✽
For correspondence. E-mail: yaoyj@im.ac.cn; Tel.: +86-10-64807496;
Fax: +86-10-64807518
§
Supplemental material for this article may be found at
http://www.springer.com/content/120956
The Journal of Microbiology (2011) Vol. 49, No. 6, pp. 913-919
Copyright ⓒ 2011, The Microbiological Society of Korea
DOI 10.1007/s12275-011-1193-z
A Survey of the Geographic Distribution of Ophiocordyceps sinensis
§
Yi Li
1,2
, Xiao-Liang Wang
1,2
, Lei Jiao
1
, Yi Jiang
1
, Hui Li
3
, Si-Ping Jiang
3
, Ngarong Lhosumtseiring
4
,
Shen-Zhan Fu
1
, Cai-Hong Dong
1
, Yu Zhan
1
,
and Yi-Jian Yao
1
*
1
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, P. R. China
2
Graduate University of Chinese Academy of Sciences, Beijing 100049, P. R. China
3
Tibet Plateau Institute of Biology, Lhasa 850001, P. R. China
4
The Bureau of Science and Technology, Yushu Tibetan Autonomous Prefecture, Qinghai 815000, P. R. China
(Received April 14, 2011 / Accepted August 6, 2011)
Ophiocordyceps sinensis is one of the best known fungi in Traditional Chinese Medicine. Many efforts have
been devoted to locating the production areas of this species resulting in various reports; however, its geo-
graphic distribution remains incompletely understood. Distribution of O. sinensis at the county level is clari-
fied in this work based on both a literature search and fieldwork. More than 3600 publications related
to O. sinensis were investigated, including scientific papers, books, and online information. Herbarium speci-
mens of O. sinensis and field collections made by this research group during the years 2000-2010 were
examined to verify the distribution sites. A total of 203 localities for O. sinensis have been found, of which
106 are considered as confirmed distribution sites, 65 as possible distribution sites, 29 as excluded distribution
sites and three as suspicious distribution sites. The results show that O. sinensis is confined to the Tibetan
Plateau and its surrounding regions, including Tibet, Gansu, Qinghai, Sichuan, and Yunnan provinces in
China and in certain areas of the southern flank of the Himalayas, in the countries of Bhutan, India and
Nepal, with 3,000 m as the lowest altitude for the distribution. The fungus is distributed from the southern-
most site in Yulong Naxi Autonomous County in northwestern Yunnan Province to the northernmost site
in the Qilian Mountains in Qilian County, Qinghai Province, and from the east edge of the Tibetan Plateau
in Wudu County, Gansu Province to the westernmost site in Uttarakhand, India. The clarification of the
geographic distribution of O. sinensis will lay the foundation for conservation and sustainable use of the
species.
Keywords: distribution, Ophiocordyceps sinensis, Tibetan Plateau
Ophiocordyceps sinensis (Berk.) G.H. Sung, J.M. Sung, Hywel-
Jones & Spatafora [≡ Cordyceps sinensis (Berk.) Sacc.] has
long been recorded as one of the most valued Traditional
Chinese Medicines in a number of ancient codices (Wang,
1694; Wu, 1757; Zhao, 1765). Recent studies have also dem-
onstrated that the fungus can be used to treat a wide range
of conditions (Wang, 1995; Zhu et al., 1998a, 1998b; Holliday
and Cleaver, 2008). It has been officially classified as a drug
in the Chinese Pharmacopoeia since 1964 (Committee of
Pharmacopoeia, Chinese Ministry of Health, 1964, 2005). The
price of natural products of O. sinensis has increased sharply
in recent years and is now sold at the price of gold or higher.
However, the natural resource of this fungus is limited due
to its strict host-specificity on moth insects, confined geo-
graphical distribution, and over exploitation by humans in re-
cent decades. It has, therefore, been listed as an endangered
species under the second class of state protection since 1999
(State Forestry Administration and Ministry of Agriculture,
1999).
