Agaricales

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Agaricales
Temporal range: Upper Aptian – Holocene, 113–0  Ma [1]
Agaricus campestris.jpg
Agaricus campestris (field mushroom) (Agaricaceae)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Subclass: Agaricomycetidae
Order: Agaricales
Underw. (1899) [2]
Subdivisions
46 families, 400+ genera
Synonyms [3]

AmanitalesJülich (1981)
CortinarialesJülich (1981)
EntolomatalesJülich (1981)
FistulinalesJülich (1981)
SchizophyllalesNuss (1980)

Contents

The Agaricales are an order of fungi in the division Basidiomycota. As originally conceived, the order contained all the agarics (gilled mushrooms), 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 (clubs and corals) and gasteroid fungi (puffballs and false truffles) belong within the Agaricales. The order has 46 extant families, more than 400 genera, and over 25,000 described species, [4] along with six extinct genera known only from the fossil record. [5] [6] [1] Species in the Agaricales range from the familiar Agaricus bisporus (cultivated mushroom) and the deadly Amanita virosa (destroying angel) to the coral-like Clavaria zollingeri (violet coral) and bracket-like Fistulina hepatica (beefsteak fungus).

History, classification and phylogeny

Agaricoid clade

Strophariaceae s. str.

Hymenogastraceae

Inocybaceae

Crepidotaceae

Tubarieae

Panaeoleae

Gymnopileae

Cortinariaceae s. str.

Bolbitiaceae

Psathyrellaceae

Hydnangiaceae

Agaricaceae

Nidulariaceae

Cystodermateae

Tricholomatoid clade

Lyophyllaceae

Entolomataceae

Clitocybe candicans , C. subditopoda

Tricholomataceae

Dendrocollybia racemosa

Neohygrophorus angelesianus

Catathelasma clade

Mycenaceae

Marasmioid clade

Omphalotaceae

Marasmiaceae

hydropoid clade

Cyphellaceae

Physalacriaceae

Schizophyllaceae

Lachnellaceae

Hygrophoroid clade

Hygrophoraceae

Pterulaceae

Typhulaceae

Pluteoid clade

Pleurotaceae

Amanitaceae

Pluteaceae

Limnoperdaceae

Plicaturopsidoid clade

Atheliaceae

Clavariaceae

Cladogram of the Agaricales, showing the division into six major clades, based on Matheny et al., 2006. [7]

In his three volumes of Systema Mycologicum published between 1821 and 1832, Elias Fries put almost all of the fleshy, 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, Quélet, 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. [8] In the twentieth century, Rolf Singer's influential work The Agaricales in Modern Taxonomy, published in four editions spanning from 1951 to 1986, used both Fries's macroscopic characters and Fayod's microscopic characters to reorganize families and genera; his final classification included 230 genera within 18 families. [9] Singer treated three major groups within the Agaricales sensu lato : the Agaricales sensu stricto, Boletineae, and Russulales. These groups are still accepted by modern treatments based on DNA analysis, as the euagarics clade, bolete clade, and russuloid clade. [10]

Molecular phylogenetics research has demonstrated that the euagarics clade is roughly equivalent to Singer's Agaricales sensu stricto. [11] [12] [13] A 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. [7]

The bird's nest fungus Cyathus striatus Cyathus striatus 65278694.jpg
The bird's nest fungus Cyathus striatus

Molecular studies have shown that agarics are more divergent than once thought. Agarics in the genera Russula and Lactarius , for example, belong to the order Russulales, whilst agarics in the genera Paxillus and Hygrophoropsis belong in the Boletales. Conversely some genera with non-agaric fruit bodies, such as the puffballs, bird's nest fungi, and many clavarioid fungi, belong in the Agaricales.

Distribution and habitat

Members of the Agaricales are ubiquitous, with species found in all continents. The great majority are terrestrial, in almost every habitat from woodland and grassland to deserts and dunes. Agaricoid species were long thought to be solely terrestrial, until the 2005 discovery of Psathyrella aquatica , the only gilled mushroom known to fruit underwater. [14] Species are variously saprotrophic or ectomycorrhizal, occasionally parasitic on plants or other fungi, and sometimes lichenized.

the Agaricales include six monotypic fossil genera mostly found fossilized in amber. The oldest records are from three Cretaceous age genera; the late Aptian Gondwanagaricites magnificus from the Crato Formation (Brazil), [1] the Albian age (approximately 100 Ma) Palaeoagaracites antiquus from Burmese amber and the slightly younger Turonian New Jersey Amber species Archaeomarasmius leggeti . [5] The three other species, Aureofungus yaniguaensis , Coprinites dominicana and Protomycena electra are known from single specimens found in the Dominican amber mines of Hispaniola. [6]

Genera Incertae sedis

Setchelliogaster is a genus of gasteroid fungi in the Agaricales that is incertae sedis with respect to familial placement. Setchelliogaster 92786.jpg
Setchelliogaster is a genus of gasteroid fungi in the Agaricales that is incertae sedis with respect to familial placement.

