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Agaricales

(Order)

Overview

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The fungal order Agaricales, also known as gilled mushrooms (for their distinctive gills), or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species,4] along with five extinct genera known only from the fossil record.[1][5] They range from the ubiquitous common mushroom to the deadly destroying angel and the hallucinogenic fly agaric to the bioluminescent jack-o-lantern mushroom.

History, classification and phylogeny

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In his three volumes of Systema Mycologicum published between 1821 and 1832, Elias Fries put almost all of the flesh y, gill-forming mushrooms in the genus Agaricus. He organized the large genus into "tribes", the names of many of which still exist as common genera of today. Fries later elevated several of these tribes to generic level, but later authors?including Gillet, Karsten, Kummer, Qu?let, and Staude?made most of the changes. Fries based his classification on macroscopic characters of the fruit bodies and color of the spore print. His system had been widely used as it had the advantage that many genera could be readily identified based on characters observable in the field. Fries's classification was later challenged when microscopic studies of basidiocarp structure, initiated by Fayod and Patouillard, demonstrated several of Fries's groupings were unnatural.[4] In more recent history, Rolf Singer's influential work The Agaricales in Modern Taxonomy, published in four editions spanning from 1951 to 1986, used bot h Fries's macroscopic characters and Fayod's microscopic characters to reorganize families and genera; his most recent classification included 230 genera within 18 families.[7] Singer treated three major groups within the Agaricales sensu lato: the Agaricales sensu stricto, Boletales, and Russulales. These groups are still accepted by modern treatments based on DNA analysis, as the euagarics clade, bolete clade, and russuloid clade.[8]

Molecular phylogenetics research has demonstrated that the euagarics clade is roughly equivalent to Singer's Agaricales sensu stricto.[9][10][11] A recent (2006) large-scale study by Brandon Matheny and colleagues used nucleic acid sequences representing six gene regions from 238 species in 146 genera to explore the phylogenetic grouping within the Agaricales. The analysis showed that most of the species tested could be grouped into six clades that were named the Agaricoid, Tricholomatoid, Marasmioid, Pluteoid, Hygrophoroid and Plicaturopsidoid clades.[6]

Some notable fungi with gill-like structures, such as chanterelles, have long been recognized as being substantially different from usual Agaricales. Interestingly, molecular studies are showing more groups of agarics as being more divergent than previously thought, such as the genera Russula and Lactarius belonging to a separate order Russulales, and other gilled fungi, including such species as Paxi llus involutus and Hygrophoropsis aurantiaca showing a closer affinity with Boletes in the order Boletales.

Also, some other quite distinctive fungi, the puffballs, and some clavaroid fungi, e.g. Typhula, and the Beefsteak fungus have been recently been shown to lie within the Agaricales.

The term agaric had traditionally referred to Agaricales, which were defined as exactly those fungi with gills. Given the discoveries described above, those two categories are not synonymous (although there is a very large overlap between the two groups).

Distribution and habitat

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Agarics are ubiquitous, being found across all continents. Most are terrestrial, their habitats including all types of woodland and grassland, varying largely from one genus to another. Agarics were long thought to be solely terrestrial, until the 2005 discovery of Psathyrella aquatica, the only gilled mushro om known to fruit underwater.[12]

Agaricals are known from five monotypic genera found fossilized in amber. The oldest records are from two Cretaceous age genera; the Albian age (approximately 100 Ma) Palaeoagaracites antiquus from Burmese amber and the slightly younger Turonian New Jersey Amber species Archaeomarasmius leggeti.[1] The three other species, Aureofungus yaniguaensis, Coprinites dominicana and Protomycena electra are known from single specimens found in the Dominican amber mines of Hispaniola.[5]

Cortinarius archeri

Characteristics

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Basidiocarps of the agarics are typically fleshy, with a stipe, often called a stem or stalk, a pileus (or cap) and lamellae (or gills), where basidiospores are produced. This is the stereotypical structure of a mushroom.

