Amanita muscaria

Amanita muscaria
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Basidiomycota
Class:	Agaricomycetes
Order:	Agaricales
Family:	Amanitaceae
Genus:	Amanita
Species:	A. muscaria
Binomial name
Amanita muscaria (L.) Lam. (1783)
Subspecies and varieties
A. muscaria var. alba A. muscaria subsp. flavivolvata Singer[1][2] A. muscaria var. guessowii Veselý[3] A. muscaria var. inzengae Neville & Poumarat[4][2]

Amanita muscaria mycorrhizal
	Edibility is poisonous or psychoactive

Amanita muscaria, commonly known as the fly agaric or fly amanita,

Despite its easily distinguishable features, A. muscaria is a fungus with several known variations, or subspecies. These subspecies are slightly different, some having yellow or white caps, but are all usually called fly agarics, most often recognizable by their notable white spots. Recent DNA fungi research, however, has shown that some mushrooms called 'fly agaric' are in fact unique species, such as A. persicina (the peach-colored fly agaric).

Native throughout the

The 16th-century Flemish botanist

The English mycologist John Ramsbottom reported that Amanita muscaria was used for getting rid of bugs in England and Sweden, and bug agaric was an old alternative name for the species.^[12] French mycologist Pierre Bulliard reported having tried without success to replicate its fly-killing properties in his work Histoire des plantes vénéneuses et suspectes de la France (1784), and proposed a new binomial name Agaricus pseudo-aurantiacus because of this.^[18] One compound isolated from the fungus is 1,3-diolein (1,3-di(cis-9-octadecenoyl)glycerol), which attracts insects.^[19] It has been hypothesised that the flies intentionally seek out the fly agaric for its intoxicating properties.^[20] An alternative derivation proposes that the term fly- refers not to insects as such but rather the delirium resulting from consumption of the fungus. This is based on the medieval belief that flies could enter a person's head and cause mental illness.^[21] Several regional names appear to be linked with this connotation, meaning the "mad" or "fool's" version of the highly regarded edible mushroom Amanita caesarea. Hence there is oriol foll "mad oriol" in Catalan, mujolo folo from Toulouse, concourlo fouolo from the Aveyron department in Southern France, ovolo matto from Trentino in Italy. A local dialect name in Fribourg in Switzerland is tsapi de diablhou, which translates as "Devil's hat".^[22]

Classification

Amanita muscaria is the

A large, conspicuous mushroom, Amanita muscaria is generally common and numerous where it grows, and is often found in groups with basidiocarps in all stages of development. Fly agaric fruiting bodies emerge from the soil looking like white eggs. After emerging from the ground, the cap is covered with numerous small white to yellow pyramid-shaped warts. These are remnants of the universal veil, a membrane that encloses the entire mushroom when it is still very young. Dissecting the mushroom at this stage reveals a characteristic yellowish layer of skin under the veil, which helps identification. As the fungus grows, the red colour appears through the broken veil and the warts become less prominent; they do not change in size, but are reduced relative to the expanding skin area. The cap changes from globose to hemispherical, and finally to plate-like and flat in mature specimens.^[28] Fully grown, the bright red cap is usually around 8–20 centimetres (3–8 inches) in diameter, although larger specimens have been found. The red colour may fade after rain and in older mushrooms.

The free gills are white, as is the spore print. The oval spores measure 9–13 by 6.5–9 μm; they do not turn blue with the application of iodine.^[29] The stipe is white, 5–20 cm (2–8 in) high by 1–2 cm (1⁄2–1 in) wide, and has the slightly brittle, fibrous texture typical of many large mushrooms. At the base is a bulb that bears universal veil remnants in the form of two to four distinct rings or ruffs. Between the basal universal veil remnants and gills are remnants of the partial veil (which covers the gills during development) in the form of a white ring. It can be quite wide and flaccid with age. There is generally no associated smell other than a mild earthiness.^[30][31]

Although very distinctive in appearance, the fly agaric has been mistaken for other yellow to red mushroom species in the Americas, such as