O. sinensis parasitizes underground dwelling larvae of moths
(Lepidoptera), especially species of Thitarodes. The body of
the insect host is used by the fungus as substrate to form
the mycelium, which is, finally converted into a sclerotium,
leaving the exoskeleton intact. When the stroma of the fungus
grows from the sclerotium and emerges above the ground,
it is collected with the sclerotium as a whole for medicinal
use. The larvae of the host insect live underground for their
entire larval stage of three to four years or longer, feeding
on roots and caudexes of alpine plants. If infected by the
fungus, they usually die in the winter. The fungal stroma
comes out in the spring or summer of the following year (Yao,
2004). Hyphae of O. sinensis can grow at about 2°C with the
optimum temperature of 15-18°C, but the hyphal growth is
restrained when the temperature reaches 25°C, and stopped
below 0°C (Liu et al., 1989; Dong and Yao, 2010).
There is a great volume of accounts on O. sinensis in the
literature (primarily in Chinese), whilst there are not many
reports based on fieldwork in the production areas to deter-
mine the distribution range of the species. Shen et al. (1980)
reported the distribution of O. sinensis in Yushu Tibetan
Autonomous Prefecture, Qinghai Province, China, at an alti-
tude from 4,200-5,000 m on both sides of the watershed.
Based on two field trips to Kangding County, Sichuan
Province, Xiao et al. (1983) reported that O. sinensis was dis-
tributed from 3,000 m up to the snow line. The vertical dis-
tribution of the species was then reported in Sichuan Province
from 3,700-4,700 m in Garzê Tibetan Autonomous Prefecture
(Yin et al., 1990), variable in different locations with the low
914 Li et al.
range of 3,450 m in Kangding County and of 4,200 m in Sêrxü
County (Li and Cao, 1990), and growing well above 4,500
m and up to about 5,000 m in Baiyü County (Li and Cao,
1990). After investigating the natural distribution and eco-
logical environment of O. sinensis in the seven prefectures
of Ngari, Qamdo, Lhasa, Nyingch, Nagqu, Xigazê, and
Shannan in Tibet, Jiang (1989) reported that the species was
distributed in all the prefectures from the altitude of 3,000
to 4,700 m. The distribution range of the fungus was shown
as almost the same as the distribution of host insects in Nagqu
Prefecture, Tibet, where the host insects were distributed from
4,100-5,000 m altitude, but sparsely between 4,100-4,400 m
and above 4,900 m (Chen et al., 1999). The vertical distribu-
tion of O. sinensis along the altitude was also reported to
be accordant with that of its host insect in Baima Mountain,
Yunnan, from 3,850-5,000 m (Shen et al., 1988). According
to Yang et al. (1993), O. sinensis was distributed only in the
northwest of Yunnan with an altitude above 4,000 m, but not
in the center and northeast of the province even if at the same
altitude. Balfour-Browne (1955) described some Himalayan
fungi, including O. sinensis from the southeast of Tibet, Bhutan
and Nepal from 4,200-4,650 m altitude. Kobayasi (1980) re-
ported that the species was collected on the south side of the
Himalayas in Bhutan, India and Nepal, with altitudes recorded
as 3,500 m in the Annapurna Snow Mountain of Nepal and
4,000 m in Kharka, India. Recently, O. sinensis was reported
from India on high hills along the border lines with Nepal
and China at an altitude ranging from 3,300-4,600 m (Negi
et al., 2009).
The record of exceptional low altitude for the species was
made as 2,900 m by Liu et al. (1986), and even as low as
2,260 m and 2,700 m mentioned by Cheng et al. (2007) and
Liang et al. (2008), whilst the majority of the reports were
from above 3,000 m. In a recent review, an altitude of 2,600
m for O. sinensis was also added by Yang et al. (2010).
