Agaricomycetes Temporal range: | |
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Amanita muscaria (Agaricales) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Subdivision: | Agaricomycotina |
Class: | Agaricomycetes Doweld (2001) [1] |
Subclasses/orders | |
incertae sedis (no subclass)
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The Agaricomycetes are a class of fungi in the division Basidiomycota. The taxon is roughly identical to that defined for the Homobasidiomycetes (alternatively called holobasidiomycetes) by Hibbett & Thorn, [2] with the inclusion of Auriculariales and Sebacinales. It includes not only mushroom-forming fungi, but also most species placed in the deprecated taxa Gasteromycetes and Homobasidiomycetes. [3] Within the subdivision Agaricomycotina, which already excludes the smut and rust fungi, the Agaricomycetes can be further defined by the exclusion of the classes Tremellomycetes and Dacrymycetes, which are generally considered to be jelly fungi. However, a few former "jelly fungi", such as Auricularia , are classified in the Agaricomycetes. According to a 2008 estimate, Agaricomycetes include 17 orders, 100 families, 1147 genera, and about 21000 species. [4] Modern molecular phylogenetic analyses have been since used to help define several new orders in the Agaricomycetes: Amylocorticiales, Jaapiales, [5] Stereopsidales, [6] and Lepidostromatales. [7]
Although morphology of the mushroom or fruit body (basidiocarp) was the basis of early classification of the Agaricomycetes, [8] this is no longer the case. As an example, the distinction between the Gasteromycetes (including puffballs) and Agaricomycetes (most other agaric mushrooms) is no longer recognized as a natural one—various puffball species have apparently evolved independently from agaricomycete fungi. However, most mushroom guide books still group the puffballs or gasteroid forms separate from other mushrooms because the older Friesian classification is still convenient for categorizing fruit body forms. Similarly, modern classifications divide the gasteroid order Lycoperdales between Agaricales and Phallales.
All members of the class produce basidiocarps which range in size from tiny cups a few millimeters across to a giant polypore ( Phellinus ellipsoideus ) greater than several meters across and weigh up to 500 kilograms (1,100 lb). [9] The group also includes what are arguably the largest and oldest individual organisms on earth: the mycelium of one individual Armillaria gallica has been estimated to extend over 15 hectares (37 acres) with a mass of 10,000 kilograms (22,000 pounds) and an age of 1,500 years. [10]
Agaricomycetes also have antibacterial properties. Agaricomycetes can help in research in treating bacteria.[ clarification needed ] [11]
Nearly all species are terrestrial (a few are aquatic), occurring in a wide range of environments where most function as decayers, especially of wood. However, some species are pathogenic or parasitic, and yet others are symbiotic (i.e., mutualistic), these including the important ectomycorrhizal symbionts of forest trees. General discussions on the forms and life cycles of these fungi are developed in the article on mushrooms, in the treatments of the various orders (links in table at right), and in individual species accounts.
A study of 5,284 species with a backbone phylogeny based on 104 genomes [12] has suggested the following dates of evolution:
Agaricomycetidae ~ 185 million years ago ( 174 million years ago– 192 million years ago)
Cantharellales 184 million years ago ( 144 million years ago– 261 million years ago)
Agaricales 173 million years ago ( 160 million years ago- 182 million years ago)
Hymenochaetales 167 million years ago ( 130 million years ago– 180 million years ago)
Boletales 142 million years ago ( 133 million years ago– 153 million years ago)
The fruit bodies of Agaricomycetes are extremely rare in the fossil record, and the class does not yet pre-date the Early Cretaceous (146–100 Ma). [13] The oldest Agaricomycetes fossil, dating from the lower Cretaceous (130–125 Ma) is Quatsinoporites . It is a fragment of a poroid fruit body with features that suggest it could be a member of the family Hymenochaetaceae. [14] Based on molecular clock analysis, the Agaricomycetes are estimated to be about 290 million years old. [15]
Modern molecular phylogenetics suggest the following relationships: [16]
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There are many genera in the Agaricomycetes that have not been classified in any order or family. These include:
Basidiomycota is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basidiomycota includes these groups: agarics, puffballs, stinkhorns, bracket fungi, other polypores, jelly fungi, boletes, chanterelles, earth stars, smuts, bunts, rusts, mirror yeasts, and Cryptococcus, the human pathogenic yeast.
