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Amanita muscaria

Amanita muscaria, commonly known as the fly agaric or fly Amanita is a poisonous and psychoactive basidiomycete fungus, one of many in the genus Amanita. Native throughout the temperate and boreal regions of the Northern Hemisphere, Amanita muscaria has been unintentionally conveyed to many countries in the Southern Hemisphere, generally as a symbiont with pine plantations, and is now a true cosmopolitan species. It associates with various deciduous and coniferous trees. The quintessential toadstool, it is a large imposing white-gilled, white-spotted, usually deep red mushroom, one of the most recognizable and widely encountered in popular culture.

Though generally considered poisonous, Amanita muscaria is otherwise famed for its hallucinogenic properties with its main psychoactive constituent being the compound muscimol. Used as an intoxicant by the Koryaks of the Kamchatka Krai of eastern Siberia, the mushroom has had a religious significance in Siberian culture and possibly also in ancient Scandinavian culture. The American banker and amateur ethnomycologist R. Gordon Wasson proposed the fly agaric was in fact the Soma talked about in the ancient Rig Veda texts of India; although this theory has been refuted by anthropologists, it gained common credence when first published in 1968.

The common name in English is thought to have been derived from its European use as an insecticide, sprinkled in milk. The fly-killing agent is now known to be ibotenic acid. Another compound isolated from the fungus is 1,3-diolein which is an insect attractor. 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.

Taxonomy and naming

Similar to its English common name, the German Fliegenpilz, Dutch Vliegenzwam, Swedish Röd flugsvamp, Danish Rød fluesvamp, Finnish punakärpässieni, Polish muchomor, Slovak muchotrávka, Croatian muhara, Hungarian légyölő galóca and French Amanite tue-mouches, are derived from its use in Europe as an insecticide, sprinkled in milk. This practice has been recorded from Germanic- (bar England) and Slavic-speaking parts of Europe, as well as the bilingual Vosges region and pockets elsewhere in France, and Romania. Albertus Magnus was the first to record it in his work De vegetabilibus sometime before 1256, commenting:

The 16th century Flemish botanist Charles de l'Écluse localised the practice to Frankfurt in Germany, while the father of taxonomy Linnaeus reported it from Småland in southern Sweden where he had lived as a child. He officially described it in Volume Two of his Species Plantarum in 1753, giving it the name Agaricus muscarius, the specific epithet deriving from Latin musca meaning "fly". It gained its current name in 1783, when placed in the genus Amanita by Jean-Baptiste Lamarck and sanctioned by Elias Magnus Fries.

The starting date had been formerly set as January 1st 1821, the date of the works of the 'father of mycology', Swedish naturalist Elias Magnus Fries, and under these conditions, the full name was Amanita muscaria (L.:Fr.) Hook.. However, a recent revision of the International Code of Botanical Nomenclature in 1987 changed the rules regarding the starting date and primary work for names of fungi, and now names can be considered valid as far back as May 1st 1753, the date of publication of Linneaus' seminal work. Hence, Linnaeus and Lamarck become the authorities and the name has become Amanita muscaria (L.) Lam.

English mycologist John Ramsbottom reported it was used for getting rid of bugs in England and Sweden, and bug agaric was an old alternate name. French mycologist Pierre Bulliard tried to replicate its fly-killing properties without success in his work Histoire des plantes vénéneuses et suspectes de la France, and proposed a new binomial name Agaricus pseudo-aurantiacus because of this.

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. Several regional names appear to be linked with this connotation, meaning either 'mad- or 'fool's 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 the Province of Trento in Italy. A local dialect name in Fribourg in Switzerland is tsapi de diablhou 'Devil's hat'.

The word toadstool in English does not refer to any particular species, yet it has a more definite specific connotation with A. muscaria in continental Europe. Yet another name is crapaudin in many parts of France, and a Basque term from Guipúzcoa and Biscay is amoroto, all alluding to toads. Finally, a common name from China is ha-ma chün 'toad mushroom' (蛤蟆菌). Interestingly, the toad lacks the negative connotations in Chinese culture and symbolism.

Wasson proposed this was due to its being a shamanic and also taboo object and hence unable to be named specifically in ancient Celtic culture. He speculates that the power of this taboo may have perpetuated its malign reputation while other lethal fungi such as the death cap (A. phalloides) have had little cultural connotations throughout European history.