Life cycle

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The fungus fruit body is the spore-producing stage of the life cycle. Most fungi reproduce by spores and the fruit bodies are developed specifically for the production and dispersal of spores. The spores produced by fruit bodies are usually the result of sexual reproduction.[13]

The fruit body is the visible part of the growing fungus. It is supported by and d evelops from an extensive network of thread-like filaments called hyphae. Hyphae are often collectively termed the mycelium; the food-absorbing part of the fungus?as opposed to the spore-producing fruit body of the fungus?is called the vegetative mycelium. The individual hyphae that compose the mycelium absorb nutrients and water from the substratum in which they are growing. When the nutrient supply is adequte and envirnmental conditions are favorable, some fungi may grow in the same location for several years. Fungi cannot make their own food, namely carbohydrates, as can green plants. Some species are saprobic, obtaining nutrients from dead organic material, whereas others are parasitic on living plants or animals or even on other fungi. Many fungi, especially gilled mushroomes and boletes, have an extensive mycelium that lives in association with the roots of woody plants. This association, which is beneficial to both the fungus and host plant, is termed a mycorrhiza.[13]

When the environmental conditions are favorable and the mycelium is at the proper stage of development, one or more fruit bodies are produced by the fungus. The actual conditions necessary for fruit body formation and spore production are not clearly understood. Humidity, light, temperature, aeration, and nutrition are all factors thought to be important in fruit body formation. The genetic makeup and the general physiology of the fungus hyphae are also important in the initiation and formation of young fruit bodies and their development to a mature stage. The spores produced by a fruit body are released when it is mature. When they land in a suitable environment, the spores germinate and the hyphae grow to initiate the life cycle anew.[13]

Genera Incertae sedis

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There are several genera classified in the Agaricales that are poorly-known or have not been subjected to DNA analysis, and have not been assigned to a specific family (i.e., Incertae sedis with respect to familial placement). These include:

See also

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opolous1996_12-0" class="reference">[13]

The fruit body is the visible part of the growing fungus. It is supported by and develops from an extensive network of thread-like filaments called hyphae. Hyphae are often collectively termed the mycelium; the food-absorbing part of the fungus?as opposed to the spore-producing fruit body of the fungus?is called the vegetative mycelium. The individual hyphae that compose the mycelium absorb nutrients and water from the substratum in which they are growing. When the nutrient supply is adequte and envirnmental conditions are favorable, some fungi may grow in the same location for several years. Fungi cannot make their own food, namely carbohydrates, as can green plants. Some species are saprobic, obtaining nutrients from dead organic material, whereas others are parasitic on living plants or animals or even on other fungi. Many fungi, especially gilled mushroomes and boletes, have an extensive mycelium that lives in association with the roots of woody plants. This association, which is beneficial to both the fungus and host plant, i s termed a mycorrhiza.[13]

When the environmental conditions are favorable and the mycelium is at the proper stage of development, one or more fruit bodies are produced by the fungus. The actual conditions necessary for fruit body formation and spore production are not clearly understood. Humidity, light, temperature, aeration, and nutrition are all factors thought to be important in fruit body formation. The genetic makeup and the general physiology of the fungus hyphae are also important in the initiation and formation of young fruit bodies and their development to a mature stage. The spores produced by a fruit body are released when it is mature. When they land in a suitable environment, the spores germinate and the hyphae grow to initiate the life cycle anew.[13]

Genera Incerta e sedis

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There are several genera classified in the Agaricales that are poorly-known or have not been subjected to DNA analysis, and have not been assigned to a specific family (i.e., Incertae sedis with respect to familial placement). These include:

