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Agaricus

Agaricus is a genus of mushroom-forming fungi containing both edible and poisonous species, with over 400 members worldwide and possibly again as many disputed or newly discovered species. The genus includes the common ("button") mushroom (A. bisporus) and the field mushroom (A. campestris), the dominant cultivated mushrooms of the West.

Taxonomy
Several origins of genus name Agaricus have been proposed. It possibly originates from ancient Sarmatia Europaea, where people Agari, promontory Agarum and a river Agarus were known (all located on the northern shore of Sea of Azov, probably, near modern Berdiansk in Ukraine). The alternate name for the genus, Psalliota, derived from the Greek psalion/ψάλιον, "ring", was first published by Fries (1821) as trib. Psalliota. The type is Agaricus campestris (widely accepted, except by Earle, who proposed A. cretaceus). Paul Kummer (not Quélet, who merely excluded Stropharia) was the first to elevate the tribe to a genus. Psalliota was the tribe containing the type of Agaricus, so when separated, it should have caused the rest of the genus to be renamed, but this is not what happened. Phylogeny The use of phylogenetic analysis to determine evolutionary relationships amongst Agaricus species has increased the understanding of this taxonomically difficult genus, although much work remains to be done to fully delineate infrageneric relationships. Prior to these analyses, the genus Agaricus, as circumscribed by Rolf Singer, was divided into 42 species grouped into five sections based on reactions of mushroom tissue to air or various chemical reagents, as well as subtle differences in mushroom morphology. Restriction fragment length polymorphism analysis demonstrated this classification scheme needed revision. ==Subdivisions==
Subdivisions
As of 2018, this genus is divided into 6 subgenera and more than 20 sections: are found in a variety of often rather arid habitats, and typically have a smooth white to scaly light brown cap. The flesh, when bruised, usually turns distinctly yellow to pinkish in particular on the cap, while the end of the stalk may remain white; a marked yellow stain is caused by applying KOH. Their sweetish smell of almond extract or marzipan due to benzaldehyde and derived compounds distinguishes them from the section Xanthodermatei, as does a bright dark-orange to brownish-red coloration in Schäffer's test. Many members of this subgenus are highly regarded as food, and even medically beneficial, but at least some are known to accumulate cadmium and other highly toxic chemicals from the environment, and may not always be safe to eat. Peck 1873'' Fr. 1836/1838'' 'Subgenus Minores''''' :A group of buff-white to reddish-brown species. Often delicate and slender, the typical members of this subgenus do not resemble the larger Agaricus species at a casual glance, but have the same telltale chocolate-brown gills at spore maturity. Their flesh has a barely noticeable to pronounced sweetish smell, typically almond-like, turns yellowish to brownish-red when cut or bruised at least in the lower stalk, yellow to orange with KOH, and orange to red in Schäffer's test. Species such as A. aridicola (formerly known as Gyrophragmium dunalii), A. colpeteii, A. columellatus (formerly Araneosa columellata), A. diminutivus, A. dulcidulus, A. lamelliperditus, A. luteomaculatus, A. porphyrizon, A. semotus and A. xantholepis are included here, but delimitation to and indeed distinctness from subgenus Flavoagaricus is a long-standing controversy. Unlike these however, subgenus Minores contains no choice edible species, and may even include some slightly poisonous ones; most are simply too small to make collecting them for food worthwhile, and their edibility is unknown. • 'Section Leucocarpi''''' :Includes A. leucocarpus. • 'Section Minores''''' :Includes A. comtulus and A. huijsmanii. • 'Section Pantropicales''''' :Includes A. candidolutescens and an undescribed relative. • 'Section Duploannulatae''' (also known as section Bivelares or Hortenses'') :Traditionally often included in section Agaricus as subsection Bitorques, it seems to belong to a much younger radiation. It unites robust species, usually with a thick, almost fleshy ring, which inhabit diverse but often nutrient-rich locations. Some are well-known edibles; as they are frequently found along roads and in similar polluted places, they may not be safe to eat if collected