Ethnobotanist and ethnomycologist Giorgio Samorini has proposed a symbiotic relationship between toads, flies and the fly agaric. After a lick of A. muscaria, flies become inebriated and delirious prey for hungry toads that may have learned this, therefore hanging out around toadstools. This relationship within nature illuminates an etymological keystone and example of zoopharmacognosy. This would also provide further biosemiotic insight into the ancient mystery of toads, flies and mushrooms appearing together in popular mythology and fairy lore. However it is more commonly thought that toads are associated with the mushroom because they symbolise toxicity and chthonic forces in the same way that serpents do.

Classification

Amanita muscaria is the type species of the genus Amanita. By extension, it is also the type species of Amanita subgenus Amanita, as well as section Amanita within this subgenus. Amanita subgenus Amanita includes all Amanita with inamyloid spores. Amanita section Amanita includes those species with patchy universal veil remnants, including a volva that is reduced to a series of concentric rings and the veil remnants on the pileus being a series of patches or warts. Most species in this group also have a bulbous base.

Amanita section Amanita consists of A. muscaria and its close relatives, including A. pantherina (the panther cap), Amanita gemmata, A. farinosa, and A. xanthocephala. Modern fungal taxonomists have classified Amanita muscaria and its allies this way based on gross morphology and spore inamyloidy. Two recent molecular phylogenetic studies have confirmed this classification as natural.

Amanita muscaria varies considerably in its morphology and many authorities recognize a number of subspecies or varieties within the species. In The Agaricales in Modern Taxonomy, Rolf Singer listed three subspecies, though without description: A. muscaria ssp. muscaria, A. muscaria ssp. americana, and A. muscaria ssp. flavivolvata.

Contemporary authorities recognize up to seven varieties:

var. muscaria, the typical red-and-white spotted variety. Some authorities, such as Rodham Tulloss, only use this name for Eurasian and western Alaskan populations.
var. flavivolvata is red, with yellow to yellowish-white warts, and occurs in the western regions of the North American continent, from southern Alaska down through the Rocky Mountains, through Central America, to at least Andean Colombia. Rodham Tulloss uses this name to describe all "typical" A. muscaria from indigenous New World populations from Alaska southward.
var. alba, an uncommon fungus, has a white to silvery white cap with white warts but otherwise similar to the usual form.
var. formosa, has a yellow to orange-yellow cap with yellowish or tan warts and stem. Some authorities use this name for all A. muscaria fitting this description worldwide (cf, Jenkins), others (cf, Tulloss) restrict its use to Eurasian populations.
var. guessowii is yellow to orange, with center of cap more orange or reddish orange than the outer part. It is found throughout North America, but is most common in northeastern North America, from Newfoundland and Quebec down to Tennessee. Some authorities (cf, Jenkins) treat these populations as part of A. muscaria var. formosa, while others (cf, Tulloss) recognize it as a distinct variety.
var. persicina is pinkish to orangish "melon" colored with poorly formed or absent remnants of universal veil on the stem and vasal bulb, known from the Southeastern Coastal areas of the U.S.A, described in 1977.
var. regalis (= Amanita regalis (Fr.) Michael), from Scandinavia and Alaska, is liver-brown and has yellow warts. It appears to be uniformly distinctive and some authorities (cf, Tulloss) treat it as a separate species, while others (cf, Jenkins) treat it as a variety of A. muscaria.

A 2006 molecular phylogenetic study of different regional populations of A. muscaria by Geml, et al. found three distinct clades within this species representing, roughly, Eurasian, Eurasian "subalpine", and North American populations. (Alaska contains examples of all three clades, leading to the hypothesis that this was the center of diversification of this species.) The study also looked at four named varieties of this species; var. alba, var. flavivolvata, var. formosa (including var. guessowii), and var. regalis from both areas. All four varieties were found within both the Eurasian and North American clades, evidence that these morphological forms are simply polymorphisms found throughout the species rather than distinct subspecies or varieties.

Description

A large conspicuous mushroom, Amanita muscaria is generally common and numerous where it grows, often being found in groups with basidiocarps in all stages of development. Fully grown, the bright red cap is usually around 8-20 cm (3-8 in) in diameter, though larger specimens have been found. The red colour may fade after rain and in older mushrooms. After emerging from the ground, the cap is covered with numerous small white to yellow flecks (warts) which are remnants of the universal veil, a membrane that encloses the entire mushroom when it is still very young. The gills are white, as is the spore print. The stem is white, 5-20 cm high (approximately 2-8 in), with a basal bulb that bears universal veil remnants (more or less distinct rings or ruffs), and has the slightly brittle, fibrous texture typical of many large mushrooms. 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.