Several attempts have been made to summarize the distribu-
tion of O. sinensis. Zang (1979) gave the first general dis-
tribution of the species in China as latitudinally from Central
Yunnan Plateau to the Qilian Mountains in Qinghai Province
and transmeridionally from Mount Daloushan in Guizhou
Province to the wide areas of the Himalayas. Recently, coor-
dinates of longitude and latitude for distribution of O. sinensis
was reported by Xishuangbanna Tropical Botanical Garden,
Chinese Academy of Sciences (2008), indicating Cangshan
Mountain (Dali, Yunnan, 25°45′N, 99°34′E) as the southern-
most and Qilian Mountains (South of Hexi Corridor in Gansu,
38°49′N, 102°90′E) as the northernmost boundaries. The pro-
vincial distribution of the species in China was summarized
and illustrated by Pegler et al. (1994) as Gansu, Qinghai,
Sichuan, Yunnan and Tibet. Winkler (2008) also illustrated
the distribution range of O. sinensis, including the eastern part
in the north, down to the south-eastern part and the south
edge of the Tibetan Plateau, with a line to separate the dis-
tribution and non-distribution areas starting roughly from the
east of Xining City in Qinghai, passing through the west of
Nagqu Town and down to Gyangzê or Kangmar county in
Tibet, and then around the Himalayas westwards and ended
in Uttarakhand in India. In Qinghai Province, an investigation
was carried out by the General Investigation Office of Qinghai
for Chinese Traditional Medicine Resource in 1990 (Ye et
al.,
1995), with a report of distribution sites and estimation
of yield of O. sinensis in 29 counties of eight prefectures. In
Yushu and Guoluo prefectures of Qinghai Province, the dis-
tribution and yield of O. sinensis have been well investigated.
All the 47 villages and towns of Yushu Prefecture, except
eight, were reported to harvest the fungus by Lei (1995) and
the amount of harvest in 33 villages was also reported recently
by Tseiga et al. (2005). A report on five of the six counties
in Guoluo Prefecture, except Madoi County, was made by
Zhang (2003) with estimations of annual yield. The distribu-
tion of O. sinensis in Gansu Province was first reported from
Wudu and Wenxian counties (Liu et al., 1986), and then fol-
lowed by Luqu and Maqu counties (Dong and Luo, 1996;
Fang and Zhang, 2005). For Sichuan Province, Yin et al. (1990)
reported the distribution in almost every county of Garzê
Tibetan Autonomous Prefecture. In addition to the distribution
of O. sinensis in the seven prefectures of Tibet Autonomous
Region reported by Jiang (1989), its distribution at the county
level were given by Jin (2003) including 43 counties in eastern
and central Tibet. For the distribution of O. sinensis in
Yunnan Province, Dêqên, Zhongdian (= Xamgyi′nyilha) and
Lijiang counties were mentioned by Xu (1991), whereas most
counties in northwestern Yunnan were reported by Yang
(1997a and 1997b). In addition, the distribution of O. sinensis
has been spread widely online since the year 2000. Most of
the materials online are short articles with a list of localities,
often at county level or above, without any analysis (e.g. Jin,
2003). These lists have then been copied and displayed on
various websites (e.g. Anon, 2009a, 2009b; and so on) and
also reproduced in scientific publications (e.g. Hu et al., 2005;
Ya ng et al., 2010). The distribution and the impact of O. si-
nensis on local economies in the Himalayan countries, Bhutan,
Nepal and India, have also drawn much attention in recent
years (Cannon et al., 2009; Kumar et al., 2010; Pant, 2010).
Since the latter part of 1970s, much intensive research work
on O. sinensis has been carried out, but the distribution of the
species was not fully established. The reports or the analyses
of the distribution were either on a local scale, up to the
provincial level, or in a general statement without specified
location. Furthermore, places far from the Tibetan Plateau
were also regarded as distribution areas of O. sinensis, e.g.