from the wild. Their flesh is rather firm, white, with no characteristic smell, in some species turning markedly reddish when bruised or cut (though this may soon fade again), and generally changing color barely if at all after application of KOH or Schäffer's test. Based on DNA analysis of ITS1, ITS2, and 5.8S sequences, the studied species of this section could be divided into six distinct clades, four of which correspond to well-known species from the temperate Northern Hemisphere: A. bisporus, A. bitorquis (and the doubtfully distinct A. edulis), A. cupressicola and A. vaporarius. The other two clades comprise the A. devoniensis (including A. subperonatus) and A. subfloccosus (including A. agrinferus) species complexes. Additional members of this section not included in that study are A. cappellianus, A. cupressophilus, A. subsubensis, A. taeniatus, A. tlaxcalensis, and at least one undescribed species. The section forms a single clade based on analysis of ITS1+2. They are either bright white all over, or have a cap densely flecked with brownish scales or tufts of fibers. The ring is usually large but thin and veil-like. Most inhabit woodland, and in general they have a more or less pronounced unpleasant smell of phenolic compounds such as hydroquinone. As food, they should all be avoided, because even though they are occasionally reported to be eaten without ill effect, the chemicals they contain give them a acrid, metallic taste, especially when cooked, and are liable to cause severe gastrointestinal upset. Their flesh at least in the lower stalk turns pale yellow to intensely reddish-ochre when bruised or cut; more characteristic however is the a bright yellow reaction with KOH while Schäffer's test is negative. Apart from A. xanthodermus, the core group of this section contains species such as A. atrodiscus, A. californicus, A. endoxanthus and the doubtfully distinct A. rotalis, A. fuscopunctatus, A. iodosmus, A. laskibarii, A. microvolvatulus, A. menieri, A. moelleri, A. murinocephalus, A. parvitigrinus, A. placomyces, A. pocillator, A. pseudopratensis, A. tibetensis, A. tollocanensis, A. tytthocarpus, A. xanthodermulus, A. xanthosarcus, as well as at least 4 undescribed species, and possibly A. cervinifolius and the doubtfully distinct A. infidus. Whether such species as A. bisporiticus, A. nigrogracilis and A. pilatianus are more closely related to the mostly Eurasian core group, or to the more basal lineage here separated as section Hondenses, requires clarification. ''(F.H.Møller & Jul.Schäff. 1938) Hlaváček 1949 ex Pilát/Singer 1951 'Subgenus Spissicaules''''' :The flesh of members of this subgenus tends to turn more or less pronouncedly yellowish in the lower stalk, where the skin is often rough and scaly, and reddish in the cap. They typically resemble the darker members of subgenus Flavoagaricus, with a sweet smell and mild taste; like that subgenus, Spissicaules belongs to the smaller of the two main groups of the genus, but they form entirely different branch therein. While some species are held to be edible, others are considered unappetizing or even slightly poisonous. Also includes A. lanipes and A. maskae, which probably belong to section Rarolentes or Spissicaules, and possibly also A. bresadolanus and its doubtfully distinct relatives A. radicatus/romagnesii. • 'Section Amoeni''''' :Includes A. amoenus and A. gratolens. • 'Section Rarolentes''''' :Includes A. albosquamosus and A. leucolepidotus. • 'Section Spissicaules''''' (Hainem.) Kerrigan :Includes species such as A. leucotrichus/litoralis (of which A. spissicaulis is a synonym, but see also Geml et al. 2004 but since then, molecular phylogenetic studies have revalidated several disputed species, as well as resolved some species complexes, and aided in discovery and description of a wide range of mostly tropical species that were formerly unknown to science. As of 2020, the genus is believed to contain no fewer than 400 species, and possibly many more. The medicinal mushroom known in Japan as Echigoshirayukidake was initially also thought to be an Agaricus, either a subspecies of Agaricus "blazei" (i.e. A. subrufescens), or a new species. It was eventually identified as sclerotium of the crust-forming bark fungus Ceraceomyces tessulatus, which is not particularly closely related to Agaricus. Several secotioid (puffball-like) fungi have in recent times be recognized as highly aberrant members of Agaricus, and are now included here. These