Ophiocordyceps sinensis

{{Short description|Species of fungus}} {{Redirect|Vegetable caterpillar|the other fungus with this common name|Ophiocordyceps robertsii{{!}}''Ophiocordyceps robertsii''}} {{Speciesbox | image = Cordyceps sinensis.jpg | image_caption = ''Ophiocordyceps sinensis'' (left) growing out of the head of a dead caterpillar | status = VU | status_system = IUCN3.1 | status_ref = <ref name=IUCN>{{cite iucn |last1=Yang |first1=Z.-L. |year=2020 |title=''Ophiocordyceps sinensis'' |amends=2020 |volume=2020 |article-number=e.T58514773A179197748 |doi=10.2305/IUCN.UK.2020-3.RLTS.T58514773A179197748.en |access-date=14 February 2025}}</ref> | genus = Ophiocordyceps | species = sinensis | authority = (Berk.) G.H.Sung, J.M.Sung, Hywel-Jones & Spatafora (2007) | synonyms_ref = <ref name="urlMycoBank: Ophiocordyceps sinensis"/> | synonyms = ''Sphaeria sinensis'' Berk. (1843) ''Cordyceps sinensis'' (Berk.) Sacc. (1878) }} {{Infobox Chinese | name = ''Ophiocordyceps sinensis'' | t = 冬蟲夏草 | s = 冬虫夏草 | p = dōng chóng xià cǎo | l = winter worm, summer grass | tib = དབྱར་རྩྭ་དགུན་འབུ་ | wylie = Dbyar Rtswa Dgun'bu | zwpy = Yarza Günbu | nep = यार्सागुम्बा | my = ရှီးပတီး | vie= = Đông trùng hạ thảo | t2=蟲草 | s2=虫草 | p2 = chóngcǎo | l2 = worm-grass }} thumb|Microscopic detail of the fruiting body [[File:Yarchagumba.JPG|thumb|Fruiting body of ''O. sinensis'' यार्सागुम्बा in the Dolpo region of Nepal]]

'''''Ophiocordyceps sinensis''''' (synonym '''''Cordyceps sinensis'''''), known colloquially as '''caterpillar fungus''', is an entomopathogenic fungus (a fungus that grows on insects) in the family Ophiocordycipitaceae. It is mainly found in the meadows above {{convert|3500|m}} on the Tibetan Plateau regions of Tibet, Qinghai, Sichuan, Yunnan and Himalayan regions of Bhutan and Nepal. It parasitizes larvae of ghost moths and produces a fruiting body which is valued in traditional Chinese medicine and Nepalese medicine as an aphrodisiac.<ref>Interview with Britt Bunyard on {{cite news |title=Newshour |url=https://www.bbc.co.uk/programmes/w172z09lhvttylh |agency=BBC |date=Sep 9, 2023}}</ref> However, naturally harvested fruiting bodies often contain high amounts of arsenic and other heavy metals, making them potentially toxic. As a result, their sale has been strictly regulated by China's State Administration for Market Regulation since 2016.<ref>《总局关于停止冬虫夏草用于保健食品试点工作的通知 -食药监食监三〔2016〕21号》</ref>

''O. sinensis'' parasitizes the larvae of moths within the family Hepialidae, specifically genera found on the Tibetan Plateau and in the Himalayas, between elevations of {{convert|3000|and|5000|m|-3}}. The fungus germinates in the living larva, kills and mummifies it, and then a dark brown stalk-like fruiting body which is a few centimeters long emerges from the corpse and stands upright.

''O. sinensis'' is classified as a medicinal mushroom, and its use has a long history in traditional Chinese medicine as well as in traditional Tibetan medicine.<ref name=MM> {{cite book |author= Halpern, Miller |title= Medicinal Mushrooms |pages= 64–65 |publisher= M. Evans and Company, Inc. |location= New York, New York |year= 2002 |isbn= 978-0-87131-981-4 }}</ref> It is marketed for various health benefits but lacks sufficient scientific evidence for safety or effectiveness, and quality can vary due to inconsistent processing and labeling.<ref name="opss">{{cite web |title=Mushrooms in dietary supplements |url=https://www.opss.org/article/mushrooms-dietary-supplements |website=Operation Supplement Safety |publisher=Consortium for Health and Military Performance |date=2020-07-29 |access-date=2025-05-19}}</ref> The hand-collected, intact fungus-caterpillar body is valued by herbalists as medicine, and because of its cost, its use is also a status symbol.<ref>{{Cite web|url=https://www.npr.org/2011/10/09/141164173/caterpillar-fungus-the-viagra-of-the-himalayas|title=Caterpillar Fungus: The Viagra of the Himalayas|website=NPR.org }}</ref><ref name=yong2018> {{cite journal |url=https://www.theatlantic.com/science/archive/2018/10/tibetan-caterpillar-fungus-trouble/573607/ |title=The World's Most Valuable Parasite Is in Trouble. And so are the livelihoods of the people who depend on it. |first=Ed|last=Yong|journal= The Atlantic|date=2018-10-22}}</ref>

The fruiting bodies of the fungus are not cultivated commercially outside of China,<ref name="Hsieh">Hsieh, C., et al., A Systematic Review of the Mysterious Caterpillar Fungus Ophiocordyceps sinensis in Dong-ChongXiaCao and Related Bioactive Ingredients. Vol. 3. 2013. 16–32.</ref> but the mycelium form can be cultivated in vitro.<ref name="Yan">{{cite journal|year=2014|title=Recent advances in Cordyceps sinensis polysaccharides: mycelial fermentation, isolation, structure, and bioactivities: a review|journal=J Funct Foods|volume=6|pages=33–47|vauthors=Yan JK, Wang WQ, Wu JY|doi=10.1016/j.jff.2013.11.024|pmid=32362940|pmc=7185505}}</ref><ref name="Martel">{{cite journal|year=2017|title=Myths and realities surrounding the mysterious caterpillar fungus|journal=Trends Biotechnol|volume=35|issue=11|pages=1017–1021|vauthors=Martel J, Ko YF, Liau JC, Lee CS, Ojcius DM, Lai HC, Young JD|pmid=29055355|doi=10.1016/j.tibtech.2017.06.011}}</ref> Overharvesting and overexploitation have led to the classification of ''O. sinensis'' as an endangered species in China.<ref name="Xiao">{{cite journal|year=2011|title=Host insect species of Ophiocordyceps sinensis: a review|journal=ZooKeys|issue=127|pages=12743–59|doi=10.3897/zookeys.127.802|pmid=21998547|pmc=3175130|author1=Xiao-Liang W.|author2=Yi-Jian Y.|bibcode=2011ZooK..127...43W |doi-access=free}}</ref> Additional research needs to be carried out in order to understand its morphology and growth habits for conservation and optimum utilization.

