Editing Basidiomycota

{{Short description|Division of fungi}}
{{Automatic taxobox
|fossil_range = [[Carboniferous]]-present{{fossil range|330|0|<ref>{{Cite journal|last1=Krings|first1=Michael|last2=Dotzler|first2=Nora|last3=Galtier|first3=Jean|last4=Taylor|first4=Thomas N.|date=Jan 2011|title=Oldest fossil basidiomycete clamp connections|url=https://www.jstage.jst.go.jp/article/mycosci/52/1/52_MYC52018/_article/-char/en|journal=Mycoscience|language=en|volume=52|issue=1|pages=18–23|doi=10.1007/S10267-010-0065-4|hdl=1808/13681|hdl-access=free}}</ref>}}
| image = Haeckel Basimycetes.jpg
| image_caption = Basidiomycetes from [[Ernst Haeckel]]'s 1904 ''[[Kunstformen der Natur]]''
| taxon = Basidiomycota
| authority = [[Royall T. Moore|R.T.Moore]] (1980)<ref name=Moore>{{cite journal | last=Moore | first = R. T. | title=Taxonomic proposals for the classification of marine yeasts and other yeast-like fungi including the smuts | date = 1980 | journal=Botanica Marina| volume=23 | issue = 6 | page=371 | doi = 10.1515/bot-1980-230605 | bibcode = 1980BoMar..23..361M }}</ref>
| subdivision_ranks = Subdivisions and classes
| subdivision =
:[[Agaricomycotina]]
:[[Pucciniomycotina]]
:[[Ustilaginomycotina]]
}}

'''Basidiomycota''' ({{IPAc-en|b|ə|ˌ|s|ɪ|d|i|.|oʊ|m|aɪ|ˈ|k|oʊ|t|ə}}){{refn|{{cite Dictionary.com|Basidiomycota}}}} is one of two large [[division (mycology)|divisions]] that, together with the [[Ascomycota]], constitute the subkingdom [[Dikarya]] (often referred to as the "'''higher fungi'''") within the kingdom [[Fungi]]. Members are known as '''basidiomycetes'''.<ref name="Li-et-al-2007" /> More specifically, Basidiomycota includes these groups: [[agaric]]s, [[puffball]]s, [[Phallaceae|stinkhorns]], [[Bracket fungus|bracket fungi]], other [[polypore]]s, [[Jelly fungus|jelly fungi]], [[bolete]]s, [[Cantharellus|chanterelles]], [[Geastraceae|earth stars]], [[Smut (fungus)|smuts]], [[Common bunt|bunts]], [[Rust (fungus)|rusts]], [[Sporobolomyces|mirror yeasts]], and ''[[Cryptococcus (fungus)|Cryptococcus]]'', the human pathogenic yeast.

Basidiomycota are filamentous fungi composed of [[hypha]]e (except for [[Yeast|basidiomycota-yeast]]) and reproduce sexually via the formation of specialized club-shaped end [[Cell (biology)|cells]] called [[Basidium|basidia]] that normally bear external [[Meiosis|meiospores]] (usually four). These specialized [[spore]]s are called [[basidiospores]].<ref>{{cite journal |last1=Rivera-Mariani |first1=F.E. |last2=Bolaños-Rosero |first2=B. |title=Allergenicity of airborne basidiospores and ascospores: need for further studies |journal=Aerobiologia |date=2011 |volume=28 |issue=2 |pages=83–97 |doi=10.1007/s10453-011-9234-y |s2cid=84615057 |url=https://www.researchgate.net/publication/255823215 |access-date=16 January 2019}}</ref> However, some Basidiomycota are obligate [[Asexual reproduction|asexual]] reproducers. Basidiomycota that reproduce asexually (discussed below) can typically be recognized as members of this division by gross similarity to others, by the formation of a distinctive anatomical feature (the [[clamp connection]]), [[cell wall]] components, and definitively by [[phylogenetic]] molecular analysis of [[DNA sequence]] data.