Controversy

Amanita muscaria varies considerably in its morphology, and many authorities recognize several subspecies or varieties within the species. In The Agaricales in Modern Taxonomy, German mycologist Rolf Singer listed three subspecies, though without description: A. muscaria ssp. muscaria, A. muscaria ssp. americana, and A. muscaria ssp. flavivolvata.^[23]

However, a 2006 molecular phylogenetic study of different regional populations of A. muscaria by mycologist József Geml and colleagues found three distinct

species complex.^[38] The complex also includes at least three other closely related taxa that are currently regarded as species:^[1] A. breckonii is a buff-capped mushroom associated with conifers from the Pacific Northwest,^[39] and the brown-capped A. gioiosa and A. heterochroma from the Mediterranean Basin and from Sardinia respectively. Both of these last two are found with Eucalyptus and Cistus trees, and it is unclear whether they are native or introduced from Australia.^[40][41]

Amanitaceae.org lists four varieties as of May 2019[update], but says that they will be segregated into their own taxa "in the near future". They are:^[2]

Image	Reference name	Common name	Synonym	Description
	Amanita muscaria var. muscaria[1]	Euro-Asian fly agaric		Bright red fly agaric from northern Europe and Asia. Cap might be orange or yellow due to slow development of the purple pigment. Wide cap with white or yellow warts which are removed by rain. Known to be toxic but used by shamans in northern cultures. Associated predominantly with Birch and diverse conifers in forest.
	Amanita muscaria subsp. flavivolvata[3]	American fly agaric		red, with yellow to yellowish-white warts. It is found from southern Alaska down through the Rocky Mountains, through Central America, all the way to Andean Colombia. Rodham Tulloss uses this name to describe all "typical" A. muscaria from indigenous New World populations.
	Amanita muscaria var. guessowii[4]	American fly agaric (yellow variant)	Amanita muscaria var. formosa	has a yellow to orange cap, with the centre more orange or perhaps even reddish orange. It is found most commonly in northeastern North America, from Newfoundland and Quebec south all the way to the state of Tennessee. Some authorities (cf. Jenkins) treat these populations as A. muscaria var. formosa, while others (cf. Tulloss) recognise them as a distinct variety.
	Amanita muscaria var. inzengae[42]	Inzenga's fly agaric		it has a pale yellow to orange-yellow cap with yellowish warts and stem which may be tan.

Distribution and habitat

A. muscaria is a

temperate and boreal regions of the Northern Hemisphere,^[37] including higher elevations of warmer latitudes in regions such as Hindu Kush, the Mediterranean and also Central America. A recent molecular study proposes that it had an ancestral origin in the Siberian–Beringian region in the Tertiary period, before radiating outwards across Asia, Europe and North America.^[37] The season for fruiting varies in different climates: fruiting occurs in summer and autumn across most of North America, but later in autumn and early winter on the Pacific coast. This species is often found in similar locations to Boletus edulis, and may appear in fairy rings.^[43] Conveyed with pine seedlings, it has been widely transported into the southern hemisphere, including Australia,^[44] New Zealand,^[45] South Africa^[46] and South America, where it can be found in the Brazilian states of Paraná,^[37] São Paulo, Minas Gerais, Rio Grande do Sul.^[47]

Manjimup, Western Australia in 2010.^[51] Although it has apparently not spread to eucalypts in Australia, it has been recorded associating with them in Portugal. Commonly found throughout the great Southern region of western Australia, it is regularly found growing on Pinus radiata.^[52]

Toxicity

A. muscaria poisoning has occurred in young children and in people who ingested the mushrooms for a

hallucinogenic experience,^[21][53][54]

or who confused it with an edible species.