Guangdong, Guangxi, Guizhou, Hainan, Hubei, Jilin, Shaanxi,
Shanxi, Taiwan, Xinjiang, Zhejiang in China (Guo and Liu,
1958; Liu, 1984; Zhao and Mao, n.d.; Wang, 1995; Zhi, 1998;
McKenna et al., 2002; Zhuang, 2005) and also Australia,
Canada, Finland, Ghana, Italy, Kenya, Japan, Mexico, New
Zealand, Norway, Russia, Sweden, The Netherlands, The
United States and Tanzania (Wang, 1995; McKenna et al.,
2002; Dahal, 2004), although they are very doubtful. To clarify
this situation, a survey through extensive fieldwork and thor-
ough literature searches was undertaken. The literature was
analyzed with the field records made by this group over the
past 11 years and specimens preserved in HMAS (Mycological
Herbarium, Institute of Microbiology, Chinese Academy of
Sciences) and HKAS (Herbarium of Cryptograms, Kunming
Institute of Botany, Chinese Academy of Sciences). The re-
sults are presented in this paper.
Distribution of O. sinensis 915
Materials and Methods
This survey covers the reports on O. sinensis and also the five species
described from the Tibetan Plateau during 1989-1998, including O.
gansuensis (K.Y. Zhang, C.J. Wang & M.S. Yan) G.H. Sung, J.M.
Sung, Hywel-Jones & Spatafora (≡ Cordyceps gansuensis K.Y. Zhang
et al.), O. crassispora (M. Zang, D.R. Yang & C.D. Li) G.H. Sung
et al. (≡ Cordyceps crassispora M. Zang et al.), O. multiaxialis (M. Zang
& Kinjo) G.H. Sung et al. (≡ Cordyceps multiaxialis M. Zang & Kinjo),
O. nepalensis (M. Zang & Kinjo) G.H. Sung et al. (≡ Cordyceps nepal-
ensis M. Zang & Kinjo) and O. kangdingensis (M. Zang & Kinjo)
G.H. Sung et al. (≡ Cordyceps kangdingensis M. Zang & Kinjo). Among
these names, the former four have been proved to be synonyms of
O. sinensis by several independent investigations using ITS sequence
analyses (Kang et al., 2000; Liu et al., 2001; Kinjo and Zang, 2001);
however, the relationship between O. kangdingensis and O. sinensis
still requires clarification (Jiang and Yao, 2004). It is likely that O.
kangdingensis will be proved to be another synonym of O. sinensis,
because the former was described from an area within the dis-
tribution range of the latter, as seen in the cases of the other synon-
ymous names.
More than 3,600 publications (over 3,100 in Chinese and over 500
in English) related to O. sinensis were investigated, including scien-
tific papers, books and online information. The literature search was
conducted to find all the available information on the distribution
of O. sinensis, including the material on the Internet. However, only
the earliest and the most representative reports were listed in the
Supplementary data Table 1. Field expeditions to Gansu, Qinghai,
Sichuan, Yunnan Provinces and Tibet Autonomous Region were car-
ried out by this research group during the years 2000-2010 and the
locality, altitude, latitude, and longitude were noted as completely
as possible. The fungal collections made during the fieldwork and
specimens of O. sinensis preserved in HMAS and HKAS, including
those under synonyms as listed above, were examined and used to
determine the distribution sites of this species. All the related speci-
mens were examined morphologically for identification and some
were subjected to molecular analyses.
For the general pattern of distribution of O. sinensis
, the location
s
i
tes were based on the county level within Tibet, Gansu, Qinghai,
Sichuan and Yunnan, except for Ngari Prefecture in Tibet and
Kunming Municipality in Yunnan because there is no further infor-
mation on distribution in counties available. Locations reported out-
side of the Tibetan Plateau in China were indicated by the province
concerned and those outside of China by the country. The name
and the boundary of some counties in China have been changed in
recent years, causing some confusion in the analysis. To update the
geographic information, the new edition of regionalism for admin-
istration promulgated on the website of Ministry of Civil Affairs of
the People's Republic of China (2010) is adopted in this survey.
