Amanita bisporigera | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Class: | Agaricomycetes |
Order: | Agaricales |
Family: | Amanitaceae |
Genus: | Amanita |
Species: | A. bisporigera |
Binomial name | |
Amanita bisporigera G.F.Atk. (1906) | |
Synonyms [1] [2] | |
|
Amanita bisporigera | |
---|---|
Gills on hymenium | |
Cap is convex or flat | |
Hymenium is free | |
Stipe has a ring and volva | |
Spore print is white | |
Ecology is mycorrhizal | |
Edibility is deadly |
Amanita bisporigera is a deadly poisonous species of fungus in the family Amanitaceae. It is commonly known as the eastern destroying angel amanita, [3] the eastern North American destroying angel or just as the destroying angel , although the fungus shares this latter name with three other lethal white Amanita species, A. ocreata , A. verna and A. virosa . The mushroom has a smooth white cap that can reach up to 10 centimetres (4 inches) across and a stipe up to 14 cm (5+1⁄2 in) tall with a white skirt-like ring near the top. The bulbous stipe base is covered with a membranous sac-like volva. The white gills are free from attachment to the stalk and crowded closely together. As the species name suggests, A. bisporigera typically bears two spores on the basidia, although this characteristic is not immutable. A. bisporigera closely resembles a few other white amanitas, including the equally deadly A. virosa and A. verna.
A. bisporigera was described as a new species in 1906. It is classified in the section Phalloideae of the genus Amanita together with other amatoxin-containing species. The species is found in mixed coniferous and deciduous forests of eastern North America south to Mexico, but are rare in western North America. The first symptoms of poisoning appear 6 to 24 hours after consumption, followed by a period of apparent improvement, then by symptoms of liver and kidney failure, and death after four days or more.
The cap is 3–10 centimetres (1–4 inches) in diameter and, depending on its age, ranges in shape from egg-shaped to convex to somewhat flattened. The cap surface is smooth and white, sometimes with a pale tan- or cream-colored tint in the center. The surface is either dry or, when the environment is moist, slightly sticky. The flesh is thin and white, and does not change color when bruised. [4] The margin of the cap, which is rolled inwards in young specimens, does not have striations (grooves), and lacks volval remnants. The gills, also white, are crowded closely together. They are either free from attachment to the stipe or just barely reach it. The lamellulae (short gills that do not extend all the way to the stipe) are numerous, and gradually narrow.
The white stipe is 6–14 cm (2+1⁄2–5+1⁄2 in) by 0.7–1.8 cm (1⁄4–3⁄4 in) thick, solid (i.e., not hollow), and tapers slightly upward. The surface, in young specimens especially, is frequently floccose (covered with tufts of soft hair), fibrillose (covered with small slender fibers), or squamulose (covered with small scales); there may be fine grooves along its length. The bulb at the base of the stipe is spherical or nearly so. The delicate ring on the upper part of the stipe is a remnant of the partial veil that extends from the cap margin to the stalk and covers the gills during development. It is white, thin, membranous, and hangs like a skirt. When young, the mushrooms are enveloped in a membrane called the universal veil, which stretches from the top of the cap to the bottom of the stipe, imparting an oval, egg-like appearance. In mature fruit bodies, the veil's remnants form a membrane around the base, the volva, like an eggshell-shaped cup. On occasion, however, the volva remains underground or gets torn up during development. It is white, sometimes lobed, and may become pressed closely to the stipe. [5] The volva is up to 3.8 cm (1+1⁄2 in) in height (measured from the base of the bulb), and is about 2 mm thick midway between the top and the base attachment. [6] The mushroom's odor has been described as "pleasant to somewhat nauseous", [4] becoming more cloying as the fruit body ages. [6] The cap flesh turns yellow when a solution of potassium hydroxide (KOH, 5–10%) is applied (a common chemical test used in mushroom identification). This characteristic chemical reaction is shared with A. ocreata and A. virosa, although some authors have expressed doubt about the identity of North American A. virosa, suggesting those collections may represent four-spored A. bisporigera. [6] [7] Tulloss suggests that reports of A. bisporigera that do not turn yellow with KOH were actually based on white forms of A. phalloides . [6] Findings from the Chiricahua Mountains of Arizona and in central Mexico, although "nearly identical" to A. bisporigera, do not stain yellow with KOH; their taxonomic status has not been investigated in detail. [2]