The Polyporales are an order of about 1,800 species of fungi in the division Basidiomycota. The order includes some polypores as well as many corticioid fungi and a few agarics. Many species within the order are saprotrophic, most of them wood-rotters. Some genera, such as Ganoderma and Fomes, contain species that attack living tissues and then continue to degrade the wood of their dead hosts. Those of economic importance include several important pathogens of trees and a few species that cause damage by rotting structural timber. Some of the Polyporales are commercially cultivated and marketed for use as food items or in traditional Chinese medicine.
The Agaricales are an order of fungi in the division Basidiomycota. As originally conceived, the order contained all the agarics, but subsequent research has shown that not all agarics are closely related and some belong in other orders, such as the Russulales and Boletales. Conversely, DNA research has also shown that many non-agarics, including some of the clavarioid fungi and gasteroid fungi belong within the Agaricales. The order has 46 extant families, more than 400 genera, and over 25,000 described species, along with six extinct genera known only from the fossil record. Species in the Agaricales range from the familiar Agaricus bisporus and the deadly Amanita virosa to the coral-like Clavaria zollingeri and bracket-like Fistulina hepatica.
The Hymenochaetales are an order of fungi in the class Agaricomycetes. The order in its current sense is based on molecular research and not on any unifying morphological characteristics. According to one 2008 estimate, the Hymenochaetales contain around 600 species worldwide, mostly corticioid fungi and poroid fungi, but also including several clavarioid fungi and agarics. Species of economic importance include wood decay fungi in the genera Phellinus and Inonotus sensu lato, some of which may cause losses in forestry. Therapeutic properties are claimed for Inonotus obliquus ("chaga") and Phellinus linteus, both of which are now commercially marketed.
Neolentinus is a genus of wood-decaying agarics with tough fruit bodies composed of dimitic tissue, serrated lamella edges, and nonamyloid white binucleate basidiospores among other features. It was segregated from Lentinus in the broad taxonomic sense, hence the derivation of the name. Biologically Neolentinus species produce a brown rot type of decay of wood, whereas Lentinus causes a white rot. Molecular phylogenetic analysis shows that the two genera are unrelated. Neolentinus is phylogenetically allied to other brown rot genera such as Gloeophyllum, Heliocybe, and Veluticeps. A new order, the Gloeophyllales, has been described for these fungi. Heliocybe had been placed in synonymy but it differs phylogenetically and anatomically by the lack of clamp connections that all Neolentinus produce on their generative hyphae.
The Clavariaceae are a family of fungi in the order Agaricales. Originally the family contained most of the clavarioid fungi, but in its current sense is more restricted, albeit with a greater diversity of basidiocarp forms. Basidiocarps are variously clavarioid or agaricoid (mushroom-shaped), less commonly corticioid or hydnoid.
The Clavulinaceae are a family of fungi in the order Cantharellales. The family is not well defined, but currently comprises species of clavarioid fungi as well as some corticioid fungi. These species are nutritionally diverse, some being ectomycorrhizal, others wood-rotting saprotrophs, others lichenized, and yet others lichenicolous.
The Thelephorales are an order of fungi in the class Agaricomycetes. The order includes corticioid and hydnoid fungi, together with a few polypores and clavarioid species. Most fungi within the Thelephorales are ectomycorrhizal. None is of any great economic importance, though Sarcodon imbricatus is edible and commercially marketed, whilst several species have been used for craft dyeing.
The Tricholomataceae are a large family of fungi within the order Agaricales. Originally a classic "wastebasket taxon", the family included any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to e.g. the Amanitaceae, Lepiotaceae, Hygrophoraceae, Pluteaceae, or Entolomataceae.
The Sclerodermataceae are a family of fungi in the order Boletales, containing several genera of unusual fungi that little resemble boletes. Taxa, which include species commonly known as the ‘hard-skinned puffballs’, ‘earthballs’, or 'earthstars', are widespread in both temperate and tropical regions. The best known members include the earthball Scleroderma citrinum, the dye fungus Pisolithus tinctorius and the 'prettymouths' of the genus Calostoma.