English translations for all the localities within China were obtained
from the Atlas of China (Di, 2008).
The reported localities for O. sinensis were divided into four cate-
gories: confirmed distribution site (C), possible distribution site (P),
suspicious distribution site (S) and excluded site (E). Confirmed dis-
tribution sites were backed by supporting materials in four ways: 1)
specimens collected in the fieldwork conducted by this group; 2)
specimens preserved in HMAS, HKAS or other herbaria (excluding
misidentifications); 3) specimens studied in published research ar-
ticles and cited with clear information related to the determination
and the location of the fungus; 4) ITS sequences from GenBank
(http://www.ncbi.nlm.nih.gov/genbank/index.html) which provide clear
locality information with correct identification confirmed by the ITS
sequence data generated by this research group. Reported localities
on the Tibetan Plateau with an altitude above 3,000 m (some of these
localities were surrounded by confirmed production sites), were clas-
sified as possible distribution sites if no supporting material was
found. Localities on the edge of the whole production region with
limited acreage higher than 3,000 m, but without supporting evidence
of fungal specimen, were considered as suspicious distribution sites.
Localities with an altitude lower than 3,000 m and/or far away from
the Tibetan Plateau were excluded.
Results
In the fieldwork carried out by this research group since the
year 2000, 441 collections were made on the Tibetan Plateau.
Among the collections, some 240 were recorded with clear
altitude information. The vertical distribution of O. sinensis
varied in different areas. The lowest altitude recorded was
3,087 m with supporting collections (CS 465, CS 466) made
in Xiaojin County, Sichuan Province. It is conceivable that
3,000 m is in general the reliable lowest altitude for the dis-
tribution of O. sinensis. The highest altitude was recorded as
5,048 m with supporting specimen (CS 187) collected in
Nagqu County, Tibet.
Some 53 herbarium specimens under Cordycpes sinensis
and its synonyms mentioned above, preserved in HMAS (33
specimens) and HKAS (20 specimens), were examined. Among
the specimens, six were found to be misidentified (two in
HMAS and four in HKAS) including two sites which have
never been reported in the literature for the distribution of
O. sinensis, i.e. Jiangxi Province (HKAS 18226 and 18227) and
Daguan County in Yunnan Province (HKAS 13178). Forty
seven specimens with correct identification were used in the
analysis of distribution sites of the species.
A total of 203 localities (county or above) for the dis-
tribution of O. sinensis (marked in Figs. 1A, B) were gathered
during this survey, including 106 confirmed distribution sites,
65 possible distribution sites, three suspicious distribution
sites and 29 excluded sites, which are listed in Supplementary
data Table 1 with references, specimen support and notes on
distribution. Among the 106 confirmed distribution sites, 88
were supported by voucher collections in addition to other
evidence, three by sequence data in GenBank and publications,
and 15 by research articles with credible supporting materials.
For the 88 sites supported by voucher collections, 80 sites
were recorded in 441 field collections made by this group
and 20 sites in 47 specimens preserved in both HMAS and
HKAS. There are 12 sites supported by both the field collec-
tions and the herbarium specimens. A total of 12 sites in Tibet
(Chengguan District, and Damxung, Markam, Zogang, Cona,
Lhozhag, Qusum, Dinggyê, Gamba, Lhazê, Nyalam, and
Sa′gya counties) are first reported here based on the fieldwork
carried out by this group (Supplementary data Table 1). O.
sinensis is confined to the Tibetan Plateau and its surrounding
regions, including Tibet, Gansu, Qinghai, Sichuan, and Yunnan
provinces in China and in certain areas of the southern flank
of the Himalayas, in the countries of Bhutan, India and
Nepal. In this survey, it was found that the fungus was dis-
tributed from alpine region of northwestern Yunnan, with
916 Li et al.
Fig. 1. Geographic distribution of O. sinensis. (A) World distribution. (B) Distribution in China. Numbers 1-12 indicate excluded distribution
sites at the provincial level of Xinjiang, Jilin, Shanxi, Shaanxi, Hubei, Zhejiang, Jiangxi, Guizhou, Taiwan, Guangdong, Guangxi and Hainan
respectively. Numbers 13-14 indicate excluded distribution sites at the city level of Jiangyou City and Kunming Municipality. Numbers 15-17
indicate suspicious distribution sites of Dulan and Madoi counties in Qinghai Province and Mount Emei in Sichuan Province.