Fly agaric fruiting bodies emerge from the soil looking like a white egg, covered in the white warty material of the universal veil. As the fungus grows, the red colour appears through the broken veil, and the cap changes from hemispherical to plate-like and flat in mature specimens.

Though very distinctive, the fly agaric has been mistaken for other yellow to red species in the Americas such as Armillaria cf. mellea and the edible Amanita basii, a Mexican species similar to A. caesarea of Europe. Poison control centers in the U.S. and Canada are aware that amarill is a common name of A. caesarea-like species in Mexico, not just the Spanish for 'yellow'.

Amanita caesarea can be distinguished as it has an entire orange red cap, lacking the numerous white warty spots of the fly agaric. Furthermore the stem, gills and ring are bright yellow, not white. Finally the volva is a distinct white bag, not broken into scales.

In Australia, the introduced fly agaric may be confused with the native vermilion grisette (Amanita xanthocephala), which grows in association with Eucalypts. The latter species generally lacks the white warts of A. muscaria and bears no ring.

Distribution and habitat

A. muscaria is a cosmopolitan mushroom, native to birch, pine, spruce, fir and cedar woodlands throughout the temperate and boreal regions of the Northern Hemisphere, including high elevations of warmer latitudes in regions like the Hindu Kush, the Mediterranean and Central America. Interestingly, a recent molecular study proposes an ancestral origin in the Siberian–Beringian region in the Tertiary period before radiating outwards across Asia, Europe and North America. Though generally encountered in autumn, the season can vary 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. It is often found in similar locations to Boletus edulis, and may appear in fairy rings. It has been widely transported into the southern hemisphere, including Australia, New Zealand, South Africa and South America, where it usually occurs under introduced pine trees.

The fungal equivalent of a weed in southeastern Australia, it appears to have formed new associations with southern beech (Nothofagus) in Tasmania and Victoria and invading native rain forest, where there are concerns it may be displacing native species. Furthermore it appears to be spreading northwards with recent reports near Port Macquarie on the New South Wales north coast.

When imported to a new country, A. muscaria can jump to native species (for example, Eucalyptus in Australia). It can then be exported with its new symbiont (for example, from Australia to Argentina).

Toxicity

Victims of Amanita muscaria poisoning are generally either young children or people ingesting it for a hallucinogenic experience. Occasionally, immature button forms have been mistaken for edible puffballs. About of A. muscaria or 50-100 mg ibotenic acid is considered a toxic dose. Ibotenic acid, a compound present in A. muscaria, has shown to be highly neurotoxic when injected directly into the brains of mice and rats.

Deaths from A. muscaria are extremely rare. A historical journal article reported two fatalities occurring in North America. With modern medical treatment the prognosis is generally good. A fatal dose has been calculated at approximately 15 fly agaric caps, although the amount and ratio of chemical compounds per mushroom varies widely from region to region, season to season, further confusing the issue. It has been reported that spring and summer mushrooms may contain up to 10 times as much ibotenic acid and muscimol compared to fall fruitings. Many older books mistakenly list it as deadly, giving the impression that it is far more toxic than it really is. The vast majority of mushroom poisoning fatalities (90% or more) are from having eaten either the greenish to yellowish death cap (A. phalloides) or one of the several Amanita species known as destroying angel.

The active constituents of this species are water soluble, and boiling and then discarding the cooking water will at least partly detoxify A. muscaria. However, drying may increase potency as the process facilitates the conversion of ibotenic acid to the more potent muscimol. According to some sources, once detoxified, the mushroom becomes edible. In Sanada, Japan, fly agarics are detoxified by pickling them.

Pharmacology

Amanita muscaria contains a number of biologically active agents, at least two of which are known to be psychoactive. Muscimol (3hydroxy-5-aminomethy-1 isoxazole, an unsaturated cyclic hydroxamic acid) is the most significant. It is the product of the decarboxylation or drying of ibotenic acid, another important compound in the biochemistry of the fly agaric. Muscarine, discovered in 1869, was long thought to be the active hallucinogenic agent in A. muscaria until the mid 20th century, when researchers in England, Japan, and Switzerland recognized that these effects were due mainly to ibotenic acid and muscimol.