The spore print of A. bisporigera, like most Amanita, is white. The spores are roughly spherical, thin-walled, hyaline (translucent), amyloid, and measure 7.8–9.6 by 7.0–9.0 μm. The cap cuticle is made of partially gelatinized, filamentous interwoven hyphae, 2–6 μm in diameter. The tissue of the gill is bilateral, meaning it diverges from the center of the gill to its outer edge. The subhymenium is ramose—composed of relatively thin branching, unclamped hyphae. The spore-bearing cells, the basidia, are club-shaped, thin-walled, without clamps, with dimensions of 34–45 by 4–11 μm. They are typically two-spored, although rarely three- or four-spored forms have been found. [5] Although the two-spored basidia are a defining characteristic of the species, there is evidence of a tendency to shift towards producing four-spored basidia as the fruiting season progresses. [6] The volva is composed almost exclusively of densely interwoven filamentous hyphae, 2–10 μm in diameter, that are sparsely to moderately branched. There are few small inflated cells, which are mostly spherical to broadly elliptic. The tissue of the stipe is made of abundant, sparsely branched, filamentous hyphae, without clamps, measuring 2–5 μm in diameter. The inflated cells are club-shaped, longitudinally oriented, and up to 2–3 by 15.7 μm. The annulus is made of abundant moderately branched filamentous hyphae, measuring 2–6 μm in diameter. The inflated cells are sparse, broadly elliptic to pear-shaped, and are rarely larger than 31 by 22 μm. [5] Pleurocystidia and cheilocystidia (cystidia found on the gill faces and edges, respectively) are absent, but there may be cylindrical to sac-like cells of the partial veil on the gill edges; these cells are hyaline and measure 24–34 by 7–16 μm. [4]
In 1906 Charles E. Lewis studied and illustrated the development of the basidia in order to compare the nuclear behavior of the two-spored with that of the four-spored forms. Initially (1), the young basidium, appearing as a club-shaped branch from the subhymenium, is filled with cytoplasm and contains two primary nuclei, which have distinct nucleoli. As the basidium grows larger, the membranes of the two nuclei contact (2), and then the membrane disappears at the point of contact (3). The two primary nuclei remain distinct for a short time, but eventually the two nuclei fuse completely to form a larger secondary nucleus with a single secondary nucleolus (4, 5). The basidium increases in size after the primary nuclei fuse, and the nucleus migrates towards the end of the basidia (6, 7). During this time, the nucleus develops vacuoles "filled by the nuclear sap in the living cell". Chromosomes are produced from the nucleolar threads, and align transversely near the apex of the basidium, connected by spindles (8–10). The chromosomes then move to the poles, forming the daughter nuclei that occupy different positions in the basidium; the daughters now have a structure similar to that of the parent nuclei (11). The two nuclei then divide to form four nuclei, similar to fungi with four-spored basidia (12, 13). The four nuclei crowd together at some distance from the end of the basidium to form an irregular mass (14). Shortly thereafter, the sterigmata (slender projections of the basidia that attach the spores) begin to form (15), and cytoplasm begins to pass through the sterigmata to form the spores (16). Although Lewis was not able to clearly determine from observation alone whether the contents of two or four nuclei passed through the sterigmata, he deduced, by examining older basidia with mature spores, that only two nuclei enter the spores (16, 17). [8]
The Amanita Genome Project was begun in Jonathan Walton's lab at Michigan State University in 2004 as part of their ongoing studies of A. bisporigera. [9] The purpose of the project is to determine the genes and genetic controls associated with the formation of mycorrhizae, and to elucidate the biochemical mechanisms of toxin production. The genome of A. bisporigera has been sequenced [10] using a combination of automated Sanger sequencing and pyrosequencing, and the genome sequence information is publicly searchable. [11] The sequence data enabled the researchers to identify the genes responsible for amatoxin and phallotoxin biosynthesis, AMA1 and PHA1. The cyclic peptides are synthesized on ribosomes, and require proline-specific peptidases from the prolyl oligopeptidase family for processing. [12] [13]