Yulong Naxi Autonomous County as the southernmost boun-
dary (27°04′14′′, 110°11′43′′E, confirmed by field collections
CS 17-CS 24 by this research group), to the Qilian Mountains,
with Qilian County in Qinghai as the northernmost locality
(38°17′46.92′′N, 99°17′26.22′′E, confirmed by collection CS 262);
and from the east edge of the Tibetan Plateau, with Wudu
County in Gansu as the easternmost boundary (33°23′N,
104°55′E, supported by HMAS 163296, 163298-163301, and
163303; longitude and latitude cited here were generated by
Google Earth Version 5.2.1.1329 beta (http://earth.google.com)
using the county locality records of the specimens), to District
Pithoragarh of Uttarakhand, India as the westernmost locality
(29°34′N, 80°13′E, see Singh and Bhatt, 2010; longitude and
latitude coordinates from Google Earth).
Discussion
In this work, all the reported distribution areas of O. sinensis
were investigated through the literature search in combina-
tion with examination of herbarium specimens and field col-
lections gathered in expeditions to the Tibetan Plateau. This
is a full-scale survey covering all the production regions of
the fungus, and the location sites were reviewed at the county
level within the five production provinces in China, including
Gansu, Qinghai, Sichuan, Tibet, and Yunnan. The distribution
of O. sinensis is clarified here as confined to the Tibetan
Plateau and its surrounding regions at an altitude above 3,000
m. The reports of the fungus from other regions were either
based on misidentification of the specimen concerned or
lacked the support of any fungal collection.
It is evident that the upper altitude limit of the species
distribution may reach the snowline, whilst the low altitude
limit is variable, usually higher in the south and lower in the
north. The lowest altitude for the distribution of O. sinensis,
3,000 m, was determined based on both the previous reports
and the collections made by this research group from the entire
production regions of this fungus for 11 years. To verify the
altitude record of 2,900 m for the species distribution by Liu
et al. (1986), an expedition to Gansu was conducted by this
group in 2006. Attempts were made to visit collecting sites
with the kind help of the senior author of that report, Mr.
Liu, but none of the sites were below the altitude of 3,000 m.
Furthermore, another research group of Gansu Provincial
Grassland Supervision Station (Zhao et al., 2010) reported
recently the vertical distribution of the fungus in the province
as 3,000-4,250 m. For the extreme low altitude records of
2,260 m and 2,700 m, the information on localities of the
specimens cited by Cheng et al. (2007) and Liang et al. (2008),
both from a research group of To n g j i U n i v e r s i t y, w a s c he ck ed
against geographic information (Google Earth Version
5.2.1.1329 beta, http://earth.google.com) based on the provided
latitude and longitude for accuracy. The Tongji University
group investigated the phylogenetic relationship of host in-
sects of O. sinensis using Cytb gene sequences (Cheng et al.,
2007), and the genetic diversity and structure of the fungus
using ISSR technique (Liang et al., 2008). The identical mate-
rials were used in the two articles as shown by the information
of collecting sites and altitudes (Cheng et al., 2007; Liang et
al., 2008) but some of the corresponding longitudes and lat-
itudes for each specimen were inconsistent. For the material
collected in Qilian County, Qinghai Province, marked as at
the altitude of 2,700 m with the longitude and latitude
(38°02′N, 100°22′E) given in Cheng et al. (2007) pointed to
a site above 3,400 m altitude in Google Earth. When using
the longitude and latitude information for the same locality