A. muscaria contains several biologically active agents, at least one of which,

psychoactive. Ibotenic acid, a neurotoxin, serves as a prodrug to muscimol, with a small amount likely converting to muscimol after ingestion. An active dose in adults is approximately 6 mg muscimol or 30 to 60 mg ibotenic acid;^[55][56] this is typically about the amount found in one cap of Amanita muscaria.^[57] The amount and ratio of chemical compounds per mushroom varies widely from region to region and season to season, which can further confuse the issue. Spring and summer mushrooms have been reported to contain up to 10 times more ibotenic acid and muscimol than autumn fruitings.^[53]

Deaths from A. muscaria have been reported in historical journal articles and newspaper reports,[58]^[59][60] but with modern medical treatment, fatal poisoning from ingesting this mushroom is extremely rare.^[61] Many books list A. muscaria as deadly,^[62] but according to David Arora, this is an error that implies the mushroom is far more toxic than it is.^[63] Furthermore, The North American Mycological Association has stated that there were "no reliably documented cases of death from toxins in these mushrooms in the past 100 years".^[64]

The active constituents of this species are water-soluble, and boiling and then discarding the cooking water at least partly detoxifies A. muscaria.^[65] Drying may increase potency, as the process facilitates the conversion of ibotenic acid to the more potent muscimol.^[66] According to some sources, once detoxified, the mushroom becomes edible.^[67][68] Patrick Harding describes the Sami custom of processing the fly agaric through reindeer.^[69]

Pharmacology

Muscarine, discovered in 1869,^[70] was long thought to be the active hallucinogenic agent in A. muscaria. Muscarine binds with muscarinic acetylcholine receptors leading to the excitation of neurons bearing these receptors. The levels of muscarine in Amanita muscaria are minute when compared with other poisonous fungi^[71] such as Inosperma erubescens, the small white Clitocybe species C. dealbata and C. rivulosa. The level of muscarine in A. muscaria is too low to play a role in the symptoms of poisoning.^[72]

The major toxins involved in A. muscaria poisoning are

unsaturated cyclic hydroxamic acid) and the related amino acid ibotenic acid. Muscimol is the product of the decarboxylation (usually by drying) of ibotenic acid. Muscimol and ibotenic acid were discovered in the mid-20th century.^[73][74] Researchers in England,^[75] Japan,^[76] and Switzerland^[74] showed that the effects produced were due mainly to ibotenic acid and muscimol, not muscarine.^[19][73] These toxins are not distributed uniformly in the mushroom. Most are detected in the cap of the fruit, a moderate amount in the base, with the smallest amount in the stalk.^[77][78] Quite rapidly, between 20 and 90 minutes after ingestion, a substantial fraction of ibotenic acid is excreted unmetabolised in the urine of the consumer. Almost no muscimol is excreted when pure ibotenic acid is eaten, but muscimol is detectable in the urine after eating A. muscaria, which contains both ibotenic acid and muscimol.^[56]

Ibotenic acid and muscimol are structurally related to each other and to two major

GABA_A agonist, while ibotenic acid is an agonist of NMDA glutamate receptors and certain metabotropic glutamate receptors^[79] which are involved in the control of neuronal activity. It is these interactions which are thought to cause the psychoactive effects found in intoxication.^[21][57]

amavadine.^[81] The biological importance of the accumulation process is unknown.^[82]

Symptoms

Fly agarics are best known for the unpredictability of their effects. Depending on habitat and the amount ingested per body weight, effects can range from mild

loss of equilibrium (like with tetanus.)^{[53][54][57][59]}

In cases of serious poisoning the mushroom causes

Seizures and coma may also occur in severe poisonings.^[54][57] Symptoms typically appear after around 30 to 90 minutes and peak within three hours, but certain effects can last for several days.^[32][56] In the majority of cases recovery is complete within 12 to 24 hours.^[65] The effect is highly variable between individuals, with similar doses potentially causing quite different reactions.^[53][56][83] Some people suffering intoxication have exhibited headaches up to ten hours afterwards.^[56] Retrograde amnesia and somnolence can result following recovery.^[57]

Treatment

Medical attention should be sought in cases of suspected poisoning. If the delay between ingestion and treatment is less than four hours,

activated charcoal is given. Gastric lavage can be considered if the patient presents within one hour of ingestion.^[84] Inducing vomiting with syrup of ipecac is no longer recommended in any poisoning situation.^[85]