The genetic sequence information from A. bisporigera has been used to identify molecular polymorphisms in the related A. phalloides. These single-nucleotide polymorphisms may be used as population genetic markers to study phylogeography and population genetics. [14] Sequence information has also been employed to show that A. bisporigera lacks many of the major classes of secreted enzymes that break down the complex polysaccharides of plant cell walls, like cellulose. In contrast, saprobic fungi like Coprinopsis cinerea and Galerina marginata , which break down organic matter to obtain nutrients, have a more complete complement of cell wall-degrading enzymes. Although few ectomycorrhizal fungi have yet been tested in this way, the authors suggest that the absence of plant cell wall-degrading ability may correlate with the ectomycorrhizal ecological niche. [15]
The color and general appearance of A. bisporigera are similar to those of A. verna and A. virosa . A. bisporigera is at times smaller and more slender than either A. verna or A. virosa, but it varies considerably in size; therefore size is not a reliable diagnostic characteristic. [4] A. virosa fruits in autumn—later than A. bisporigera. [16] A. elliptosperma is less common but widely distributed in the southeastern United States, while A. ocreata is found on the West Coast and in the Southwest. Other similar toxic North American species include Amanita magnivelaris , which has a cream-colored, rather thick, felted-submembranous, skirt-like ring, [17] and A. virosiformis , which has elongated spores that are 3.9–4.7 by 11.7–13.4 μm. [18] Neither A. elliptosperma nor A. magnivelaris typically turn yellow with the application of KOH; [19] [20] the KOH reaction of A. virosiformis has not been reported. [21]
Leucoagaricus leucothites is another all-white mushroom with an annulus, free gills, and white spore print, but it lacks a volva and has thick-walled dextrinoid (staining red-brown in Melzer's reagent) egg-shaped spores with a pore. [22] [23] A. bisporigera may also be confused with the larger edible species Agaricus silvicola , the "horse-mushroom". Like many white amanitas, young fruit bodies of A. bisporigera, still enveloped in the universal veil, can be confused with puffball species, but a longitudinal cut of the fruit body reveals internal structures in the Amanita that are absent in puffballs. [24] In 2006, seven members of the Hmong community living in Minnesota were poisoned with A. bisporigera because they had confused it with edible paddy straw mushrooms ( Volvariella volvacea ) that grow in Southeast Asia. [25]
| ||||||||||||||||||||||||||||||||||||
Relationships of Amanita bisporigera and related species based on ITS sequence data. The A. virosa specimen was collected from Japan, A. bisporigera from the US, and the other species from China. [26] |
Amanita bisporigera was first described scientifically in 1906 by American botanist George Francis Atkinson in a publication by Cornell University colleague Charles E. Lewis. The type locality was Ithaca, New York, where several collections were made. [8] In his 1941 monograph of world Amanita species, Édouard-Jean Gilbert transferred the species to his new genus Amanitina, [27] but this genus is now considered synonymous with Amanita. [28] In 1944, William Murrill described the species Amanita vernella, collected from Gainesville, Florida; [29] that species is now thought to be synonymous with A. bisporigera after a 1979 examination of its type material revealed basidia that were mostly 2-spored. [2] [30] Amanita phalloides var. striatula, a poorly known taxon originally described from the United States in 1902 by Charles Horton Peck, [31] is considered by Amanita authority Rodham Tulloss to be synonymous with A. bisporigera. [2] Vernacular names for the mushroom include "destroying angel", "deadly amanita", "white death cap", "angel of death" [4] and "eastern North American destroying angel". [6]
Amanita bisporigera belongs to section Phalloideae of the genus Amanita, which contains some of the deadliest Amanita species, including A. phalloides and A. virosa. This classification has been upheld with phylogenetic analyses, which demonstrate that the toxin-producing members of section Phalloideae form a clade—that is, they derive from a common ancestor. [32] [33] In 2005, Zhang and colleagues performed a phylogenetic analysis based on the internal transcribed spacer (ITS) sequences of several white-bodied toxic Amanita species, most of which are found in Asia. Their results support a clade containing A. bisporigera, A. subjunquillea var. alba, A. exitialis, and A. virosa. The Guangzhou destroying angel (Amanita exitialis) has two-spored basidia, like A. bisporigera. [26]