==Taxonomic history and systematics== thumb|Caterpillars with emerging ''Ophiocordyceps sinensis''

===Morphological features=== ''Ophiocordyceps sinensis'' consists of two parts, a fungal endosclerotium (within the caterpillar) and stroma.<ref name="Shrestha">{{cite journal |author1=Shrestha B. |author2=Weimin Z. |author3=Yongjie Z. |author4=Xingzhong L. |year= 2010 |title= What is the Chinese caterpillar fungus Ophiocordyceps sinensis (Ophiocordycipitaceae)? |journal= Mycology |volume= 1 |issue= 4 |pages= 228–236 |doi= 10.1080/21501203.2010.536791|doi-access= free }}</ref> The stroma is the upper fungal part and is dark brown or black, but can be a yellow color when fresh, and longer than the caterpillar itself, usually 4–10 cm. It grows singly from the larval head, and is clavate, sublanceolate or fusiform, and distinct from the stipe (stalk).<ref name=Sung>{{cite journal |author= Sung G. H. |year= 2007 |title= A multi-gene phylogeny of Clavicipitaceae (Ascomycota, Fungi): identification of localized incongruence using a combinational bootstrap approach |journal= Molecular Phylogenetics and Evolution |volume= 44 |issue= 3 |pages= 1204–1223 |doi=10.1016/j.ympev.2007.03.011|display-authors=etal |pmid=17555990|bibcode= 2007MolPE..44.1204S }}</ref> The stipe is slender, glabrous, and longitudinally furrowed or ridged.

The fertile part of the stroma is the head. The head is granular because of the ostioles of the embedded perithecia.<ref name=Shrestha/> The perithecia are ordinally arranged and ovoid.<ref name=Sung/> The asci are cylindrical or slightly tapering at both ends, and may be straight or curved, with a capitate and hemispheroid apex, and may be two to four spored.<ref name=Shrestha/> Similarly, ascospores are hyaline, filiform, multiseptate at a length of 5–12 μm and subattenuated on both sides.<ref name=Sung/> Perithecial, ascus and ascospore characters in the fruiting bodies are the key identification characteristics of ''O. sinensis''.

''Ophiocordyceps'' (Petch) Kobayasi species produce whole ascospores and do not separate into part spores. This is different from other ''Cordyceps'' species, which produce either immersed or superficial perithecia perpendicular to stromal surface, and the ascospores at maturity are disarticulated into part spores.<ref name=Sung2007/> Generally ''Cordyceps'' species possess brightly colored and fleshy stromata, but ''O. sinensis'' has dark pigments and tough to pliant stromata, a typical characteristic feature of most of the ''Ophiocordyceps'' species.<ref name=Hsieh/>

===Developments in classification=== The species was first described scientifically by Miles Berkeley in 1843 as ''Sphaeria sinensis'';<ref name=Berkeley1843/> Pier Andrea Saccardo transferred the species to the genus ''Cordyceps'' in 1878.<ref name=Saccardo1878/> The fungus was known as ''Cordyceps sinensis'' until 2007, when molecular analysis was used to amend the classification of the Cordycipitaceae and the Clavicipitaceae, resulting in the naming of a new family Ophiocordycipitaceae and the transfer of several ''Cordyceps'' species including ''C. sinensis'' to the genus ''Ophiocordyceps''.<ref name=Sung2007/>

===Common names=== In Tibet, it is known as '''yartsa gunbu''', {{bo-textonly|དབྱར་རྩྭ་དགུན་འབུ་}} ({{bo|z=''yarza g̈unbu''|w=dbyar rtswa dgun 'bu|l=jɑ̀ːt͡sɑ kỹ̀pu}}, "summer grass winter worm"). The name was first recorded in the 15th century by the Tibetan doctor Zurkhar Namnyi Dorje. In colloquial Tibetan yartsa gunbu is often shortened to simply "bu" or "yartsa". The Tibetan name is transliterated in Nepali as यार्चागुन्बू, यार्चागुन्बा, '''''yarshagumba''''', '''''yarchagumba''''' or '''''yarsagumba'''''. The transliteration in Bhutan is '''{{Transliteration|dz|yartsa guenboob}}'''.

In India, it is known as '''''keera jhar''''', '''''keeda jadi''''', '''''keeda ghas''''' or '''{{Transliteration|ne|ghaas fafoond}}''' in Nepali, Hindi and Garhwali.

It is known in Chinese as '''{{Transliteration|zh|dōng chóng xià cǎo}}''' (冬蟲夏草), meaning "winter worm, summer grass", which is a literal translation of the original Tibetan name.<ref name=Lu2023>{{cite book |author=Lu D |title=The Global Circulation of Chinese Materia Medica, 1700-1949: A Microhistory of the Caterpillar Fungus |series=Medicine and Biomedical Sciences in Modern History |year=2023 |publisher=Palgrave Macmillan |pages=1–3 |doi=10.1007/978-3-031-24723-1 |isbn=978-3-031-24722-4 |s2cid=256618310 |url=https://link.springer.com/book/10.1007/978-3-031-24723-1 |language=en}}</ref> In traditional Chinese medicine, its name is often abbreviated as '''''chong cao''''' (蟲草 "insect plant"), a name that also applies to other ''Cordyceps'' species, such as ''C. militaris''. In Japanese, it is known by the Japanese reading of the characters for the Chinese name, {{nihongo|'''''tōchūkasō'''''|冬虫夏草}}, while in Korean and Vietnamese it is known by the transliterated forms of the Chinese word, '''''dongchunghacho''''' (동충하초) and '''''đông trùng hạ thảo''''' respectively. Strangely, sometimes in Chinese English-language texts ''Cordyceps sinensis'' is referred to as '''''aweto''''', which is the Māori name for ''Ophiocordyceps robertsii'', a species from south-eastern Australia and New Zealand.<ref>Hill H. Art. XXXVI: The Vegetable Caterpillar (Cordiceps robertsii). Transactions and Proceedings of the Royal Society of New Zealand. Vol 34, 1901;396–401</ref>

The English term "vegetable caterpillar" is a misnomer, as no plant is involved. "Caterpillar fungus" is a preferred term.

