Amanita muscaria


Amanita muscaria, commonly known as the fly agaric or fly amanita, is a basidiomycete fungus of the genus Amanita. Its common name derives from its traditional use as an insecticide. It is a distinctive, large white-gilled mushroom typically featuring a bright red cap covered with white warts. The complex genetic diversity of A.muscaria suggests that it is a species complex. It is a widely distributed mushroom native to temperate and boreal forests of the Northern Hemisphere, now also naturalised in the Southern Hemisphere, forming symbiotic relationships with various trees and spreading invasively in some regions.
Ingestion of the mushroom can cause poisoning, especially in children and those seeking its hallucinogenic effects, due to psychoactive compounds like muscimol and the ibotenic acid; however, fatal poisonings are extremely rare. Parboiling reduces toxicity, though drying converts ibotenic acid into muscimol, retaining psychoactive effects. Some cultures use it as food after preparation. Indigenous peoples of Siberia used A.muscaria as an inebriant and entheogen. It has been controversially linked to Santa Claus, Viking berserkers, Vedic soma, and early Christianity, though evidence is sparse and disputed. Its rise in the 2020s as a legal hallucinogen alternative has led to Food and Drug Administration scrutiny in the United States of America.
A.muscaria has appeared in art and literature since the Renaissance, becoming iconic in fairy tales, children’s books, and media like Disney’s Fantasia and the Super Mario video games. It has also influenced literary depictions of altered perception—most notably in Alice’s Adventures in Wonderland—and has been referenced in novels by writers including Oliver Goldsmith, Thomas Pynchon, and Alan Garner.

Taxonomy

The name of the mushroom in many European languages is thought to derive from its use as an insecticide when sprinkled in milk. This practice has been recorded from Germanic- and Slavic-speaking parts of Europe, as well as the Vosges region and pockets elsewhere in France, and Romania. Albertus Magnus was the first to record it in his work De vegetabilibus some time before 1256, commenting, "it is called the fly mushroom because it is powdered in milk to kill flies".
The 16th-century Flemish botanist Carolus Clusius traced the practice of sprinkling it into milk to Frankfurt in Germany, while Carl Linnaeus, the "father of taxonomy", reported it from Småland in southern Sweden, where he had lived as a child. He 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, a name sanctioned in 1821 by the "father of mycology", Swedish naturalist Elias Magnus Fries. The starting date for all the mycota had been set by general agreement as January 1, 1821, the date of Fries's work, and so the full name was then Amanita muscaria Hook. The 1987 edition of the International Code of Botanical Nomenclature changed the rules on the starting date and primary work for names of fungi, and names can now be considered valid as far back as May 1, 1753, the date of publication of Linnaeus's work. Hence, Linnaeus and Lamarck are now taken as the namers of Amanita muscaria Lam..
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. 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, and proposed a new binomial name Agaricus pseudo-aurantiacus because of this. One compound isolated from the fungus is 1,3-diolein, which attracts insects.
It has been hypothesised that the flies intentionally seek out the fly agaric for its intoxicating properties.
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 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".

Classification

Amanita muscaria is the type species of the genus. 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 the species with patchy universal veil remnants, including a volva that is reduced to a series of concentric rings, and the veil remnants on the cap to 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, A. gemmata, A. farinosa, and A. xanthocephala. Modern fungal taxonomists have classified A. muscaria and its allies this way based on gross morphology and spore inamyloidy. Two recent molecular phylogenetic studies have confirmed this classification as natural.

Subdivision

Amanita muscaria varies considerably in its morphology and many authorities recognise several subspecies or varieties within the species. In 1986, German mycologist Rolf Singer listed three subspecies : A. muscaria ssp. muscaria, A. muscaria ssp. americana, and A. muscaria ssp. flavivolvata.
A 2006 molecular phylogenetic study of different regional populations of A. muscaria by mycologist József Geml and colleagues found three distinct clades within the species, roughly representing Eurasian, Eurasian "subalpine", and North American populations. Specimens belonging to all three clades have been found in Alaska; this has led to the hypothesis that this was the centre of diversification for this species. The study also looked at four named varieties of the species: var. alba, var. flavivolvata, var. formosa, and var. regalis from both areas. All four were found within both the Eurasian and North American clades, evidence that the forms are polymorphisms rather than distinct subspecies or varieties. Further molecular study by Geml and colleagues published in 2008 show that these three genetic groups, plus a fourth associated with oak–hickory–pine forest in the southeastern United States and two more on Santa Cruz Island, California, are delineated from each other enough genetically to be considered separate species. Thus A. muscaria as it stands currently is, evidently, a species complex. The complex also includes at least three other closely related taxa that are currently regarded as species: A. breckonii is a buff-capped mushroom associated with conifers from the Pacific Northwest, 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.

Description

A large, conspicuous mushroom, A. 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 has an irregular distribution 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 in mature specimens. The bright red cap ranges from in diameter. Age and rain may cause the red colour to fade and the warts to fall off.
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. The stipe is white, high by 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 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.

Similar species

Although very distinctive in appearance, the fly agaric has been mistaken for other yellow to red mushroom species in the Americas, such as Armillaria cf. mellea and the edible A. basii—a Mexican species similar to A. caesarea of Europe. Poison control centres in the U.S. and Canada have become aware that is a common name for the A. caesarea-like species in Mexico. A. caesarea is distinguished by its entirely orange to red cap, which lacks the numerous white warty spots of the fly agaric. Furthermore, the stem, gills and ring of A. caesarea are bright yellow, not white. The volva is a distinct white bag, not broken into scales. In Australia, the introduced fly agaric may be confused with the native A. xanthocephala, which grows in association with eucalypts. The latter species generally lacks the white warts of A. muscaria and bears no ring. Additionally, immature button forms resemble puffballs. A. muscaria is also morphologically similar to closely related species in A. stirps Muscaria, including A. persicina, and A. chrysoblema.