Like most other Amanita species, A. bisporigera is thought to form mycorrhizal relationships with trees. [7] This is a mutually beneficial relationship where the hyphae of the fungus grow around the roots of trees, enabling the fungus to receive moisture, protection and nutritive byproducts of the tree, and giving the tree greater access to soil nutrients. [34] Fruit bodies of Amanita bisporigera are found on the ground growing either solitarily, scattered, or in groups in mixed coniferous and deciduous forests; [5] they tend to appear during summer and early fall. [16] The fruit bodies are commonly found near oak, but have been reported in birch-aspen areas in the west. It is most commonly found in eastern North America, and rare in western North America. It is widely distributed in Canada, [4] and its range extends south to Mexico. [35] The species has also been found in Colombia, where it may have been introduced from trees exported for use in pine plantations. [6]
A. bisporigera is considered the most toxic North American Amanita mushroom, with little variation in toxin content between different fruit bodies. [36] [37] Three subtypes of amatoxin have been described: α-, β, and γ-amanitin. The principal amatoxin, α-amanitin, is readily absorbed across the intestine, and 60% of the absorbed toxin is excreted into bile and undergoes enterohepatic circulation; the kidneys clear the remaining 40%. The toxin inhibits the enzyme RNA polymerase II, thereby interfering with DNA transcription, which suppresses RNA production and protein synthesis. This causes cellular necrosis, especially in cells which are initially exposed and have rapid rates of protein synthesis. This process results in severe acute liver dysfunction and, ultimately, liver failure. [38] Amatoxins are not broken down by boiling, freezing, or drying. [39] [40] Roughly 0.2 to 0.4 milligrams of α-amanitin is present in 1 gram of A. bisporigera; the lethal dose in humans is less than 0.1 mg/kg body weight. [38] One mature fruit body can contain 10–12 mg of α-amanitin, enough for a lethal dose. [12] The α-amanitin concentration in the spores is about 17% that of the fruit body tissues. [41] A. bisporigera also contains the phallotoxin phallacidin, structurally related to the amatoxins but considered less poisonous because of poor absorption. [12] Poisonings (from similar white amanitas) have also been reported in domestic animals, including dogs, cats, and cows. [42]
The first reported poisonings resulting in death from the consumption of A. bisporigera were from near San Antonio, Mexico, in 1957, where a rancher, his wife, and three children consumed the fungus; only the man survived. [43] Amanita poisoning is characterized by the following distinct stages: [44] the incubation stage is an asymptomatic period which ranges from 6 to 12 hours after ingestion. In the gastrointestinal stage, about 6 to 16 hours after ingestion, there is onset of abdominal pain, explosive vomiting, and diarrhea for up to 24 hours, which may lead to dehydration, severe electrolyte imbalances, and shock. These early symptoms may be related to other toxins such as phalloidin. In the cytotoxic stage, 24 to 48 hours after ingestion, clinical and biochemical signs of liver damage are observed, but the patient is typically free of gastrointestinal symptoms. The signs of liver dysfunction such as jaundice, hypoglycemia, acidosis, and hemorrhage appear. Later, there is an increase in the levels of prothrombin and blood levels of ammonia, and the signs of hepatic encephalopathy and/or kidney failure appear. The risk factors for mortality that have been reported are age younger than 10 years, short latency period between ingestion and onset of symptoms, severe coagulopathy (blood clotting disorder), severe hyperbilirubinemia (jaundice), and rising serum creatinine levels. [38]
Amanita phalloides, commonly known as the death cap, is a deadly poisonous basidiomycete fungus, one of many in the genus Amanita. Widely distributed across Europe, but introduced to other parts of the world since the late twentieth century, A. phalloides forms ectomycorrhizas with various broadleaved trees. In some cases, the death cap has been introduced to new regions with the cultivation of non-native species of oak, chestnut, and pine. The large fruiting bodies (mushrooms) appear in summer and autumn; the caps are generally greenish in colour with a white stipe and gills. The cap colour is variable, including white forms, and is thus not a reliable identifier.