===Synonyms=== Since the 1980s, 22 species in 13 genera have been attributed to the anamorph (asexually reproducing mold-like form) of ''O. sinensis''.

{|class=wikitable |+Anamorphs attributed to ''O. sinensis'' |- ! Anamorph ! Correct Teleomorph ! Method for identification/Reference |- | ''Cephalosporium acreomonium'' | ''Umbelopsis'' | <ref name=JYY/> |- | ''Chrysosporium sinense'' | ? (not ''O. sinensis'') | RAPD polymorphism similarity<ref name=JYY/> |- | '''''Cephalosporium dongchongxiacae''''' (invalid name) | ''O. sinensis'' | <ref name=JYY/> |- | '''''Cephalosporium'' sp. sensu{{who|date=April 2025}}''' (incomplete name) | ''O. sinensis'' | <ref name=JYY/> |- | '''''Hirsutella sinensis''''' (invalid name but commonly used) | ''O. sinensis'' |;<ref name=JYY/> ITS sequence;<ref name="Liu">{{cite journal |author1=Liu Z. |author2=Liang Z. |author3=Liu A. |author4=Yao Y. |author5=Hyde K. D. |author6=Yu Z. |year= 2002|title= Molecular evidence for teleomorph-anamorph connections in Cordyceps based on ITS-5.8S rDNA sequences |journal= Mycological Research |volume= 106 |issue= 9 |pages= 1100–1108 |doi=10.1017/s0953756202006378|s2cid=86704330 }}</ref> microcyclic conidiation from ascospores and molecular studies<ref name="Shrestha" /> |- | '''''Hirsutella hepiali''''' | ''O. sinensis'' | <ref name=JYY/> |- | ''Cephalosporium sinensis'' | Possibly ''O. sinensis'' | there is lack of valid information <ref name=JYY/> |- | ''Isaria farinosa'' | ''Paecilomyces farinosus'' | <ref name=JYY/> |- | Isolates reported as ''Isaria'' sp., ''Verticella'' sp., ''Scydalium'' sp. | Unknown, identification is dubious | <ref name=JYY/> |- | ''Mortierella hepiali'' | Unknown Zygomycota | <ref name=JYY/> |- | ''Paecilomyces sinensis'' | ? (not ''O. sinensis'') | molecular evidence<ref name=JYY/> |- | ''Sporothrix insectorum'' | ? (not ''O. sinensis'') | molecular evidence<ref name=JYY/> |- | ''Paecilomyces lingi'' | ? | incomplete information, only appeared in one article<ref name=JYY/> |- | ''Tolypocladium sinense'' | ? | no valid information as of 2002<ref name=JYY/> |- | ''Paecilomyces hepiali'' | ? | Confirmed to be its own species, now ''Samsoniella hepiali''<ref name=Wang20/> |- | ''Tolypocladium sinense'' | ? | no valid information as of 2002<ref name=JYY>Jiang, Y. Y., & Yao, Y. J. (2002). Names related to Cordyceps sinensis anamorph. ''Mycotaxon'', 84:245-254.</ref> |}

Additional synonyms for the teleomorph are ''Cordyceps nepalensis'' and ''C. multiaxialis''. They have similar morphological characteristics to ''O. sinensis'', also had almost identical or identical ITS sequences and its presumed anamorph, ''H. sinensis''.<ref name="Liu"/>

==Ecology and life cycle== thumb|right|''Ophiocordyceps sinensis'' The caterpillars prone to infection by ''O. sinensis'' generally live {{convert|15|cm|in|abbr=on}} underground<ref name="Zhu">{{cite journal|year=1998|title=The scientific rediscovery of an ancient Chinese herbal medicine: Cordyceps sinensis: part I|journal=J Altern Complement Med|volume=4|issue=3|pages=289–303|doi=10.1089/acm.1998.4.3-289|pmid=9764768|vauthors=Zhu JS, Halpern GM, Jones K }}</ref> in alpine grass and shrub-lands on the Tibetan Plateau and the Himalayas at an altitude between {{convert|3000|and|5000|m|abbr=on|ft}}. The fungus is reported from the northern range of Nepal, Bhutan, and also from the northern states of India, apart from northern Yunnan, eastern Qinghai, eastern Tibet, western Sichuan, southwestern Gansu provinces.<ref name="Zhu"/> Fifty-seven taxa from several genera (37 ''Thitarodes'', 1 ''Bipectilus'', 1 ''Endoclita'', 1 ''Gazoryctra'', 3 ''Pharmacis'', and 14 others not correctly identified to genus<ref name="Hsieh" />) are recognized as potential hosts of ''O. sinensis''.

