of the People’s Republic of China (2007BAI32B03), the Key Research Project of Innovation Programmes (KSCX2-YW-G-076, KSCX2-YW-G-074-04, KSCX2-SW-101C) and the scheme of Introduction of Overseas Outstanding Talents operated by the Chinese Academy of Sciences, and the National Science Funds for Distinguished Young Scholars from the National Natural Science Foundation of China (30025002).