Agaricomycetes

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]

Agaricomycetes Temporal range: Barremian–present PreꞒ Ꞓ O S D C P T J K Pg N
Amanita muscaria (Agaricales)
Scientific classification
Kingdom:	Fungi
Division:	Basidiomycota
Subdivision:	Agaricomycotina
Class:	Agaricomycetes Doweld (2001)[1]
Subclasses/Orders
Agaricomycetidae Agaricales (32 fam., 410+ gen.) Amylocorticiales (1 fam., 14 gen.) Atheliales (1 fam., 22 gen.) Boletales (16 fam., 95+ gen.) Jaapiales (1 fam., 1 gen.) Lepidostromatales (1 fam., 3 gen.) Phallomycetidae Geastrales (1 fam., 8 gen.) Gomphales (3 fam., 18 gen.) Hysterangiales (5 fam., 18 gen.) Phallales (2 fam., 26 gen.) incertae sedis (no subclass) Auriculariales (6–7 fam., 30+ gen.) Cantharellales (7 fam., 39 gen.) Corticiales (3 fam., 30+ gen.) Gloeophyllales (1 fam., 7 gen.) Hymenochaetales (3 fam., 50+ gen.) Polyporales (9 fam., ~200 gen.) Russulales (12 fam., 80+ gen.) Sebacinales (1 fam., 8 gen.) Stereopsidales (1 fam., 2 gen.) Thelephorales (2 fam., 18 gen.) Trechisporales (1 fam., 15 gen.)

Classification

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.

Features

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 150,000 square metres (37 acres) with a mass of 10,000 kg (22,000 lb) and an age of 1,500 years.[10]

Ecology

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.

Evolution

A study of 5,284 species with a backbone phylogeny based on 104 genomes[11] 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)

Fossil record

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).[12] 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.[13] Based on molecular clock analysis, the Agaricomycetes are estimated to be about 290 million years old.[14]

Phylogeny

Modern molecular phylogenetics suggest the following relationships:[15]

Basidiomycetes (outgroup)   Agaricomycetes   Cantharellales       Sebacinales       Auriculariales         Stereopsidales   Phallomycetidae   Geastrales       Hysterangiales       Gomphales     Phallales               Trechisporales       Hymenochaetales         Thelephorales     Polyporales           Corticiales       Jaapiales     Gloeophyllales           Russulales   Agaricomycetidae   Agaricales       Boletales       Amylocorticiales       Lepidostromatales     Atheliales

Genera incertae sedis

There are many genera in the Agaricomycetes that have not been classified in any order or family. These include:

• Akenomyces
• Aldridgea
• Anixia
• Arrasia[16]
• Arthrodochium
• Arualis
• Atraporiella
• Cenangiomyces
• Ceraceopsis
• Corticomyces
• Cruciger
• Dendrosporomyces
• Ellula
• Fibulochlamys
• Fibulocoela
• Fibulotaeniella
• Geotrichopsis[17]
• Gloeoradulum
• Gloeosynnema
• Glomerulomyces
• Glutinoagger
• Grandinia
• Granulocystis
• Hallenbergia
• Hyphobasidiofera
• Hypolyssus
• Intextomyces
• Korupella
• Minostroscyta[18]
• Mylittopsis
• Odonticium
• Pagidospora
• Peniophorella
• Phlyctibasidium
• Pseudasterodon
• Purpureocorticium S.H.Wu (2017)[19]
• Pycnovellomyces
• Resinicium
• Riessia
• Riessiella
• Skvortzovia
• Taiwanoporia[20]
• Timgrovea
• Titaeella
• Trechinothus
• Tricladiomyces
• Trimitiella
• Tubulicrinopsis
• Xanthoporus
• Xenosoma

References

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2. Hibbett DS, Thorn RG (2001). McLaughlin DJ, et al. (eds.). The Mycota, Vol. VII. Part B., Systematics and Evolution. Berlin, Germany: Springer-Verlag. pp. 121–168.
3. Hibbett DS; et al. (2007). "A higher level phylogenetic classification of the Fungi". Mycological Research. 111 (5): 509–547. CiteSeerX 10.1.1.626.9582. doi:10.1016/j.mycres.2007.03.004. PMID 17572334.
4. Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CAB International. pp. 12–13. ISBN 978-0-85199-826-8.
5. Binder M, Larsson KH, Matheny PB, Hibbett DS (2010). "Amylocorticiales ord. nov. and Jaapiales ord. nov.: Early diverging clades of Agaricomycetidae dominated by corticioid forms". Mycologia. 102 (4): 865–880. doi:10.3852/09-288. PMID 20648753.
6. Sjökvist E, Pfeil BE, Larsson E, Larsson K-H (2014). "Stereopsidales – a new order of mushroom-forming fungi". PLOS ONE. 9 (8): e106204. Bibcode:2014PLoSO...995227S. doi:10.1371/journal.pone.0095227. PMC 4002437. PMID 24777067.
7. Hodkinson BP, Moncada B, Lücking R (2014). "Lepidostromatales, a new order of lichenized fungi (Basidiomycota, Agaricomycetes), with two new genera, Ertzia and Sulzbacheromyces, and one new species, Lepidostroma winklerianum". Fungal Diversity. 64 (1): 165–179. doi:10.1007/s13225-013-0267-0.
8. Fries EM (1874). Hymenomycetes Europaei (in Latin). Uppsala: Typis Descripsit Ed. Berling. p. 1.
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10. Smith M, Bruhn JH, Anderson JB (1992). "The fungus Armillaria bulbosa is among the largest and oldest living organisms". Nature. 356 (6368): 428–431. Bibcode:1992Natur.356..428S. doi:10.1038/356428a0.
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15. Hibbett D, Bauer R, Binder M, Giachini AJ, Hosaka K, Justo A, Larsson E, Larsson K-H, Lawrey JD, Miettinen O, Nagy LG, Nilsson RH, Weiss M, Thorn RG (2014). "Agaricomycetes". In McLaughlin DJ, Spatafora JW (eds.). Systematics and Evolution. The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research. 7A (2nd ed.). Berlin, Heidelberg: Springer-Verlag. pp. 373–429. doi:10.1007/978-3-642-55318-9_14. ISBN 978-3-642-55317-2.
16. Berniccia, Annarosa; Gorjón, Sergio P.; Nakasone, Karen K. (2011). "Arrasia rostrata (Basidiomycota), a new corticioid genus and species from Italy" (PDF). Mycotaxon. 118: 257–264. doi:10.5248/118.257.
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18. Hjortstam, Kurt; Ryvarden, Leif (2001). "Corticioid species (Basidiomycotina, Aphyllophorales) from Colombia III". Mycotaxon. 79: 189–200.
19. Wu, Sheng-Hua; Wang, Dong-Me; Chen, Yu-Ping (2018). "Purpureocorticium microsporum (Basidiomycota) gen. et sp. nov. from East Asia". Mycological Progress. 17 (3): 357–364. doi:10.1007/s11557-017-1362-5.
20. Chang, TunTschu; Chou, Wen Neng (2003). "Taiwanoporia, a new aphyllophoralean genus". Mycologia. 95 (6): 1215–1218. doi:10.1080/15572536.2004.11833029. JSTOR 3761921. PMID 21149022.

External links

• Data related to Agaricomycetes at Wikispecies
• Tree of Life Agaricomycetes by David S. Hibbett
• Overview of the Basidiomycota from Aarhus University, Denmark
• Evolution & Morphology in the Homobasidiomycetes