See also

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References

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  1. ^ a b c Poinar, GO, Buckley R. (2007). "Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber". Mycological Research 111 (4): 503?506. doi:10.1016/j.mycres.2007.02.004. PMID 17512712. 
  2. ^ Underwood LM. (1899). Moulds, mildews and mushrooms: a guide to the systematic study of the Fungi and Mycetozoa and their literature. New York, Ne w York: Henry Holt. p. 97. 
  3. ^ "Agaricales Underw. 1899". MycoBank. International Mycological Association. http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=90508. Retrieved 2010-12-30. 
  4. ^ a b Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CABI. p. 12. ISBN 0-85199-826-7. 
  5. ^ a b Hibbett DS, Binder M, Wang Z, Goldma n Y. (2003). "Another Fossil Agaric from Dominican Amber". Mycologia 95 (4): 685?687. doi:10.2307/3761943. JSTOR 3761943. PMID 21148976. (subscription required)
  6. ^ a b Matheny PB, Curtis JM, Hofstetter V, Aime MC, Moncalvo JM, Ge ZW, Slot JC, Ammirati JF, Baroni TJ, Bougher NL, Hughes KW, Lodge DJ, Kerrigan RW, Seidl MT, Aanen DK, DeNitis M, Daniele GM, Desjardin DE, Kropp BR, Norvell LL, Parker A, Vellinga EC, Vilgalys R, Hibbett DS. (2006). "Major clades of Agaricales: a multilocus phylogenetic overview" (PDF). Mycologia 98 (6): 982?95. doi :10.3852/mycologia.98.6.982. PMID 17486974. http://www.clarku.edu/faculty/dhibbett/Reprints%20PDFs/Mathenyetal_Agaricales_2006.pdf. 
  7. ^ Singer R. (1986). The Agaricales in Modern Taxonomy (4th ed.). Koenigstein K?nigstein im Taunus, Germany: Koeltz Scientific Books. ISBN 3-87429-254-1. 
  8. ^ Hibbett DH, Thorn RG.. "Basidiomycota: Homobasidiomycetes". In McLaughlin DJ, McLaughlin EG, Lemke PA. The Mycota. VIIB. Systematics and Evolution. Springer-Verlag. pp. 121?68. ISBN 978-3540580089. 
  9. ^ Hibbett DS, Pine EM, Langer E, Langer G, Donoghue MJ. (1997). "Evolution of gilled mushrooms and puffballs inferred from ribosomal DNA sequences". Proceedings of the National Academy of Sciences USA 94 (22): 12002?6. doi:10.1073/pnas.+94.22.12002. 
  10. ^ Moncalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R. (2000). "Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences" (PDF). Systematic Biology 49 (2): 278?305. doi:10.1093/sysbio/49.2.278. PMID 12118409. http://sysbio.oxfordjournals.o rg/content/49/2/278.full.pdf. 
  11. ^ Moncalvo JM, Vilgalys R, Redhead SA, Johnson JE, James TY, Catherine Aime M, Hofstetter V, Verduin SJ, Larsson E, Baroni TJ, Greg Thorn R, Jacobsson S, Cl?men?on H, Miller OK. (2002). "One hundred and seventeen clades of euagarics" (PDF). Molecular Phylogenetics and Evolution 23 (3): 357?400. doi:10.1016/S1055-7903(02)00027-1. PMID 12099793. http://www.umich.edu/~mycology/publications_assets/moncalvo.mpe.2002.pdf. 
  12. ^ Frank JL, Coffan RA, Southworth D. (20 10). "Aquatic gilled mushrooms: Psathyrella fruiting in the Rogue River in southern Oregon". Mycologia 102 (1): 93?107. doi:10.3852/07-190. PMID 20120233. 
  13. ^ a b c Alexopolous et al., pp. 508?43.