The stalk-like dark brown to black fruiting body (or mushroom) grows out of the head of the dead caterpillar and emerges from the soil in alpine meadows by early spring.<ref name="Stone">{{cite journal|year=2008|title=Last Stand for the Body Snatcher of the Himalayas?|url=http://english.xtbg.cas.cn/ns/es/201106/P020110609389437729680.pdf|journal=Science|volume=322|issue=5905|page=1182|doi=10.1126/science.322.5905.1182|pmid=19023056|author=Stone R|jstor=20145300|s2cid=206583137}}</ref> During late summer, the fruiting body disperses spores. The caterpillars, which live underground feeding on roots, are most vulnerable to the fungus after shedding their skin, during late summer. In late autumn, chemicals on the skin of the caterpillar interact with the fungal spores and release the fungal mycelia, which then infects the caterpillar.<ref name="Zhu"/>

The infected larvae tend to remain underground vertical to the soil surface with their heads up. After invading a host larva, the fungus ramifies throughout the host and eventually kills it. Gradually the host larvae become rigid because of the production of fungal sclerotia. Fungal sclerotia are multihyphal structures that can remain dormant and then germinate to produce spores. After overwintering, the fungus ruptures the host body, forming the fruiting body, a sexual sporulating structure (a perithecial stroma) from the larval head that is connected to the sclerotia in the dead larva below ground and grows upward to emerge from the soil to complete the cycle.<ref name="Zing">{{cite journal|year=2008|title=The Structure and Histochemistry of Sclerotia of Ophiocordyceps sinensis|journal=Mycologia|volume=100|issue=4|pages=616–625|author1=Xing X. K.|author2=Guo S. X.|jstor=20444986|doi=10.3852/07-007R2|pmid=18833754|s2cid=31450924}}</ref>

The slow growing ''O. sinensis'' grows at a comparatively low temperature, i.e., below 21 °C. Temperature requirements and growth rates are crucial factors that distinguish ''O. sinensis'' from other similar fungi.<ref name="Hsieh" /> Climate change is suspected to be negatively affecting the mountain organism.<ref>{{cite news |url=https://www.nytimes.com/2016/06/27/world/asia/himalayan-viagra-climate-change.html |title=Demand for 'Himalayan Viagra' Fungus Heats Up, Maybe Too Much |work=NYT |date=2016-06-26 |access-date=27 June 2016 |author=KAI SCHULTZ}}</ref><ref name=yong2018/>

== Use in traditional Asian medicines == [[File:Weighing Caterpillar fungus.jpg|thumb|upright|left|Weighing the precious caterpillar fungus in Yushu, Southern Qinghai, China]] The use of caterpillar fungus as folk medicine apparently originated in Tibet and Nepal. So far the oldest known text documenting its use was written in the late 15th century by the Tibetan doctor Zurkhar Nyamnyi Dorje (Wylie: {{Transliteration|bo|Zur mkhar mnyam nyid rdo rje}}) [1439–1475]) in his text: {{Transliteration|bo|Man ngag bye ba ring bsrel}} ("Instructions on a Myriad of Medicines"), where he describes its use as an aphrodisiac.<ref name=Winkler2008b/>

The first mention of ''Ophiocordyceps sinensis'' in traditional Chinese medicine was in Wang Ang's 1694 compendium of materia medica, ''Ben Cao Bei Yao''.<ref name=Winkler2008/> In the 18th century it was listed in Wu Yiluo's ''Ben cao cong xin'' ("New compilation of materia medica").<ref name=Wu1757/> The ethno-mycological knowledge on caterpillar fungus among the Nepalese people is documented.<ref>[http://www.nepjol.info/index.php/ON/article/viewFile/502/502 nepjol.info], Devkota(2006)</ref> The entire fungus-caterpillar combination is hand-collected for medicinal use.

In traditional Chinese medicine, it is regarded as having an excellent balance of yin and yang as it is considered to be composed of both an animal and a vegetable.<ref>{{Cite journal |last1=Siu |first1=Kai Ming |last2=Mak |first2=Duncan H. F. |last3=Chiu |first3=P. Y. |last4=Poon |first4=Michel K. T. |last5=Du |first5=Y. |last6=Ko |first6=Kam Ming |date=2004-12-10 |title=Pharmacological basis of 'Yin-nourishing' and 'Yang-invigorating' actions of Cordyceps, a Chinese tonifying herb |url=https://linkinghub.elsevier.com/retrieve/pii/S0024320504008173 |journal=Life Sciences |volume=76 |issue=4 |pages=385–395 |doi=10.1016/j.lfs.2004.07.014 |pmid=15530501 |issn=0024-3205|url-access=subscription }}</ref>

Wild-collected "cordyceps" is not always true ''O. sinensis'', even when the location and the host insect matches. This has resulted in the description of new species such as ''Cordyceps liangshanensis'' (Liangshan, China; many Nepali "cordyceps" are also incorrectly identified to be this species, and no new names for them have been proposed yet)<ref>{{cite journal |last1=Wang |first1=Y |last2=Dai |first2=YD |last3=Yang |first3=ZL |last4=Guo |first4=R |last5=Wang |first5=YB |last6=Yang |first6=ZL |last7=Ding |first7=L |last8=Yu |first8=H |title=Morphological and Molecular Phylogenetic Data of the Chinese Medicinal Fungus Cordyceps liangshanensis Reveal Its New Systematic Position in the Family Ophiocordycipitaceae. |journal=Mycobiology |date=2021 |volume=49 |issue=4 |pages=297–307 |doi=10.1080/12298093.2021.1923388 |pmid=34512076|pmc=8409936 }}</ref> and ''Samsoniella hepiali'' (Qinghai, China).<ref name=Wang20/> Despite not being the same species when examined using a modern method, these species are used in broadly the same way in traditional medicine.<ref name=culti-meta/> There are also "cordyceps" species that are traditionally known to be different from ''O. sinensis'' but nevertheless thought to be have a similar tonifying action. China is home to at least 299 species of "cordyceps" in this broadest sense.<ref name=sl>{{cite journal |title=中国广义虫草种类及其驯化栽培研究进展 |trans-title=Cordyceps Sensu Lato and Their Domestication and Cultivation in China |journal=食用菌学报 |date=2023-10-15 |volume=30 |issue=5 |doi=10.16488/j.cnki.1005-9873.2023.05.012 |url=https://www.syjxb.com/CN/10.16488/j.cnki.1005-9873.2023.05.012 |language=zh |issn=1005-9873}}</ref>

It is marketed for various health benefits but lacks sufficient scientific evidence for safety or effectiveness, and quality can vary due to inconsistent processing and labeling.<ref name=opss></ref>