There are several genera classified in the Agaricales that are i) poorly known, ii) have not been subjected to DNA analysis, or iii) if analysed phylogenetically do not group with as yet named or identified families, and have not been assigned to a specific family (i.e., Incertae sedis with respect to familial placement). These include:

See also

Related Research Articles

<span class="mw-page-title-main">Strophariaceae</span> Family of fungi

The Strophariaceae are a family of fungi in the order Agaricales. Under an older classification, the family covered 18 genera and 1316 species. The species of Strophariaceae have red-brown to dark brown spore prints, 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. The family was circumscribed in 1946 by mycologists Rolf Singer and Alexander H. Smith.

<span class="mw-page-title-main">Marasmiaceae</span> Family of fungi

The Marasmiaceae are a family of fungi in the order Agaricales. Basidiocarps are most frequently agarics, but occasionally cyphelloid. According to a 2008 estimate, the family contained 54 genera and 1590 species, but recent molecular research, based on cladistic analysis of DNA sequences, has led to a more restricted family concept so that the Marasmiaceae now includes just 13 genera and some 1205 species.

<span class="mw-page-title-main">Clavariaceae</span> Family of fungi

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.

<span class="mw-page-title-main">Tricholomataceae</span> Family of fungi

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.

<i>Stropharia</i> Genus of fungi

The genus Stropharia is a group of medium to large agarics with a distinct membranous ring on the stipe. Well-known members of this genus include the edible Stropharia rugosoannulata and the blue-green verdigris agarics. Stropharia are not generally regarded as good to eat and there are doubts over the edibility of several species. However the species Stropharia rugosoannulata is regarded as prized and delicious when young, and is now the premier mushroom for outdoor bed culture by mycophiles in temperate climates.

<span class="mw-page-title-main">Entolomataceae</span> Family of fungi

The Entolomataceae are a family of fungi in the order Agaricales. The family contains eight genera and 2250 species, the majority of which are in Entoloma. Basidiocarps are typically agaricoid, but a minority are cyphelloid. secotioid, or gasteroid. All produce pink basidiospores that are variously angular (polyhedral), ridged, or nodulose. Species are mostly saprotrophic, though a few are parasitic on other fungi. The family occurs worldwide.

<span class="mw-page-title-main">Hymenogastraceae</span> Family of fungi

The Hymenogastraceae is a family of fungi in the order Agaricales with both agaric and false-truffle shaped fruitbodies. Formerly, prior to molecular analyses, the family was restricted to the false-truffle genera. The mushroom genus Psilocybe in the Hymenogastraceae is now restricted to the hallucinogenic species while nonhallucinogenic former species are largely in the genus Deconica classified in the Strophariaceae.

<i>Melanoleuca</i> Genus of fungi

Melanoleuca is a poorly known genus of saprotrophic mushrooms traditionally classified in the family Tricholomataceae. Most are small to medium sized, white, brown, ocher or gray with a cylindrical to subcylindrical stipe and white to pale yellowish gills. The basidiospores are ellipsoid and ornamented with amyloid warts. Melanoleuca is considered a difficult group to study due to their macroscopic similarities among species and the need of a thorough microscopic analysis to separate species. DNA studies have determined that this genus is closely related to Amanita and Pluteus and that it does not belong to the family Tricholomataceae.

<i>Pholiota</i> Genus of mushrooms

Pholiota is a genus of small to medium-sized, fleshy mushrooms in the family Strophariaceae. They are saprobes that typically live on wood. The genus has a widespread distribution, especially in temperate regions, and contains about 150 species.

<span class="mw-page-title-main">Physalacriaceae</span> Family of fungi

The Physalacriaceae are a family of fungi in the order Agaricales. Species in the family have a widespread distribution, ranging from the Arctic, (Rhizomarasmius), to the tropics, e.g. Gloiocephala, and from marine sites (Mycaureola) and fresh waters (Gloiocephala) to semiarid forests (Xerula).