typically inhabit deserts where few fungiand even fewer of the familiar cap-and-stalk mushroom shapegrow. Another desert species, A. zelleri, was erroneously placed in the present genus and is now known as Gyrophragmium californicum. In addition, the scientific names Agaricus and – even more so – Psalliota were historically often used as a "wastebasket taxon" for any and all similar mushrooms, regardless of their actual relationships. Species either confirmed or suspected to belong into this genus include: • Agaricus abramsii •  Agaricus abruptibulbus – abruptly-bulbous agaricus, flat-bulb mushroom (disputed) •  Agaricus aestivalisAgaricus agrinferus (disputed) • Agaricus alabamensis •  Agaricus albolutescens (disputed) • Agaricus alligator •  Agaricus amicosusAgaricus angusticystidiatusAgaricus annaeAgaricus arorae •  Agaricus arvensis – horse mushroom •  Agaricus augustus – the prince •  Agaricus aurantioviolaceusAgaricus benesii •  Agaricus bernardii – salt-loving mushroom •  Agaricus bisporus – cultivated/button/portobello mushroom (includes A.brunnescens) •  Agaricus bitorquis – pavement mushroom, banded agaric • Agaricus braendleiAgaricus bresadolanus •  Agaricus brunneofibrillosus (formerly in A.fuscofibrillosus) •  Agaricus californicus – California agaricus •  Agaricus campestris – field/meadow mushroom • Agaricus columellatus (formerly in Araneosa) •  Agaricus cupreobrunneus – brown field mushroom •  Agaricus deserticola G.Moreno, Esqueda & Lizárraga (2010) – gasteroid agaricus (formerly in Longula) •  Agaricus dulcidulus – rosy wood mushroom (sometimes in A.semotus) •  Agaricus excellens (disputed) • Agaricus freirei •  Agaricus hondensis – felt-ringed agaricus •  Agaricus impudicus – tufted wood mushroom • Agaricus inapertus (formerly in Endoptychum) • Agaricus juliusAgaricus lanatonigerAgaricus langei (= A.fuscofibrillosus) •  Agaricus lilaceps – giant cypress agaricus • Agaricus litoralis – coastal mushroom (includes A.spissicaulis) •  Agaricus macrosporus (disputed) •   Agaricus moelleri – inky/dark-scaled mushroom (formerly in A.placomyces, includes A.meleagris) • Agaricus nebularum •  Agaricus pattersoniae •  Agaricus perobscurus – American princess •  Agaricus phaeolepidotus •  Agaricus pilatianus •  Agaricus placomyces (includes A.praeclaresquamosus) •  Agaricus pocillator •  Agaricus santacatalinensis •  Agaricus semotus •  Agaricus silvaticus – scaly/blushing wood mushroom, pinewood mushroom (= A.sylvaticus, includes A.haemorrhoidarius) •  Agaricus silvicola – wood mushroom (= A.sylvicola) •  Agaricus subrufescens (includes A.rufotegulis, often confused with A.blazei and A.brasiliensis) – almond mushroom, royal sun agaricus, and various fanciful names •  Agaricus subrutilescens – wine-colored agaricus • Agaricus taeniatus •  Agaricus urinascens •  Agaricus xanthodermus – yellow-staining mushroom == Description ==
Description
Members of Agaricus are characterized by having a fleshy cap or pileus, from the underside of which grow a number of radiating plates or gills, on which are produced the naked spores. They are distinguished from other members of their family, Agaricaceae, by their chocolate-brown spores. Members of Agaricus also have a stem or stipe, which elevates it above the object on which the mushroom grows, or substrate, and a partial veil, which protects the developing gills and later forms a ring or annulus on the stalk. == Toxicity ==
Toxicity
Amanita is often mistaken for edible Agaricus, with fatal results A notable group of poisonous Agaricus is the clade around the yellow-staining mushroom, A. xanthodermus. One species reported from Africa, A. aurantioviolaceus, is reportedly deadly poisonous. The species most often sold as such quack cures is A. subrufescens, which is often referred to by the erroneous name "Agaricus Blazei" and advertised by fanciful trade names such as "God's mushroom" or "mushroom of life", but can cause allergic reactions and even liver damage if consumed in excessive amounts. == Uses ==
Uses
The genus contains the most widely consumed and best-known mushroom today, A. bisporus. Other well-known and highly regarded species include A. arvensis, A. augustus, and A. campestris. Agaricus porphyrocephalus is a choice edible when young, while A. subrufescens may be edible in some cases. Many other species are of culinary value, especially within sections Agaricus, Arvense, Duploannulatae and Sanguinolenti. ==References==
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