=== Secondary metabolites === [[File:cordycepin.svg|thumb|Cordycepin]] A 2008 source reports that ''O. sinensis'' contains cordycepin, an adenosine derivative originally discovered in ''C. militaris''.<ref name="National Cancer Institute">{{cite web|author=National Cancer Institute|title=Definition of cordycepin|url=http://www.cancer.gov/publications/dictionaries/cancer-drug?cdrid=42667|website=NCI Drug Dictionary|access-date=21 December 2015|date=2011-02-02}}</ref> However, this study uses store-bought material labeled as ''O. sinensis'' without any molecular confirmation that it is indeed the species.<ref>{{cite journal |last1=Zhou |first1=X |last2=Luo |first2=L |last3=Dressel |first3=W |last4=Shadier |first4=G |last5=Krumbiegel |first5=D |last6=Schmidtke |first6=P |last7=Zepp |first7=F |last8=Meyer |first8=CU |title=Cordycepin is an immunoregulatory active ingredient of Cordyceps sinensis |journal=The American Journal of Chinese Medicine |date=2008 |volume=36 |issue=5 |pages=967–80 |doi=10.1142/S0192415X08006387 |pmid=19051361}}</ref> A more in-depth 2017 study, which fully characterized the biosynthetic machinery for cordycepin, found that ''O. sinensis'' does not produce cordycepin. This discrepancy underscores the importance of correctly identifying "cordyceps" species.<ref>{{cite journal |last1=Xia |first1=Yongliang |last2=Luo |first2=Feifei |last3=Shang |first3=Yanfang |last4=Chen |first4=Peilin |last5=Lu |first5=Yuzhen |last6=Wang |first6=Chengshu |title=Fungal Cordycepin Biosynthesis Is Coupled with the Production of the Safeguard Molecule Pentostatin |journal=Cell Chemical Biology |date=December 2017 |volume=24 |issue=12 |pages=1479–1489.e4 |doi=10.1016/j.chembiol.2017.09.001|pmid=29056419 }}</ref>

==Economics== In rural Tibet, ''yartsa gunbu'' has become the most important source of cash income. The fungi contributed 40% of the annual cash income to local households and 8.5% to the GDP in 2004. {{citation needed|date=June 2019}} Prices have increased continuously, especially since the late 1990s. In 2008, one kilogram traded for US$3,000 (lowest quality) to over US$18,000 (best quality, largest larvae). The annual production on the Tibetan Plateau was estimated in 2009 at 80–175 tons.<ref name=Winkler2009>{{cite journal |last1= Winkler |first1= D. |title= Caterpillar Fungus (Ophiocordyceps sinensis) Production and Sustainability on the Tibetan Plateau and in the Himalayas |doi= 10.1163/157342109X568829 |journal= Asian Medicine |volume= 5 |issue= 2 |pages= 291–316 |year= 2009 |url=http://app.quickblogcast.com/files/4/6/3/9/3/248966-239364/Winkler_Cordyceps_ICTAM_Bhutan.pdf|doi-access= free }}</ref> The Himalayan ''Ophiocordyceps'' production might not exceed a few tons.

In 2004 the value of a kilogram of caterpillars was estimated at 30,000 to 60,000 Nepali rupees in Nepal, and about Rs 100,000 in India.<ref name=Sharma2004/> In 2011, the value of a kilogram of caterpillars was estimated at 350,000 to 450,000 Nepali rupees in Nepal. A 2012 BBC article indicated that in north Indian villages a single fungus was worth Rs 150 (about £2 or $3), which is more than the daily wage of a manual labourer.<ref name="bbc20120707">{{cite news |url=https://www.bbc.co.uk/news/magazine-18735544 |title=The 'Viagra' transforming local economies in India |work=BBC News |date=2012-07-07 |access-date=July 9, 2012 |author=Jeffrey, Craig}}</ref> In 2012, a pound of top-quality yartsa had reached retail prices of $50,000.<ref>Finkel, M. (2012) [https://web.archive.org/web/20120719230226/http://ngm.nationalgeographic.com/2012/08/tibetan-mushroom/finkel-text Tibet's Golden "Worm"], National Geographic, August 2012</ref>

[[File:VM 5591 Lanzhou Dong chong xia cao.jpg|thumb|A shop in Lanzhou advertising ''Dōng chóng xià cǎo'' (冬虫夏草) among other local specialties.]] The price of ''Ophiocordyceps sinensis'' is reported to have increased dramatically on the Tibetan Plateau, about 900% between 1998 and 2008, an annual average of over 20% (after inflation). However, the value of large caterpillar fungus has increased more dramatically than small ''Cordyceps'', regarded as lower quality.<ref name=Winkler2008/> {| class="wikitable" style="text-align:center" ! Year !! % Price Increase !! Price/kg (Yuan) |- | 1980s|| ||align="center"|1,800 |- | 1997 || align="center"|467% (incl. inflation)|| align="center"|8,400 |- | 2004 || align="center"|429% (incl. inflation)|| align="center"|36,000 |- | 2005 |||| align="center"|10,000–60,000 |- | 2013 |||| align="center"|125,000–500,000 |- |}

In the Kingdom of Bhutan, ''Ophiocordyceps sinensis'' is recently also being harvested. The Bhutanese "cordyceps" has been molecularly confirmed to be ''O. sinensis'', with its quality shown to be equal to the Tibetan one.<ref>{{Cite journal|last1=Wu|first1=Ding-Tao|last2=Lv|first2=Guang-Ping|last3=Zheng|first3=Jian|last4=Li|first4=Qian|last5=Ma|first5=Shuang-Cheng|last6=Li|first6=Shao-Ping|last7=Zhao|first7=Jing|date=December 2016|title=Cordyceps collected from Bhutan, an appropriate alternative of Cordyceps sinensis|journal=Scientific Reports|volume=6|issue=1|article-number=37668|doi=10.1038/srep37668|issn=2045-2322|pmc=5118747|pmid=27874103|bibcode=2016NatSR...637668W}}</ref>

== Impacts of wild collection == === Societal impact === Because of its high value, inter-village conflicts over access to its grassland habitats has become a headache for the local governing bodies and in several cases people were killed. In November 2011, a court in Nepal convicted 19 villagers over the murder of a group of farmers during a fight over the prized aphrodisiac fungus. Seven farmers were killed in the remote northern district of Manang in June 2009 after going to forage for Yarchagumba.<ref>Staff (14 November 2011) [https://www.bbc.co.uk/news/world-asia-15741813 'Himalayan viagra': Six men get life for Nepal murders] BBC News Asia, Retrieved 9 July 2012</ref>

Its value gave it a role in the Nepalese Civil War, as the Nepalese Maoists and government forces fought for control of the lucrative export trade during the June–July harvest season.<ref name=Baral2005/> Collecting ''yarchagumba'' in Nepal had only been legalised in 2001, and now demand is highest in countries such as China, Thailand, Vietnam, Korea and Japan. By 2002, the 'herb' was valued at R 105,000 ($1,435) per kilogram, allowing the government to charge a royalty of R 20,000 ($280) per kilogram.