provided by Liang et al. (2008, 38°01′N, 100°13′E), the altitude
of the site was above 4,000 m. In fact, some 25 specimens
of O. sinensis were collected in the same county, Qilian, in
2008 by this group, with the lowest altitude for the distribu-
tion recorded as 3,221 m. For the material from Huangzhong
County, also in Qinghai Province, cited by Cheng et al. (2007)
and Liang et al. (2008) as at an altitude of 2,260 m, the longi-
tude and latitude (36°49′N, 101°57′E) in Cheng et al. (2007)
pointed outside the boundary of the county, whilst the longi-
Distribution of O. sinensis 917
tude and latitude (36°29′N, 101°34′E) for the same collection
in Liang et al. (2008) did fall into the county but at a down-
town area with an altitude of 2,700 m. According to our ex-
pedition to the county, O. sinensis does not occur at the down-
town site. In a recent attempt to review the distribution pat-
tern and responses to environmental changes of O. sinensis,
Ya ng et al. (2010) mentioned that the fungus was found at
an elevation of 2,600 m in Gulang, Minle and Tianzhu coun-
ties of Gansu Province, located at the north-eastern edge of
the Tibetan Plateau, but no supporting evidence was provided.
There are areas above altitude of 3,000 m in those three coun-
ties and the fungus was collected in Minle County over 3,700
m altitude by this group in 2008 (CS 254-CS 261). The dis-
tribution of O. sinensis in Minle County is therefore con-
firmed, while Gulang and Tianzhu counties are considered
as possible distribution sites because there is no supporting
material available. The record of O. sinensis at the altitude
of 2,600 m in those three counties is in need of validation
by field collections of the species.
The fieldwork carried out on the Tibetan Plateau by this
group not only validated many locations of O. sinensis reported
previously, but also extended them to 12 previously un-re-
ported counties (Supplementary data Table 1). In addition,
of the confirmed 106 distribution sites, nearly three quarters
(80 counties) were supported by the field collections gathered
by this group. Of the 88 distribution sites supported by speci-
mens, almost 90% were verified by our field collections. The
specimens collected by this group over the past decade thus
serve as important supporting materials to verify the actual
distribution of the fungus.
The excluded distribution sites of O. sinensis include 11
provinces within China (Guangdong, Guangxi, Guizhou,
Hainan, Hubei, Jilin, Shaanxi, Shanxi, Taiwan, Xinjiang, and
Zhejiang) and 16 foreign countries (Australia, Canada, Finland,
Ghana, Korea, Italy, Japan, Kenya, Mexico, New Zealand,
Norway, Russia, Sweden, Tanzania, The Netherlands, and The
United States) which are far from the Tibetan Plateau (Figs.
1A, B). Among the provinces in China reported for distribution
of the species, citations of specimen support were found for
Hubei (HMAS 45960 and 46069) and Xinjiang (HKAS 7724).
Examination of these specimens revealed erroneous identi-
fication, which in some cases can be found directly from the
relevant report, e.g. the fungus illustrated in Zhao and Mao
(n.d.) from Xinjiang is apparently not O. sinensis. The distri-
bution of O. sinensis in Guizhou Province was mentioned in
a number of reports (e.g. Zang, 1979; McKenna et al., 2002;
Holliday and Cleaver, 2004), but both the specimen citation
and the detailed locality are lacking. In fact, the highest alti-
tude in Guizhou Province is 2,900.6 m at Mount Jiucaiping,
which is apparently not suitable for the growth of O. sinensis.
Furthermore, Guizhou Province was not included in the dis-
tribution areas of the species in a recent volume of ‘Flora
Fungorum Sinicorum: Cordyceps’ by Liang (2007), a group
of mycologists working on Cordyceps s. l. in that province.