Cited texts

External links

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Taxonomy

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The Order Agaricales is further organized into finer groupings including:

Families

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Agaricaceae

The Agaricaceae are a family of basidiomycete fungi and includes the genus Agaricus, as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. The genus contains 85 genera and 1340 species. [more]

Amanitaceae

Amanitaceae are a family of fungi or mushrooms. The family, also commonly called the Amanita family, is in order Agaricales, gilled mushrooms. The family consists primarily of the genus Amanita, but also includes the genera Catatrama and Limacella. [more]

Bolbitiaceae

The Bolbitiaceae are a family of basidiomycete fungi. There are 17 genera and 171 species in the family. [more]

Broomeiaceae

[more]

Clavariaceae

The Clavariaceae are a family of fungi in the Agaricales order of mushrooms. The family contains 7 genera and 120 species. Collectively, they are commonly known as coral fungi due to their resemblance to aquatic coral, although other vernacular names including antler fungi, finger fungi, worm mold, and spaghetti mushroom are sometimes used for similar reasons. [more]

Coprinaceae

[more]

Coritinariaceae

[more]

Cortinariaceae

The Cortinariaceae are a large family of gilled mushrooms found worldwide, containing over 2100 species. The family takes its name from its largest genus, the varied species of the genus Cortinarius. Many genera formerly in the Corinariaceae have been placed in various other families, including Hymenogastraceae, Inocybaceae and Bolbitiaceae. [more]

Crepidotaceae

[more]

Entolomataceae

The Entolomataceae, also known as Rhodophyllaceae are a large family of pink spored terrestrial gilled mushrooms which includes the genera Entoloma, Rhodocybe, and Clitopilus. The family collectively contains over 1500 species, the large majority of which are in Entoloma. Genera formerly known as Leptonia and Nolanea, amongst others, have been subsumed into Entoloma. Mushrooms in the Entolomataceae typically grow in woodlands or grassy areas and have attached gills, differentiating them from the Pluteaceae which have free gills. [more]

Gigaspermaceae

Gigaspermaceae is a family of mosses in order Funariales. [more]

Hydnangiaceae

The Hydnangiaceae are a family of fungi in the order of mushrooms known as the Agaricales. Widespread in temperate and tropical regions throughout the world, the family contains approximately 30 species amongst 4 genera. Species in the Hydnangiaceae form ectomycorrhizal relationships with various species of trees in both coniferous and deciduous forests. [more]

Hygrophoraceae

The Hygrophoraceae are a family of fungi in the order Agaricales. Originally conceived as containing white-spored, thick-gilled agarics (gilled mushrooms), including Hygrophorus and Hygrocybe species (the waxcaps or waxy caps), DNA evidence has extended the limits of the family, so that it now contains not only agarics but also basidiolichens and corticioid fungi. Species are thus diverse and are variously ectomycorrhizal, lichenized, associated with mosses, or saprotrophic. The family contains 18 genera and over 400 species. None is of any great economic importance, though fruit bodies of some Hygrocybe and Hygrophorus species are considered edible and may be collected for sale in local markets. [more]

Lycoperdaceae

Lycoperdaceae is a family of approximately 150 fungi now known to lie in the Agaricales. Historically they were placed in their own order Lycoperdales. Members of the Lycoperdaceae family are known as the true puffballs. Unlike other types of fungi that hold spores in gills or teeth, puffballs contain the spores inside a layer of tougher outer skin. When a puffball reaches maturity, the tough skin will split open, allowing the billions of spores to be released. [more]

Marasmiaceae

The Marasmiaceae are a family of basidiomycete fungi which have white spores. They mostly have a tough stem and the capability of shrivelling up during a dry period and later recovering. The widely consumed edible fungus Lentinula edodes, the Shiitake mushroom, is a member of this family. According to a 2008 estimate, the family contains 54 genera and 1590 species. [more]

Nidulariaceae

The Nidulariaceae are a family of fungi in the order Nidulariales. Commonly known as the bird's nest fungi, their fruiting bodies resemble tiny egg-filled birds' nests. As they are saprobic, feeding on decomposing organic matter, they are often seen growing on decaying wood and in soils enriched with wood chips or bark mulch; they have a widespread distribution in most ecological regions. The five genera within the family, namely, Crucibulum, Cyathus, Mycocalia, Nidula, and Nidularia, are distinguished from each other by differences in morphology and peridiole structure; more recently, phylogenetic analysis and comparison of DNA sequences is guiding new decisions in the taxonomic organization of this family [more]