Agaricomycotina is one of three subdivisions of the Basidiomycota, and represents all of the fungi which form macroscopic fruiting bodies. Agaricomycotina contains over 30,000 species, divided into three classes: Tremellomycetes, Dacrymycetes, and Agaricomycetes. Around 98% of the species are in the class Agaricomycetes, including all the agarics, bracket fungi, clavarioid fungi, corticioid fungi, and gasteroid fungi. Tremellomycetes contains many basidiomycete yeasts and some conspicuous jelly fungi. Dacrymycetes contains a further group of jelly fungi. These taxa are founded on molecular research, based on cladistic analysis of DNA sequences, and supersede earlier morphology-based classifications. Agaricomycotina contains nearly one third of all described species of fungi.
Gloeophyllum is a genus of fungus in the class Agaricomycetes. It is characterized by the production of leathery to corky tough, brown, shaggy-topped, revivable fruitbodies lacking a stipe and with a lamellate to daedaleoid or poroid fertile hymenial surfaces. The hyphal system is dimitic to trimitic. The genus is further characterized by the production of a brown rot of wood. Phylogenetically, it along with several other brown rot Basidiomycota, Neolentinus, Heliocybe, and Veluticeps form an order called the Gloeophyllales.
Fungi diverged from other life around 1.5 billion years ago, with the glomaleans branching from the "higher fungi" (dikaryans) at ~570 million years ago, according to DNA analysis. Fungi probably colonized the land during the Cambrian, over 500 million years ago,, and possibly 635 million years ago during the Ediacaran, but terrestrial fossils only become uncontroversial and common during the Devonian, 400 million years ago.
The Hydnaceae are a family of fungi in the order Cantharellales. Originally the family encompassed all species of fungi that produced basidiocarps having a hymenium consisting of slender, downward-hanging tapering extensions referred to as "spines" or "teeth", whether they were related or not. This artificial but often useful grouping is now more generally called the hydnoid or tooth fungi. In the strict, modern sense, the Hydnaceae are limited to the genus Hydnum and related genera, with basidiocarps having a toothed or poroid hymenium. Species in the family are ectomycorrhizal, forming a mutually beneficial relationship with the roots of trees and other plants. Hydnum repandum is an edible species, commercially collected in some countries and often marketed under the French name pied de mouton.
The Corticiaceae are a family of fungi in the order Corticiales. The family formerly included almost all the corticioid fungi, whether they were related or not, and as such was highly artificial. In its current sense, however, the name Corticiaceae is restricted to a comparatively small group of corticioid genera within the Corticiales.
The Phanerochaetaceae are a family of mostly crust fungi in the order Polyporales.
The corticioid fungi are a group of fungi in the Basidiomycota typically having effused, smooth basidiocarps that are formed on the undersides of dead tree trunks or branches. They are sometimes colloquially called crust fungi or patch fungi. Originally such fungi were referred to the genus Corticium and subsequently to the family Corticiaceae, but it is now known that all corticioid species are not necessarily closely related. The fact that they look similar is an example of convergent evolution. Since they are often studied as a group, it is convenient to retain the informal (non-taxonomic) name of "corticioid fungi" and this term is frequently used in research papers and other texts.
The gasteroid fungi are a group of fungi in the Basidiomycota. Species were formerly placed in the obsolete class Gasteromycetes Fr., or the equally obsolete order Gasteromycetales Rea, because they produce spores inside their basidiocarps rather than on an outer surface. However, the class is polyphyletic, as such species—which include puffballs, earthballs, earthstars, stinkhorns, bird's nest fungi, and false truffles—are not closely related to each other. Because they are often studied as a group, it has been convenient to retain the informal (non-taxonomic) name of "gasteroid fungi".
Amylocorticiales is an order of fungi in the class Agaricomycetes. The order was circumscribed in 2010 to contain mostly resupinate (crust-like) forms that have been referred to genera Anomoporia, Amyloathelia, Amylocorticiellum, Amylocorticium, Amyloxenasma, Anomoloma, Athelopsis, Ceraceomyces, Hypochniciellum, Leptosporomyces and Serpulomyces and the anomalous species, Athelia rolfsii, now classified in its own genus, Agroathelia.