Ibotenic acid and muscimol are structurally related to each other and to two major neurotransmitters of the central nervous system: glutamic acid and GABA respectively. Ibotenic acid and muscimol act like these neurotransmitters (muscimol is a potent GABA_A agonist, while ibotenic acid is an agonist of NMDA glutamate receptors and certain metabotropic glutamate receptors) which are involved in the control of neuronal activity. It is these interactions which are thought to cause the psychoactive effects found in intoxication. Following ingestion some of the ibotenic acid is metabolised to muscimol which would appear to be the agent responsible for the majority of the psychoactivity. Indeed, ibotenic acid's strong water solubility means that it could not distribute into the brain without an active process like a transporter; this, and the lack of any reports of permanent brain damage following A. muscaria ingestion (which would be the result of ibotenic acid entering the brain due to its effect as an NMDA receptor agonist) make it unlikely that ibotenic acid enters the brain following A. muscaria ingestion. However, it is worth noting that no studies have directly investigated ibotenic acid's ability to permeate the brain.

Muscazone is another compound more recently isolated from European specimens of the fly agaric. It is a product of the breakdown of ibotenic acid by ultra-violet radiation. It is of minor pharmacological activity compared with the other agents.

Muscarine binds with Muscarinic acetylcholine receptor and lead to the excitation of the neurons bearing these receptors. The levels in Amanita muscaria are minute when compared with other poisonous fungi, such as the deadly Inocybe erubescens or small white Clitocybe species C. dealbata and C. rivulosa, and are too insignificant to play a role in the symptoms of poisoning.

Amanita muscaria and related species are known as effective accumulators of vanadium (up to 500 mg/kg in dry weight). Vanadium is present as an organometallic compound (called amavadine) in fruit-bodies. However, the biological importance of the accumulation process is unknown.

Symptoms

Fly agarics are known for the unpredictability of their effects. Depending on habitat and the amount ingested per body weight, effects can range from nausea and twitching to drowsiness, cholinergic effects (low blood pressure, sweating and salivation), auditory and visual distortions, mood changes, euphoria, relaxation, and loss of equilibrium. Retrograde amnesia frequently results following recovery. In cases of serious poisoning it causes a delirium, characterized by bouts of marked agitation with confusion, hallucinations, and irritability followed by periods of central nervous system depression. Seizures and coma may also occur in severe poisonings. Effects typically appear after around 30 to 90 minutes and peak within three hours, but certain effects can last for a number of days. In the majority of cases recovery is complete within 12 hours. The effect is highly variable and individuals can react quite differently to the similar doses. Some cases of intoxication have exhibited headaches up to ten hours afterwards.

Treatment

Medical attention should be sought in cases of suspected poisoning. If the delay between ingestion and treatment is less than four hours, inducing vomiting with Syrup of ipecac is warranted. However, this should be avoided if there is a reduced level of consciousness and gastric lavage performed instead. Intravenous rehydration may required for recurrent vomiting but this is rare.

Psychoactive use

Unlike the hallucinogenic mushrooms of the Psilocybe, Amanita muscaria has been rarely consumed recreationally. It is classified as an unscheduled drug in the United States. Any sales of A. muscaria for human ingestion are regulated by the Food and Drug Administration. Most other countries do not have laws against the use of A. muscaria, as it is currently legal and un-controlled under United Nations international law. However, following the outlawing of psilocybin-containing mushrooms in the United Kingdom, an increased quantity of Amanita mushrooms began to be sold and consumed.

In eastern Siberia, the shaman would consume the mushrooms, and others would drink his urine. This urine, still containing active hallucinogens may actually 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. Among the Koryak, one report held the poor would consume the urine of the wealthy, who could afford to buy the mushrooms. If a fly agaric is eaten, it is usually not fresh, but in its dried or cooked form, where ibotenic acid is converted to the more stable and far less poisonous muscimol.

Siberia

A. muscaria was widely used as a hallucinogenic drug by many of the indigenous peoples of Siberia. Its use was known among almost all of the Uralic-speaking peoples of western Siberia and the Paleosiberian-speaking peoples of eastern Siberia. However, there are only isolated reports of A. muscaria use among the Tungusic and Turkic peoples of central Siberia and it is believed that hallucinogenic use of A. muscaria was largely not a practice of these peoples. In western Siberia, the use of A. muscaria was restricted to shamans, who used it as an alternate method of achieving a trance state. (Normally, Siberian shamans achieve a trance state 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.

The Koryak of eastern Siberia have a story about the fly agaric (wapaq) which enabled Big Raven to carry a whale to its home. In the story, the deity Vahiyinin ("Existence") spat onto earth, and his 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, can learn from it.

Beyond Siberia, there are only isolated and unconfirmed reports of the hallucinogenic use of A. muscaria. One source mentions that it was once used among the Sami people, but no firsthand accounts of this exist. Hartmut Geerken claims to have discovered a tradition of recreational use of this mushroom among a Parachi-speaking group in Afghanistan. There are also unconfirmed reports of religious use of A. muscaria among two Subarctic Native American tribes, the Ojibway and the Dogrib. Authors of Sacred Soma Shamans Hawk and Venus have used A. muscaria as a sacrament and daily medicinal tonic since 1979.