Physalacriaceae

[more]

Pleurotaceae

Pleurotaceae are a family of small to medium sized mushrooms which have white spores. The family contains 6 genera and 94 species. Members of Pleurotaceae can be mistaken for members of Omphalotaceae. Perhaps the best known member is the oyster mushroom, Pleurotus ostreatus. [more]

Pluteaceae

The Pluteaceae are a family of small to medium-sized mushrooms which have free gill attachment and pink spores. Members of Pluteaceae can be mistaken for members of Entolomatacae but can be distinguished by their angled spores and attached gills. The four genera in the Pluteaceae include the widely distributed Volvariella and Pluteus, the rare Chamaeota, and Volvopluteus, newly described in 2011 as a result of molecular analysis. The Dictionary of the Fungi (10th edition, 2008) estimates there are 364 species in the family. [more]

Psathyrellaceae

The Psathyrellaceae are a family of dark-spored agarics that generally have rather soft, fragile fruiting bodies, and are characterized by black or dark brown, rarely reddish, or even pastel colored spore prints. About 50% of the species produce fruiting bodies that dissolve into ink-like ooze when the spores are mature via self digestion (autodigestion). Prior to phylogenetic research based upon DNA comparisons, most of the species that autodigested were previously classified in another family called the Coprinaceae that contained all of the inky cap mushrooms. The reclassification took place because the type species of Coprinus, Coprinus comatus, and a few other species were found to belong to another family, the Agaricaceae. The former group of old Coprinus was split between two families, and the name "Coprinaceae" became a synonym of the Agaricaceae in its 21st century phylogenetic redefinition. Note that in the 19th and early 20th centuries the family name Agaricaceae had far broader application, while in the late 20th century it had a narrower application. The family name Psathyrellaceae is based upon the subfamily name Psathyrelloideae, that had been classified in the Coprinaceae. The type genus, Psathyrella consists of species that produce fruitbodies do not liquify via autodigestion. Currently Psathyrella is a polyphyletic genus that will be further fragmented and reclassified. Lacrymaria is another genus that does not autodigest its fruitbodies. It is characterized by rough basidiospores and lamellar edges that exude beads of clear liquid when in prime condition, hence the Latin reference, 'lacrym-" to crying (tears). [more]

Pterulaceae

The Pterulaceae are a family of fungi in the Agaricales order. According to a 2008 estimate, the family contains 99 species distributed among 12 genera. [more]

Schizophyllaceae

The Schizophyllaceae are a family of fungi in the Agaricales order. The family contains two genera and seven species. Species cause white rot in hardwoods. The most common member of the genus Schizophyllum is Schizophyllum commune, a widely distributed mushroom. It looks like a mini Oyster mushroom which is one-fifth the size. [more]

Strophariaceae

The Strophariaceae are a family of fungi in the order Agaricales. The family contains 18 genera and 1316 species. The species of Strophariaceae have a red-brown to dark brown spore print, while the spores themselves are smooth and have an apical germ pore. These agarics are also characterized by having a cutis-type pileipellis. Ecologically, all species in this group are saprotrophs, growing on various kinds of decaying organic matter. [more]

Tricholomataceae

The Tricholomataceae are a large family of mushrooms within the Agaricales. A classic "wastebasket taxon", the Tricholomataceae is inclusive of any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to the Amanitaceae, Lepiotaceae, Hygrophoraceae, Pluteaceae, or Entolomataceae. [more]

Tulostomataceae

The Agaricaceae are a family of basidiomycete fungi and includes the genus Agaricus, as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. The genus contains 85 genera and 1340 species. [more]

Typhulaceae

The Typhulaceae are a family of fungi in the Agaricales order. The family contains 6 genera and 229 species. [more]

At least 459 species and subspecies belong to the Family Typhulaceae.