There is no antidote, and supportive care is the mainstay of further treatment for intoxication. Though sometimes referred to as a

respiratory depressant effects of muscimol.^[87] Recurrent vomiting is rare, but if present may lead to fluid and electrolyte imbalances; intravenous rehydration or electrolyte replacement may be required.^[57][88] Serious cases may develop loss of consciousness or coma, and may need intubation and artificial ventilation.^[54][89] Hemodialysis can remove the toxins, although this intervention is generally considered unnecessary.^[65] With modern medical treatment the prognosis is typically good following supportive treatment.^[61][65]

Uses

Psychoactive

The wide range of psychoactive effects have been variously described as depressant, sedative-hypnotic, psychedelic, dissociative, or deliriant; paradoxical effects such as stimulation may occur however. Perceptual phenomena such as synesthesia, macropsia, and micropsia may occur; the latter two effects may occur either simultaneously or alternatingly, as part of Alice in Wonderland syndrome, collectively known as dysmetropsia, along with related distortions pelopsia and teleopsia. Some users report lucid dreaming under the influence of its hypnotic effects. Unlike Psilocybe cubensis, A. muscaria cannot be commercially cultivated, due to its mycorrhizal relationship with the roots of pine trees. However, following the outlawing of psilocybin mushrooms in the United Kingdom in 2006, the sale of the still legal A. muscaria began increasing.^[90]

shamanic rituals. The Lithuanian festivities are the only report that Wasson received of ingestion of fly agaric for religious use in Eastern Europe.^[91]

Siberia

A. muscaria was widely used as an

Paleosiberian-speaking peoples of the Russian Far East. There are only isolated reports of A. muscaria use among the Tungusic and Turkic peoples of central Siberia and it is believed that on the whole entheogenic use of A. muscaria was not practised by these peoples.^[6] In western Siberia, the use of A. muscaria was restricted to shamans, who used it as an alternative method of achieving a trance state. (Normally, Siberian shamans achieve trance by prolonged drumming and dancing.) In eastern Siberia, A. muscaria was used by both shamans and laypeople alike, and was used recreationally as well as religiously.^[6] In eastern Siberia, the shaman would take the mushrooms, and others would drink his urine.^[92] This urine, still containing psychoactive elements, may be more potent than the A. muscaria mushrooms with fewer negative effects such as sweating and twitching, suggesting that the initial user may act as a screening filter for other components in the mushroom.^[93]

The

spittle became the wapaq, and his saliva becomes the warts. After experiencing the power of the wapaq, Raven was so exhilarated that he told it to grow forever on earth so his children, the people, could learn from it.^[94] Among the Koryaks, one report said that the poor would consume the urine of the wealthy, who could afford to buy the mushrooms.^[95] It was reported that the local reindeer would often follow an individual intoxicated by the muscimol mushroom, and if said individual were to urinate in snow the reindeer would become similarly intoxicated and the Koryak people's would use the drunken state of the reindeer to more easily rope and hunt them.^[96]

Other reports and theories

The Finnish historian

Tlicho use of Amanita muscaria.^[102] The flying reindeer of Santa Claus, who is called Joulupukki in Finland, could symbolize the use of A. muscaria by Sámi shamans.^{[103][104][105]} However, Sámi scholars and the Sámi peoples themselves refute any connection between Santa Claus and Sámi history or culture^[106].

"The story of Santa emerging from a Sámi shamanic tradition has a critical number of flaws," asserts Tim Frandy, assistant professor of Nordic Studies at the University of British Columbia and a member of the Sámi descendent community in North America. "The theory has been widely criticized by Sámi people as a stereotypical and problematic romanticized misreading of actual Sámi culture."^[106]