<span class="mw-page-title-main">Inocybaceae</span> Family of fungi

The Inocybaceae are a family of fungi in the order Agaricales, the largest order of mushroom-forming fungi. It is one of the larger families within Agaricales. This family exhibits an ectomycorrhizal ecology. Members of this family have a widespread distribution in tropical and temperate areas.

<span class="mw-page-title-main">Mycenaceae</span> Family of fungi

The Mycenaceae are a family of fungi in the order Agaricales. According to the Dictionary of the Fungi, the family contains 10 genera and 705 species. This is one of several families that were separated from the Tricholomataceae as a result of phylogenetic analyses. Taxa in the Mycenaceae are saprobic, have a cosmopolitan distribution, and are found in almost all ecological zones. The family was circumscribed by Caspar van Overeem in 1926.

<i>Pseudoomphalina</i> Genus of fungi

Pseudoomphalina is a genus of fungi in the placed in the family Tricholomataceae for convenience. The genus contains six species that are widespread in northern temperate areas. Pseudoomphalina was circumscribed by Rolf Singer in 1956. Pseudoomphalina was found to be paraphyletic to Neohygrophorus in a molecular phylogenetics study and since Pseudoomphalina is an older name, Neohygrophyorus was synonymized with it. The type species of Neohygrophorus was Neohygrophorus angelesianus, now Pseudoomphalina angelesiana. In earlier classifications based on anatomy prior to DNA sequence-based classifications, its unusual combination of features led taxonomists to independently create two subgenera in two genera: Hygrophorus subg. Pseudohygrophorus and Clitocybe subg. Mutabiles; the latter based on Neohygrophorus angelesianus but described under a new species name which is now placed in synonymy, Clitocybe mutabilis. All species of Pseudoomphalina are united by the presence of clamp-connections in their hyphae, an interwoven gill trama and amyloid spores. Pseudoomphalina angelesiana possesses grey-violaceous pigments that turn red in alkali solutions and lacks filiform, hyphal sterile elements in its hymenium and stipitipellis. These were features used to distinguish it from Pseudoomphalina as a genus, but Pseudoomphalina umbrinopurpurascens possesses these same pigments and the filiform elements of Pseudoomphalina. Molecular phylogenetics studies have also found some former species of Pseudoomphalina to belong in other genera. Pseudoomphalina pachyphylla was moved to its own genus, Pseudolaccaria, and Pseudoomphalina clusiliformis was synonymized with it. Pseudoomphalina flavoaurantia and Pseudoomphalina lignicola were found to belong in Clitocybula. Phylogenetically, Pseudoomphalina is in a tricholomatoid clade but not in the Tricholomataceae.

<i>Deconica coprophila</i> Species of fungus

Deconica coprophila, commonly known as the dung-loving psilocybe, or dung demon, is a species of mushroom in the family Strophariaceae. First described as Agaricus coprophilus by Jean Baptiste François Pierre Bulliard in 1793, it was transferred to the genus Psilocybe by Paul Kummer in 1871. In the first decade of the 2000s, several molecular studies showed that the Psilocybe was polyphyletic, and the non-bluing (non-hallucinogenic) species were transferred to Deconica.

<i>Hemipholiota</i> Genus of fungi

Hemipholiota is a genus of agaric fungi in the order Agaricales. It was originally proposed by Rolf Singer in 1962 as a subgenus of Pholiota to contain species with absent or sparse pleurocystidia and absent chrysocystidia. Henri Romagnesi raised it to generic status in 1980, but this naming was invalid as it did not meet the requirements of the International Code of Nomenclature for algae, fungi, and plants. Marcel Bon published the genus validly in 1986.

<i>Bogbodia</i> Genus of fungi

Bogbodia is a bog-inhabiting agaric fungal genus that colonizes peat and Sphagnum and produces tan-colored fruit bodies. The only species in the genus is Bogbodia uda. Characteristically it forms chrysocystidia and rather large, finely roughened, violaceous basidiospores each with a poorly defined germ pore. The genus differs from Hypholoma which has smaller, smooth basidiospores and typically have cespitose fruit bodies and decay wood. Phylogenetically, Bogbodia is distinct from Hypholoma, Pholiota, and Leratiomyces.