More info about the Family Typhulaceae may be found here.

References

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  1. ^ a b c Poinar, GO, Buckley R. (2007). "Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber". Mycological Research 111 (4): 503?506. doi:10.1016/j.mycres.2007.02.004. PMID 17512712. 
  2. ^ Underwood LM. (1899). Moulds, mildews and mushrooms: a guide to the systematic study of the Fungi and Mycetozoa and their literature. New York, New York: Henry Holt. p.  97. 
  3. ^ "Agaricales Underw. 1899". MycoBank. International Mycological Association. http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=90508. Retrieved 2010-12-30. 
  4. ^ a b Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CABI. p. 12. ISBN 0-85199-826-7. 
  5. ^ a b Hibbett DS, Binder M, Wang Z, Goldman Y. (2003). "Anoth er Fossil Agaric from Dominican Amber". Mycologia 95 (4): 685?687. doi:10.2307/3761943. JSTOR 3761943. PMID 21148976. (subscription required)
  6. ^ a b Matheny PB, Curtis JM, Hofstetter V, Aime MC, Moncalvo JM, Ge ZW, Slot JC, Ammirati JF, Baroni TJ, Bougher NL, Hughes KW, Lodge DJ, Kerrigan RW, Seidl MT, Aanen DK, DeNitis M, Daniele GM, Desjardin DE, Kropp BR, Norvell LL, Parker A, Vellinga EC, Vilgalys R, Hibbett DS. (2006). "Major clades of Agaricales: a multilocus phylogenetic overview" (PDF). Mycologia 98 (6): 982?95. doi:10.3852/mycologia.98.6.982. PMID 17486974. http://www.clarku.edu/faculty/dhibbett/Reprints%20PDFs/Mathenyetal_Agaricales_2006.pdf. 
  7. ^ Singer R. (1986). The Agaricales in Modern Taxonomy (4th ed.). Koenigstein K?nigstein im Taunus, Germany: Koeltz Scientific Books. ISBN 3-87429-254-1. 
  8. ^ Hibbett DH, Thorn RG.. "Basidiomycota: Homobasidiomycetes". In McLaughlin DJ, McLaughlin EG, Lemke PA. The Mycota. VIIB. Systematics and Evolution. Springer-Verlag. pp. 121?68. ISBN 978-3540580089. 
  9. ^ Hib bett DS, Pine EM, Langer E, Langer G, Donoghue MJ. (1997). "Evolution of gilled mushrooms and puffballs inferred from ribosomal DNA sequences". Proceedings of the National Academy of Sciences USA 94 (22): 12002?6. doi:10.1073/pnas.+94.22.12002. 
  10. ^ Moncalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R. (2000). "Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences" (PDF). Systematic Biology 49 (2): 278?305. doi:10.1093/sysbio/49.2.278. PMID 12118409. http://sysbio.oxfordjournals.org/content/49/ 2/278.full.pdf. 
  11. ^ Moncalvo JM, Vilgalys R, Redhead SA, Johnson JE, James TY, Catherine Aime M, Hofstetter V, Verduin SJ, Larsson E, Baroni TJ, Greg Thorn R, Jacobsson S, Cl?men?on H, Miller OK. (2002). "One hundred and seventeen clades of euagarics" (PDF). Molecular Phylogenetics and Evolution 23 (3): 357?400. doi:10.1016/S1055-7903(02)00027-1. PMID 12099793. http://www.umich.edu/~mycology/publications_assets/moncalvo.mpe.2002.pdf. 
  12. ^ Frank JL, Coffan RA, Southworth D. (2010). "Aquati c gilled mushrooms: Psathyrella fruiting in the Rogue River in southern Oregon". Mycologia 102 (1): 93?107. doi:10.3852/07-190. PMID 20120233. 
  13. ^ a b c Alexopolous et al., pp. 508?43.

Sources

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Last Revised: August 24, 2012
2012/08/24 13:08:39