Vikings

The notion that

Vikings used A. muscaria to produce their berserker rages was first suggested by the Swedish professor Samuel Ödmann in 1784.^[107] Ödmann based his theories on reports about the use of fly agaric among Siberian shamans. The notion has become widespread since the 19th century, but no contemporary sources mention this use or anything similar in their description of berserkers. Muscimol is generally a mild relaxant, but it can create a range of different reactions within a group of people.^[108] It is possible that it could make a person angry, or cause them to be "very jolly or sad, jump about, dance, sing or give way to great fright".^[108] Comparative analysis of symptoms have, however, since shown Hyoscyamus niger to be a better fit to the state that characterises the berserker rage.^[109]

Soma

See also:

Botanical identity of Soma-Haoma

In 1968,

Philologist, archaeologist, and

Amanita muscaria is traditionally used for catching flies possibly due to its content of ibotenic acid and muscimol, which lead to its common name "fly agaric". Recently, an analysis of nine different methods for preparing A. muscaria for catching flies in Slovenia have shown that the release of ibotenic acid and muscimol did not depend on the solvent (milk or water) and that thermal and mechanical processing led to faster extraction of ibotenic acid and muscimol.^[123]

Culinary

The toxins in A. muscaria are water-soluble: parboiling A. muscaria fruit bodies can detoxify them and render them edible,

Smurfs, often show fly agarics used as seats, or homes.^[29][128] Fly agarics have been featured in paintings since the Renaissance,^[129] albeit in a subtle manner. For instance, in Hieronymus Bosch's painting, The Garden of Earthly Delights, the mushroom can be seen on the left-hand panel of the work.^[130] In the Victorian era they became more visible, becoming the main topic of some fairy paintings.^[131] Two of the most famous uses of the mushroom are in the Mario franchise (specifically two of the Super Mushroom power-up items and the platforms in several stages which are based on a fly agaric),^[132][133] and the dancing mushroom sequence in the 1940 Disney film Fantasia.^[134]

An account of the journeys of Philip von Strahlenberg to Siberia and his descriptions of the use of the mukhomor there was published in English in 1736. The drinking of urine of those who had consumed the mushroom was commented on by Anglo-Irish writer Oliver Goldsmith in his widely read 1762 novel, Citizen of the World.^[135] The mushroom had been identified as the fly agaric by this time.^[136] Other authors recorded the distortions of the size of perceived objects while intoxicated by the fungus, including naturalist Mordecai Cubitt Cooke in his books The Seven Sisters of Sleep and A Plain and Easy Account of British Fungi.^[137] This observation is thought to have formed the basis of the effects of eating the mushroom in the 1865 popular story Alice's Adventures in Wonderland.^[138] A hallucinogenic "scarlet toadstool" from Lappland is featured as a plot element in Charles Kingsley's 1866 novel Hereward the Wake based on the medieval figure of the same name.^[139] Thomas Pynchon's 1973 novel Gravity's Rainbow describes the fungus as a "relative of the poisonous Destroying angel" and presents a detailed description of a character preparing a cookie bake mixture from harvested Amanita muscaria.^[140] Fly agaric shamanism is also explored in the 2003 novel Thursbitch by Alan Garner.^[141]

See also

• List of Amanita species
• Legal status of psychoactive Amanita mushrooms

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Works cited

• ISBN 978-0-9825562-7-6
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• ISBN 978-0-89815-169-5
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• Benjamin, Denis R. (1995). Mushrooms: poisons and panaceas—a handbook for naturalists, mycologists and physicians. New York: WH Freeman and Company.
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• Furst, Peter T. (1976). Hallucinogens and Culture. Chandler & Sharp. pp. 98–106.
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• Letcher, Andy (2006). Shroom: A Cultural history of the magic mushroom. London: Faber and Faber.
  ISBN 978-0-571-22770-9
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• Ramsbottom, J. (1953). Mushrooms & Toadstools. Collins.
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• Wasson, R. Gordon (1968). Soma: Divine Mushroom of Immortality. Harcourt Brace Jovanovick.
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External links

Wikimedia Commons has media related to Amanita muscaria.

Wikispecies has information related to Amanita muscaria.

• Webpages on Amanita species by Tulloss and Yang Zhuliang
• Amanita on erowid.org
• Aminita muscaria, Amanita pantherina and others (Group PIM G026) by
  IPCS INCHEM