Soma

There are also claims that A. muscaria played in important role in a number of ancient religious rites, though these claims tend to be speculative and highly controversial. The best known of these claims is R. Gordon Wasson's proposition that A. muscaria was the Soma talked about in the Rig Veda of India, and is less often also thought to be the amrita talked about in Buddhist scriptures. (For more details on this topic, see Botanical identity of Soma-Haoma.)

Nordic Vikings

A single source for the notion that Nordic Vikings used A. muscaria to produce their berserker rages was first suggested by the Swedish professor Samuel Ödman in 1784. Ödman 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. Today, it is generally considered an urban legend or at best speculation that cannot be proven. On the whole, muscimol, the psychoactive ingredient, is a mild relaxant, but it is widely known, as with all drugs that it can create a range of reactions within a range of people and it is possible that it could make a person incredibly angry, as well as make them "very jolly or sad, [jump] about, [dance], [sing] or [give] way to great fright".

Catholicism

John Marco Allegro argues in The Sacred Mushroom and the Cross that the Roman Theology is derived from a sex and psychedelic mushroom cult,, although his theory has found little support by scholars outside the field of ethnomycology. In Magic Mushrooms in Religion and Alchemy (formerly called Strange Fruit) Clark Heinrich interprets A. muscaria usage by Adam and Eve, Moses, Elijah and Elisha, Isaiah, Ezekiel, Jonah, Jesus and his disciples, and John of Patmos. In the book Apples of Apollo the mushroom is identified in a wide range of mythological tales such as those involving Perseus, Prometheus, Heracles, Jason and the Argonauts, Jesus and the Holy Grail.

Cultural depictions

The red-and-white spotted toadstool is a common image in many aspects of popular culture, especially in children's books, film and more recently computer games; a partly grown A. muscaria, as shown right, is clearly the fungus upon which this icon is based.

Art

Images dating back to 3500 BC painted in caves at Tassili, Algeria, depict mushrooms, more than likely including A. muscaria mushrooms according to scholars

, although a Psilocybe species has also been suggested.

Fly agarics have been featured in paintings since the Renaissance , albeit in a subtle manner. In the Victorian era they became more visible, even becoming the main topic of some fairy paintings , usually inspired by Shakespeare's A Midsummer Night's Dream. The waning of Romanticism and the advent of World War I reduced interest in fairies along with fly agarics, reducing them to the realm of childish fantasies.

Literature and entertainment

Garden ornaments, and children's picture books depicting gnomes and fairies, such as the Smurfs, very often show fly agarics used as seats, or homes. Two of the most famous uses of the mushroom are in the video game series Super Mario Bros. and the dancing mushroom sequence in the 1940 Disney film Fantasia. They also have appearances in Metal Gear Solid 3: Snake Eater, Katamari Damacy, and The Elder Scrolls IV: Oblivion. These mushrooms also make an appearance in Quest for Glory 1 VGA as the mushroom ring. There is an Amanita Muscaria on the cover of the album "Fungus Amongus" by Incubus.

Christmas decorations and Santa Claus

Fly agarics appear on Christmas cards and New Year cards from around the world as a symbol of good luck. They also function as Christmas tree decorations , derived from their ectomycorrhizal relationship with coniferous trees. The ethnobotanist Jonathan Ott has suggested that the idea of Santa Claus and tradition of hanging stockings over the fireplace is based centrally upon the fly agaric mushroom itself. With its generally red and white color scheme, he argues that Santa Claus's suit is related to the mushroom. He also draws parallels with flying reindeer: reindeer had been reported to consume the mushroom and prance around in an intoxicated manner afterwards. Until the 20th century the red-and-white Santa suit familiar today was not firmly established, although Saint Nicholas, on which Santa Claus is partly based, was always depicted in red (see also: Origins of Santa Claus). One scholar researching possible links between religious myths and the red mushroom notes, "If Santa Claus had but one eye [like Odin], or if magic urine had been a part of his legend, his connection to the Amanita muscaria would be much easier to believe."

Ott also speculates about Santa's bag of toys. According to historians, ancient Siberia was one of the first civilizations to use fly agaric in practice. The Siberian hut, or yurt, is equipped with a smokehole at the top. Ott suggests that a shaman entered the yurt through the smokehole with a sack of mushrooms in his hand, to be placed in stockings over the fireplace where they could be dried for celebratory use.