The name destroying angel applies to several similar, closely related species of deadly all-white mushrooms in the genus Amanita. They are Amanita virosa in Europe and A. bisporigera and A. ocreata in eastern and western North America, respectively. Another European species of Amanita referred to as the destroying angel, Amanita verna—also referred to as the "Fool's mushroom"—was first described in France in 1780.
The genus Amanita contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. The genus is responsible for approximately 95% of fatalities resulting from mushroom poisoning, with the death cap accounting for about 50% on its own. The most potent toxin present in these mushrooms is α-Amanitin.
Amanita virosa is a species of fungus in the class Agaricomycetes. In the UK, it has the recommended English name of destroying angel and is known internationally as the European destroying angel. Basidiocarps are agaricoid (mushroom-shaped) and pure white with a ring on the stem and a sack-like volva at the base. The species is deadly poisonous. It occurs in Europe and northern Asia. Amanita virosa was formerly reported from North America, but research has shown that similar-looking American species, including Amanita bisporigera and A. ocreata, are distinct.
Amatoxin is the collective name of a subgroup of at least nine related toxic compounds found in three genera of poisonous mushrooms and one species of the genus Pholiotina. Amatoxins are very potent, as little as half a mushroom cap can cause severe liver injury if swallowed.
Amanita verna, commonly known as the fool's mushroom or the spring destroying angel, is a deadly poisonous basidiomycete fungus, one of many in the genus Amanita. Occurring in Europe in spring, A. verna associates with various deciduous and coniferous trees. The caps, stipes and gills are all white in colour.
Galerina marginata, known colloquially as funeral bell, deadly skullcap, autumn skullcap or deadly galerina, is a species of extremely poisonous mushroom-forming fungus in the family Hymenogastraceae of the order Agaricales. It contains the same deadly amatoxins found in the death cap. Ingestion in toxic amounts causes severe liver damage with vomiting, diarrhea, hypothermia, and eventual death if not treated rapidly. About ten poisonings have been attributed to the species now grouped as G. marginata over the last century.
Amanita ocreata, commonly known as the death angel, destroying angel, angel of death or more precisely western North American destroying angel, is a deadly poisonous basidiomycete fungus, one of many in the genus Amanita. The large fruiting bodies generally appear in spring; the cap may be white or ochre and often develops a brownish centre, while the stipe, ring, gill and volva are all white. A. ocreata resembles several edible species commonly consumed by humans, increasing the risk of accidental poisoning. Mature fruiting bodies can be confused with the edible A. velosa, A. lanei or Volvopluteus gloiocephalus, while immature specimens may be difficult to distinguish from edible Agaricus mushrooms or puffballs.
Amanita brunnescens, also known as the brown American star-footed amanita or cleft-footed amanita is a native North American mushroom of the large genus Amanita. It differs from A. phalloides by its fragile volva and tendency to bruise brown.
Amanita arocheae, also known as the Latin American death cap, is a mushroom of the large genus Amanita, which occurs in Colombia, Central America and South America. Deadly poisonous, it is a member of section Phalloideae and related to the death cap, A. phalloides.
Amanita vaginata, commonly known as the grisette or the grisette amanita, is an edible mushroom in the fungus family Amanitaceae. The cap is gray or brownish, 5 to 10 centimetres in diameter, and has furrows around the edge that duplicate the gill pattern underneath. Unlike many other Amanita mushrooms, A. vaginata lacks a ring on the stem.
Galerina sulciceps is a dangerously toxic species of fungus in the family Strophariaceae, of the order Agaricales. It is distributed in tropical Indonesia and India, but has reportedly been found fruiting in European greenhouses on occasion. More toxic than the deathcap, G. sulciceps has been shown to contain the toxins alpha- (α-), beta- (β-) and gamma- (γ-) amanitin; a series of poisonings in Indonesia in the 1930s resulted in 14 deaths from the consumption of this species. It has a typical "little brown mushroom" appearance, with few obvious external characteristics to help distinguish it from many other similar nondescript brown species. The fruit bodies of the fungus are tawny to ochre, deepening to reddish-brown at the base of the stem. The gills are well-separated, and there is no ring present on the stem.