There are two excluded sites in the five production provinces
in China, including two cities: Jiangyou City in Sichuan and
Kunming City in Yunnan. The former was excluded because
the highest altitude of the city is 2,356 m. The distribution
of the species in the latter was based on a specimen (HMAS
01131, Ying and Zang, 1994) purchased from the market, not
directly collected from the locality, and therefore the site is
excluded for the time being until further evidence is available.
The three suspicious distribution sites of O. sinensis are
Mount Emei in Sichuan, Dulan and Madoi counties in
Qinghai (Fig. 1B). The species on Mount Emei was reported
twice with field collections at an altitude above 3,000 m (Chen
et al., 1973; Zhang, 1987), but the specimens are no longer
available for study. It was later considered that Mount Emei
might be a distribution site for O. sinensis once, but the species
is not there now owing to climate changes (Li et al., 19
91).
As Mount Emei is located on the very edge of the Tibetan
Plateau with a very limited area higher than 3,000 m, it might
be that the habitat for O. sinensis has been lost. However,
it appears more probable that the collection records from
Mount Emei were another species, O. emeiensis (A.Y. Liu
& Z.Q. Liang) G.H. Sung et al. (≡ Cordyceps emeiensis A.Y.
Liu & Z.Q. Liang), which also parasitizes larvae of Thitarodes
but has smaller ascospores than that of O. sinensis. The type
specimen of that species was collected at 3,040 m in Mount
Emei in August 1991 (Liu et al., 1997). Dulan County lies
on the Plateau edge to the eastern side of the Qaidam Basin.
It was reported to have the species in Xiangride Town of the
county with a total estimated annual yield of 77 kg (Ye et
al., 1995), but an expedition to the site by this group in June
2008 revealed no evidence for the presence of the species.
It is unclear how the previous report on O. sinensis was made
because the local government officers denied any suggestion
of the species occurring in the county during our visit in that
year. Madoi County was listed recently by Yang et al. (2010)
as one of the distribution sites of O. sinensis, but it has always
been considered as the only county in Guoluo Prefecture
without any record of the species (e.g. Diao, 1996; Zhang,
2003). During our fieldwork in the prefecture, carried out
in the years of 2005, 2008 and 2010, O. sinensis was never
found in Madoi County and local government officers kept
rejecting the occurrence of the fungus in the county. Although
at a high attitude, the county is regarded as not suitable for
the growth of the fungus owing to the dry climate and soil
conditions.
Most of the possible distribution sites for O. sinensis de-
termined here are likely to have a true distribution because
they are on the Plateau and adjacent to a confirmed site.
However, although these sites may have been reported more
than once, even including an estimate of yield, they are not
supported by sufficient evidence, e.g. field collection record
and/or specimen. For this reason, a broader distribution range
(especially for the southernmost boundary) provided by Yang
et al. (2010) cannot be confirmed until identifiable collections
become available.
False distribution sites of O. sinensis, which are caused not
only by species misidentification but most often by the lack
of supporting specimens collected directly from the field, have
been found in both Chinese (e.g. Wang, 1995) and English
(McKenna et al., 2002) publications. It is often the case that
an unreliable report was cited by subsequent authors without
any checking.
It is worth notice that the 203 localities on the Tibetan
Plateau and surrounding regions investigated in the present
study have probably not covered the entire distribution area
for O. sinensis. Many more counties on the Plateau, where
918 Li et al.
the species may be well established, have not been included
in Table S1 because, like the 12 counties first reported for
the distribution of O. sinensis in this survey, they had not
been reported before. On the other hand, among the 106 con-
firmed distribution counties, there are 88 supported by speci-
mens, and for the possible distribution sites, many only require
confirmation through field collections. It is important to clarify
the distribution of O. sinensis for conservation and for sustain-
able use of this fungal resource. More fieldwork on the
Tibetan Plateau is highly desirable.
Acknowledgements
This work is supported by the National Science and
Technology Supporting Projects operated by the Ministry of
Science and Technology of China (2007BAI32B03), the Key
Research Project of Innovation Programs (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).
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