=== Ecological impact === The search for ''Ophiocordyceps sinensis'' is often perceived to threaten the environment of the Tibetan Plateau where it grows. While it has been collected for centuries and is still common in such areas, current collection rates are much higher than in historical times.

In India, fuelwood cutting by ''Ophiocordyceps sinensis'' collectors near the treeline is reported to be depleting populations of tree species such as Himalayan birch ''Betula utilis''.<ref>{{Cite journal |last1=Maletha |first1=Ajay |last2=Maikhuri |first2=R. K. |last3=Bargali |first3=S. S. |date=2020 |title=Criteria and indicator for assessing threat on Himalayan birch (B. utilis) at timberline ecotone of Nanda Devi Biosphere Reserve: A world heritage site, Western Himalaya, India |journal=Environmental and Sustainability Indicators |volume=8 |article-number=100086 |doi=10.1016/j.indic.2020.100086 |issn=2665-9727|doi-access=free |bibcode=2020EnvSI...800086M }}</ref>

== Cultivation ==

=== Mycelia === Cultivated ''O. sinensis'' mycelium is an alternative to wild-harvested ''O. sinensis'', and producers claim it may offer improved consistency. Artificial culture of ''O. sinensis'' is typically by growth of pure mycelia in liquid culture (in China) or on grains (in the West).{{citation needed|date=January 2018}}

''Ophiocordyceps sinensis'' is now cultivated on an industrial scale for their use in traditional Chinese medicine. However, no one has succeeded so far in rearing the fungus by infecting cultivated caterpillars;<ref name=Hsieh/> all products derived from cultured ''Ophiocordyceps'' are derived from mycelia grown on grains or in liquids.

In Chinese, the mycelia is used as a powder called {{lang-zh|s=发酵冬虫夏草菌粉|l=fermented winter-worm-summer-grass powder}}. However, the same term is also informally used for the powdered mycelia of the related ''Samsoniella hepiali'', also a pathogen of ghost-moths.

In Vietnam, according to the statistics of the Ministry of Agriculture and Rural Development, the production of cultivated "cordyceps" ({{lang|vi|đông trùng hạ thảo}}){{efn|The cited Vietnamese-language sources do not make it particularly clear whether the mentioned ''đông trùng hạ thảo'' is true ''O. sinensis''. Vietnam is also known to cultivate ''C. militaris''.}} in Vietnam in 2022 reached about 1,000 tons,<ref>{{Cite web |date=2022-09-01 |title=NGÀNH NÔNG NGHIỆP ĐẨY MẠNH XUẤT KHẨU, GIẢI QUYẾT CÁC RÀO CẢN KỸ THUẬT PHÁT SINH |url=https://sokhcn.baria-vungtau.gov.vn/Content/UserImages/300920221504Noi%20dung%20TBT%20T9%202022%20y.pdf |access-date=2023-12-12 |website=Bộ Nông nghiệp và Phát triển nông thôn}}</ref> an increase of five times compared to 2017. The selling price of fresh ''O. sinensis'' ranges from 10-20 million VND/kg,<ref>{{Cite web |last=NLD.COM.VN |title=Đông trùng hạ thảo, sâm Ngọc Linh.... không còn quá xa xỉ |url=https://nld.com.vn/duoc-lieu-quy-khong-con-qua-xa-xi-196231208200618016.htm |access-date=2023-12-12 |website=Báo Người Lao Động Online |language=vi}}</ref><ref>{{Cite web |last=ONLINE |first=TUOI TRE |date=2023-04-02 |title=Đông trùng hạ thảo 'đánh lận con đen' đến nhà khoa học cũng khó phân biệt thật giả |url=https://tuoitre.vn/dong-trung-ha-thao-danh-lan-con-den-den-nha-khoa-hoc-cung-kho-phan-biet-that-gia-20230402084711625.htm |access-date=2023-12-12 |website=TUOI TRE ONLINE |language=vi}}</ref> while dried ''O. sinensis'' ranges from 100-200 million VND/kg. Therefore, the economic value of cultivated "cordyceps" in Vietnam is estimated to be around 10,000 billion VND/year.<ref>{{Cite web |title=Nâng tầm giá trị cây dược liệu |url=https://lamdong.gov.vn/sites/lacduong/khkt/nong-lam-nghiep/SitePages/Nang-tam-gia-tri-cay-duoc-lieu.aspx |access-date=2023-12-12 |website=lamdong.gov.vn}}</ref><ref>{{Cite web |title=Phát triển các mô hình trồng và chế biến dược liệu giá trị cao |url=https://truyenhinhthanhhoa.vn/phat-trien-cac-mo-hinh-trong-va-che-bien-duoc-lieu-gia-tri-cao-1808387961.htm |access-date=2023-12-12 |website=Truyenhinhthanhhoa.vn |language=vi}}</ref> In the period 2017-2022, the production of cultivated "cordyceps" has grown at an average rate of 40%/year.<ref>{{Cite web |last=VnExpress |title=Việt Nam sản xuất cao từ nấm đông trùng hạ thảo |url=https://vnexpress.net/viet-nam-san-xuat-cao-tu-nam-dong-trung-ha-thao-4380022.html |access-date=2023-12-12 |website=vnexpress.net |language=vi}}</ref><ref>{{Cite web |title=Việt Nam sản xuất cao từ nấm đông trùng hạ thảo |url=https://www.most.gov.vn/vn/tin-tuc/21218/viet-nam-san-xuat-cao-tu-nam-dong-trung-ha-thao.aspx |access-date=2023-12-12 |website=www.most.gov.vn}}</ref>