Amanita onusta, commonly known as the loaded Lepidella, the gunpowder Lepidella or the gunpowder amanita, is a species of fungus in the mushroom family Amanitaceae. It is characterized by its small to medium-sized fruit bodies that have white to pale gray caps crowded with roughly conical, pyramidal, or irregular gray warts. The stipe is whitish-gray with woolly or wart-like veil remnants, and at the base is a spindle- or turnip-shaped base that is rooted somewhat deeply in the soil.
Amanita aestivalis, commonly known as the white American star-footed amanita, is a species of fungus in the mushroom family Amanitaceae. The cap of the white fruit body is 5 to 8.5 centimetres in diameter. It sits atop a stem that is 8.5 to 16 cm long. The entire fruit body will slowly stain a reddish-brown color in response to bruising. A. aestivalis may be a synonym for A. brunnescens, and may be confused with several other white-bodied amanitas. The fungus is distributed in eastern North America.
Amanita exitialis, also known as the Guangzhou destroying angel, is a mushroom of the large genus Amanita. It is distributed in eastern Asia, and probably also in India where it has been misidentified as A. verna. Deadly poisonous, it is a member of section Phalloideae and related to the death cap A. phalloides. The fruit bodies (mushrooms) are white, small to medium-sized with caps up to 7 cm (2.8 in) in diameter, a somewhat friable ring and a firm volva. Unlike most agaric mushrooms which typically have four-spored basidia, the basidia of A. exitialis are almost entirely two-spored. Eight people were fatally poisoned in China after consuming the mushroom in 2000, and another 20 have been fatally poisoned since that incident. Molecular analysis shows that the species has a close phylogenetic relationship with three other toxic white Amanitas: A. subjunquillea var. alba, A. virosa and A. bisporigera.
Amanita australis is a species of fungus in the family Amanitaceae. It produces small- to medium-sized fruit bodies, with brown caps up to 9 centimetres in diameter covered with pyramidal warts. The gills on the underside of the cap are white, closely crowded together, and free from attachment to the stem. The stem, up to 9 cm long, has a ring and a bulbous base. The mushroom may be confused with another endemic New Zealand species, A. nothofagi, but can be distinguished by differences in microscopic characteristics.
Amanita nothofagi is a species of fungus in the family Amanitaceae. Endemic to New Zealand, the species was first described by mycologist Greta Stevenson in 1962. The fruit bodies have dark brown caps that are up to 13 cm (5.1 in) in diameter and covered with patches of soft greyish-brown scales or warts. The gills underneath the cap are crowded together, free from attachment to the stem, and white, becoming tinged with yellow in age. The stem of the mushroom is 4–14 cm (1.6–5.5 in) long by 0.5–2.5 cm (0.2–1.0 in) thick, and has a ring. The spore print is white, and individual spores are spherical to ellipsoid, measuring 7.5–9 by 7.5–9 micrometres. The mushroom may be confused with another New Zealand species, A. australis, but can be distinguished by certain characteristics. Amanita nothofagi is a mycorrhizal species, and grows in association with native New Zealand trees such as Southern Beech.
Saproamanita thiersii, commonly called Thiers' lepidella, is a North American saprotrophic basidiomycete fungus in the genus Saproamanita. It is a white, small mushroom. Its cap is convex, measuring 3.5–10 centimetres across, and the stipe is 8–20 cm (3–8 in) long. The spore print is white.
Amanita fuliginea, commonly known as the east Asian brown death cap, is a species of deadly poisonous mushroom in the family Amanitaceae. The fruit bodies have convex, dark gray to blackish caps measuring 3–6 cm (1.2–2.4 in) in diameter. The gills, largely free from attachment to the stipe, are white and have short gills (lamellulae) interspersed. The spores are roughly spherical, amyloid, and typically measure 8–11 by 7–9.5 μm. The species was described as new to science by Japanese mycologist Tsuguo Hongo in 1953. A. fuliginea is classified in Amanita section Phalloideae, which contains the infamous destroying angel.
Amanita hygroscopia, also known as the pink-gilled destroying angel is a deadly poisonous fungus, one of many in the genus Amanita.