<i>Mycopan</i> Genus of fungi

Mycopan is one of several genera of agaric fungi (mushrooms) that were formerly classified in the genus Hydropus or Mycena. Mycopan is currently monotypic, containing the single species Mycopan scabripes. It produces dusky colored fruit bodies that are mycenoid, but lack amyloid or dextrinoid tissues except for the amyloid basidiospores. Its stipe is notably scruffy from cystidioid end cells and unlike true Hydropus it does not bleed clear fluid. Phylogenetically, Mycopan is distant from the Mycenaceae and the type of that family, Mycena, and it is not with the type of Hydropus, Hydropus fuliginarius. Mycopan grouped closest to Baeospora. Baeospora was shown to be in the Cyphellaceae by Matheny and colleagues. Mycopan scabripes grows from debris in forest floors in North America and Europe.

<span class="mw-page-title-main">Porotheleaceae</span> Family of fungi

The Porotheleaceae are a family of saprotrophic, mainly wood-decay fungi in the order Agaricales that are primarily agarics, but also include cyphelloid fungi. The family had been informally cited in the literature as the 'hydropoid' clade. The type genus, Porotheleum, was placed in the phylogenetically defined clade in 2002 but the clade was more strongly supported in 2006 though without including Porotheleum. Its sister group is the Cyphellaceae, both in the 'marasmioid clade'. Some included taxa are cultivated by ants. More recently the family was recognized in three analyses that included Porotheleum.

Stagnicola is an agaric fungal genus that contains the single species Stagnicola perplexa. This fungus colonizes plant debris in wet coniferous forest floor depressions and shallow pools, and fruits after the pools drain or dry in late summer to early fall in North America and Europe. The genus is characterized by smooth, yellowish brown basidiospores lacking a germ pore, and a naucorioid appearance, with brownish mycelium at the base of the stems. Phylogenetically, Stagnicola appeared rather isolated and proved to be closest to Mythicomyces and Mythicomyces was closest to the Psathyrellaceae. In 2019 the two genera were shown to be closely related and placed in a new family, Mythicomycetaceae sister to the Psathyrellaceae.

Flammula is a dark brown-spored genus of mushrooms that cause a decay of trees, on whose bases they often fruit, forming clusters of yellowish brown mushrooms.

References

  1. 1 2 3 Heads, Sam W.; Miller, Andrew N.; Crane, J. Leland; Thomas, M. Jared; Ruffatto, Danielle M.; Methven, Andrew S.; Raudabaugh, Daniel B.; Wang, Yinan (2017). "The oldest fossil mushroom". PLOS ONE. 12 (6): e0178327. Bibcode:2017PLoSO..1278327H. doi: 10.1371/journal.pone.0178327 . PMC   5462346 . PMID   28591180.
  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. Retrieved 2010-12-30.
  4. ↑ "Catalogue of Life" . Retrieved 2023-02-15.
  5. 1 2 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.
  6. 1 2 Hibbett DS, Binder M, Wang Z, Goldman Y (2003). "Another Fossil Agaric from Dominican Amber". Mycologia. 95 (4): 685–687. doi:10.2307/3761943. JSTOR   3761943. PMID   21148976.(subscription required)
  7. 1 2 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. Archived from the original (PDF) on 2016-03-03.
  8. ↑ Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CABI. p. 12. ISBN   978-0-85199-826-8.
  9. ↑ Singer R. (1986). The Agaricales in Modern Taxonomy (4th ed.). Koenigstein Königstein im Taunus, Germany: Koeltz Scientific Books. ISBN   978-3-87429-254-2.
  10. ↑ Hibbett DH, Thorn RG. "Basidiomycota: Homobasidiomycetes". In McLaughlin DJ, McLaughlin EG, Lemke PA (eds.). The Mycota. VIIB. Systematics and Evolution. Springer-Verlag. pp. 121–68. ISBN   978-3-540-58008-9.
  11. ↑ 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 of the United States of America. 94 (22): 12002–6. Bibcode:1997PNAS...9412002H. doi: 10.1073/pnas.94.22.12002 . PMC   23683 . PMID   9342352.
  12. ↑ Moncalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R (2000). "Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences". Systematic Biology. 49 (2): 278–305. doi:10.1093/sysbio/49.2.278. PMID   12118409.
  13. ↑ 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, Clémençon 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.
  14. ↑ Frank JL, Coffan RA, Southworth D (2010). "Aquatic gilled mushrooms: Psathyrella fruiting in the Rogue River in southern Oregon". Mycologia. 102 (1): 93–107. doi:10.3852/07-190. PMID   20120233. S2CID   7175296.
  15. ↑ Jacobsson S, Larsson E (2007). "Hemistropharia, a new genus in Agaricales". Mycotaxon. 102: 235–40.


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