=== Fruiting body === A fruiting body with a mature perithecium was first grown in laboratory conditions in China in 1983, using a growth media. By 2014, it was possible to obtain a mature fruiting body with a rice-based growth media in a low-altitude location. However, such amorphous culture media do not generate a product with the traditional presentation of "worm and grass".<ref name=sl/><ref>{{cite journal |last1=Cao |first1=L |last2=Ye |first2=Y |last3=Han |first3=R |title=Fruiting Body Production of the Medicinal Chinese Caterpillar Mushroom, Ophiocordyceps sinensis (Ascomycetes), in Artificial Medium. |journal=International Journal of Medicinal Mushrooms |date=2015 |volume=17 |issue=11 |pages=1107–12 |doi=10.1615/intjmedmushrooms.v17.i11.110 |pmid=26853966}}</ref>

Inoculation of caterpillars leading to a fully mature fruiting body was reported in 1991. This led to an early form of cultivation: caterpillars were artificially inoculated with the fungus, then placed into the natural habitat to induce the generation of the fruiting body. This increased the yield of the product, but was still subject to climate variations. In 2016, it became possible to mature the inoculated insects in a controlled environment. ''O. sinensis'' cultivated this way has been commercialized in China.<ref name=sl/>

Rearing of the caterpillars intended for inoculation can be affected by parasitic nematodes and other entomopathogenic fungi in the environment. Maturation of inoculated caterpillars can be disrupted by rodents eating them as food. Growth media may become a food source for ''Sciara'' flies.<ref name=sl/>

=== Cultivated alternatives === A number of related Hypocreales fungi have been collected in the field and found to be more amenable to cultivation. These fungi contain many of the same secondary metabolites as those in ''O. sinensis'' and Asian sources claim a similar spectrum of pharmacological effects. Some have also found use in traditional medicine as a substitute:

* ''Cordyceps militaris'' fruiting body is cultivated in China,<ref name=culti-meta>{{cite journal |last1=Lu |first1=Yi |last2=Zhi |first2=Yuee |last3=Miyakawa |first3=Takuya |last4=Tanokura |first4=Masaru |title=Metabolic profiling of natural and cultured Cordyceps by NMR spectroscopy |journal=Scientific Reports |date=2019-05-22 |volume=9 |issue=1 |page=7735 |doi=10.1038/s41598-019-44154-x|pmid=31118439 |pmc=6531489 |bibcode=2019NatSR...9.7735L |doi-access=free }}</ref> Vietnam,<ref>{{cite web |title=Báo VietnamNet |url=https://vietnamnet.vn/en/the-tibet-room-full-of-cordyceps-mushrooms-in-hanoi-E190386.html |website=VietNamNet News |language=vietnamese}}</ref> Taiwan, and Indonesia.<ref>{{cite web |title=Taiwanese company cultivates cordyceps in Indonesia |url=https://en.antaranews.com/news/108304/taiwanese-company-cultivates-cordyceps-in-indonesia |website=Antara News |language=en |date=10 December 2016}}</ref> The golden-colored fruiting body is eaten as an inexpensive mushroom. * ''Samsoniella hepiali'' mycelia is cultivated in China. It is used in two "Jinshuibao" products found in the Pharmacopoeia of the People's Republic of China and more than 260 healthcare products in China, with a total market worth of approximately 10 billion RMB.<ref name=Wang20>{{Cite journal |last1=Wang |first1=Yuan-Bing |last2=Wang |first2=Yao |last3=Fan |first3=Qi |last4=Duan |first4=Dong-E |last5=Zhang |first5=Guo-Dong |last6=Dai |first6=Ru-Qin |last7=Dai |first7=Yong-Dong |last8=Zeng |first8=Wen-Bo |last9=Chen |first9=Zi-Hong |last10=Li |first10=Dan-Dan |last11=Tang |first11=De-Xiang |last12=Xu |first12=Zhi-Hong |last13=Sun |first13=Tao |last14=Nguyen |first14=Thi-Tra |last15=Tran |first15=Ngoc-Lan |date=2020-07-01 |title=Multigene phylogeny of the family Cordycipitaceae (Hypocreales): new taxa and the new systematic position of the Chinese cordycipitoid fungus Paecilomyces hepiali |journal=Fungal Diversity |language=en |volume=103 |issue=1 |pages=1–46 |doi=10.1007/s13225-020-00457-3 |issn=1878-9129|doi-access=free }}</ref> * ''Paecilomyces tenuipes'' contains acetoxyscirpenediol and ergosterol peroxide.<ref name=culti-meta/> It is cultivated in Korea.<ref>{{cite journal |last1=Hong |first1=IP |last2=Nam |first2=SH |last3=Sung |first3=GB |last4=Chung |first4=IM |last5=Hur |first5=H |last6=Lee |first6=MW |last7=Kim |first7=MK |last8=Guo |first8=SX |title=Chemical Components of Paecilomyces tenuipes (Peck) Samson. |journal=Mycobiology |date=December 2007 |volume=35 |issue=4 |pages=215–8 |doi=10.4489/MYCO.2007.35.4.215 |pmid=24015100|pmc=3763175 }}</ref>

In some traditional-medicine contexts, it is acceptable to include the above alternatives in the term "cordyceps" or {{lang|zh|虫草}}.<ref name=culti-meta/><ref name=Wang20/> Li ''et al.'' (2023) (in Chinese) provides a more detailed overview of the cultivation of ''Cordyceps'' sensu lato globally.<ref name=sl/>

==References== {{notelist}} {{Reflist|colwidth=30em|refs= <ref name=Baral2005>{{cite journal |doi=10.2990/1471-5457(2005)24[2:TMPWAC]2.0.CO;2 |vauthors=Baral N, Heinen JT |year=2005 |title=The Maoist people's war and conservation in Nepal |journal=Politics and the Life Sciences |volume=24 |issue=1 |pages=2–11|pmid=17059316 |citeseerx=10.1.1.474.3446 |s2cid=21142810 }}</ref> <ref name=Berkeley1843>{{cite journal |author=Berkeley MJ. |title=On some entomogenous Sphaeriae |journal=London Journal of Botany |year=1843 |volume=2 |pages=205–11 |url=https://www.biodiversitylibrary.org/page/783487}}</ref> <ref name=Saccardo1878>{{cite journal |author=Saccardo PA. |title=Enumeratio Pyrenomycetum Hypocreaceorum hucusque cognitorum systemate carpologico dispositorum |journal=Michelia |year=1878 |volume=1 |issue=3 |pages=277–325 |url=http://cordyceps.us/files/P_A_Saccardo_Enumeratio_.pdf |language=la}}</ref> <ref name=Sharma2004>{{cite journal |author=Sharma S. |year=2004 |title=Trade of ''Cordyceps sinensis'' from high altitudes of the Indian Himalaya: Conservation and biotechnological priorities |journal=Current Science |volume=86 |issue=12 |pages=1614–9 |url=http://www.ias.ac.in/currsci/jun252004/1614.pdf }}</ref> <ref name=Sung2007>{{cite journal |vauthors=Sung GH, Hywel-Jones NL, Sung JM, Luangsa-Ard JJ, Shrestha B, Spatafora JW |title=Phylogenetic classification of ''Cordyceps'' and the clavicipitaceous fungi |journal=Studies in Mycology |year=2007 |volume=57 |pages=5–59 |pmid=18490993 |pmc=2104736 |doi=10.3114/sim.2007.57.01}}</ref> <ref name="urlMycoBank: Ophiocordyceps sinensis">{{cite web |title=''Ophiocordyceps sinensis'' (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora 2007 |url=https://www.mycobank.org/page/Name%20details%20page/438390 |website=MycoBank |publisher=International Mycological Association |access-date=2011-07-19}}</ref> <ref name=Winkler2008>{{cite journal |last=Winkler D. |year=2008 |title=Yartsa Gunbu (''Cordyceps sinensis'') and the fungal commodification of the rural economy in Tibet AR |journal=Economic Botany |volume=62 |issue=3 |pages=291–305 |doi=10.1007/s12231-008-9038-3|s2cid=29381859 }}</ref> <ref name=Winkler2008b>{{cite journal |author=Winkler D. |title=The mushrooming fungi market in Tibet exemplified by ''Cordyceps sinensis'' and ''Tricholoma matsutake'' |journal=Journal of the International Association of Tibetan Studies |year=2008 |issue=4 |url=http://www.thlib.org/collections/texts/jiats/#jiats=/04/winkler/}}</ref> <ref name=Wu1757>{{cite book |author=Wu Y |title="Ben cao cong xin" – "New compilation of materia medica" |year=1757 |language=zh}}</ref>  }}

==Further reading== *Winkler, D. 2005. Yartsa Gunbu – Cordyceps sinensis. Economy, Ecology & Ethno-mycology of a Fungus Endemic to the Tibetan Plateau. In: A. BOESI & F. CARDI (eds.). Wildlife and plants in traditional and modern Tibet: Conceptions, Exploitation and Conservation. Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, Vol. 33.1:69–85. *{{cite journal |author1=Zhang Y. |author2=Zhang S. |author3=Wang M. |author4=Bai F. |author5=Liu X. |year= 2010 |title= High Diversity of the Fungal Community Structure in Naturally-Occurring ''Ophiocordyceps sinensis'' |journal= PLoS ONE |volume= 5 |issue= 12 |article-number= e15570 |doi= 10.1371/journal.pone.0015570|pmid=21179540 |pmc=3002287 |bibcode=2010PLoSO...515570Z |doi-access=free }} *{{cite book |author=Lu D. |year=2023 |title=The Global Circulation of Chinese Materia Medica, 1700-1949: A Microhistory of the Caterpillar Fungus |series=Medicine and Biomedical Sciences in Modern History |url=https://link.springer.com/book/10.1007/978-3-031-24723-1 |publisher=Palgrave Macmillan |pages=1–294 |doi=10.1007/978-3-031-24723-1 |isbn=978-3-031-24722-4|s2cid=256618310 }} *{{cite book |author= Sułek, Emilia Róża. |year=2019 |title=Trading caterpillar fungus in Tibet: when economic boom hits rural area |series=Global Asia |place=Amsterdam |publisher=Amsterdam University Press |isbn=978-94-6298-526-1 }}

==External links== *{{Commons category-inline}} *{{Wikispecies-inline|Cordyceps sinensis}} *[http://mushroaming.com/Yartsa_Gunbu_Cordyceps Yartsa Gunbu (Cordyceps sinensis) in Tibet] *[http://cordyceps.us An Electronic Monograph of Cordyceps and Related Fungi] {{Webarchive|url=https://web.archive.org/web/20140517064543/http://cordyceps.us/ |date=2014-05-17 }} *[https://www.drugs.com/npp/cordyceps.html Cordyceps] information from Drugs.com *[http://libproject.hkbu.edu.hk/was40/detail?lang=en&channelid=1288&searchword=herb_id=D00019 Cordyceps sinensis (Berk.) Sacc.] Medicinal Plant Images Database (School of Chinese Medicine, Hong Kong Baptist University) {{in lang|en}} {{in lang|zh-hant}} *[https://web.archive.org/web/20120719230226/http://ngm.nationalgeographic.com/2012/08/tibetan-mushroom/finkel-text Tibet's Golden "Worm"] August 2012 National Geographic (magazine) {{Taxonbar|from=Q398300}} {{Authority control}}

Category:Fungi described in 1843 Category:Fungi of Asia Category:Ophiocordycipitaceae Category:Medicinal fungi Category:Fungi used in traditional Chinese medicine Category:Taxa named by Miles Joseph Berkeley Category:Parasitic fungi Category:Fungus species Category:Fungi in cultivation