==Classification==

A 2007 classification, adopted by a coalition of 67 [[mycologist]]s recognized three subphyla ([[Pucciniomycotina]], [[Ustilaginomycotina]], [[Agaricomycotina]]) and two other class level taxa ([[Wallemiomycetes]], [[Entorrhizomycetes]]) outside of these, among the Basidiomycota.<ref name="Hibbett">{{Cite journal|title = A higher-level phylogenetic classification of the Fungi|journal = [[Mycological Research]]|date = May 2007|pages = 509–547|volume = 111|issue = 5|doi = 10.1016/j.mycres.2007.03.004|first1 = David S.|last1 = Hibbett|first2 = Manfred|last2 = Binder|first3 = Joseph F.|last3 = Bischoff|first4 = Meredith|last4 = Blackwell|first5 = Paul F.|last5 = Cannon|first6 = Ove E.|last6 = Eriksson|first7 = Sabine|last7 = Huhndorf|first8 = Timothy|last8 = James|first9 = Paul M.|last9 = Kirk|pmid = 17572334|citeseerx = 10.1.1.626.9582| s2cid=4686378 }}</ref> As now classified, the subphyla join and also cut across various obsolete taxonomic groups (see below) previously commonly used to describe Basidiomycota. According to a 2008 estimate, Basidiomycota comprise three subphyla (including six unassigned classes) 16 classes, 52 orders, 177 families, 1,589 genera, and 31,515 species.<ref name="Kirk p.78-79">{{harvnb|Kirk|Cannon|Stalpers|2008|pp=78–79}}</ref>
Wijayawardene et al. 2020 produced an update that recognized 19 classes ([[Agaricomycetes]], [[Agaricostilbomycetes]], [[Atractiellomycetes]], [[Bartheletiomycetes]], [[Classiculomycetes]], [[Cryptomycocolacomycetes]], [[Cystobasidiomycetes]], [[Dacrymycetes]], [[Exobasidiomycetes]], [[Malasseziomycetes]], [[Microbotryomycetes]], [[Mixiomycetes]], [[Monilielliomycetes]], [[Pucciniomycetes]], [[Spiculogloeomycetes]], [[Tremellomycetes]], [[Tritirachiomycetes]], [[Ustilaginomycetes]] and [[Wallemiomycetes]]) with multiple orders and genera.<ref name="Wijayawardene et al. 2020">{{cite journal |display-authors=6 |last1=Wijayawardene |first1=Nalin |last2=Hyde |first2=Kevin |first3=Laith Khalil Tawfeeq |last3=Al-Ani |last4=Somayeh |first4=Dolatabadi |last5=Stadler |first5=Marc |last6=Haelewaters |first6=Danny |last7=Tsurykau |first7=Andrei |last8=Mesic |first8=Armin |last9=Navathe |first9=Sudhir |last10=Papp |first10=Viktor |last11=Oliveira Fiuza |first11=Patrícia |last12=Vázquez |first12=Víctor |last13=Gautam |first13=Ajay |last14=Becerra |first14=Alejandra G. |last15=Ekanayaka |first15=Anusha |last16=K. C. |first16=Rajeshkumar |last17=Bezerra |first17=Jadson |last18=Matočec |first18=Neven |last19=Maharachchikumbura |first19=Sajeewa |last20=Suetrong |first20=Satinee |year=2020 |title=Outline of Fungi and fungus-like taxa |journal=Mycosphere |volume=11 |pages=1060–1456 |doi=10.5943/mycosphere/11/1/8 |doi-access=free|hdl=10481/61998 |hdl-access=free }}</ref>

Traditionally, the Basidiomycota were divided into two classes, now obsolete:{{cn|date=February 2025}}
* [[Agaricomycetes|Homobasidiomycetes]] (alternatively called holobasidiomycetes), including true [[mushroom]]s
* [[Heterobasidiomycetes]], including the [[Jelly fungus|jelly]], [[Rust (fungus)|rust]] and [[Smut (fungus)|smut]] fungi
Nonetheless these former concepts continue to be used as two types of [[growth habit]] groupings, the "mushrooms" (e.g. ''[[Schizophyllum commune]]'') and the non-mushrooms (e.g. ''[[Mycosarcoma maydis]]'').<ref name="Li-et-al-2007">{{cite journal | last1=Li | first1=Liande | last2=Wright | first2=Sara J. | last3=Krystofova | first3=Svetlana | last4=Park | first4=Gyungsoon | last5=Borkovich | first5=Katherine A. | title=Heterotrimeric G Protein Signaling in Filamentous Fungi | journal=[[Annual Review of Microbiology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=61 | issue=1 | year=2007 | issn=0066-4227 | doi=10.1146/annurev.micro.61.080706.093432 | pages=423–452| pmid=17506673 }}</ref>

===Agaricomycotina===
The [[Agaricomycotina]] include what had previously been called the [[Hymenomycetes]] (an obsolete morphological based class of Basidiomycota that formed [[hymenial]] layers on their [[sporocarp (fungus)|fruitbodies]]), the [[Gasteromycetes]] (another obsolete class that included species mostly lacking hymenia and mostly forming spores in enclosed [[sporocarp (fungus)|fruitbodies]]), as well as most of the [[jelly fungi]]. This sub-phyla also includes the "classic" mushrooms, polypores, corals, chanterelles, crusts, puffballs and stinkhorns.<ref>{{cite web|url=https://cpb-us-e1.wpmucdn.com/blogs.uoregon.edu/dist/c/8944/files/2014/09/Lecture-7-Ustilaginomycets-1w4kndx.pdf |archive-url=https://web.archive.org/web/20200730231800/https://cpb-us-e1.wpmucdn.com/blogs.uoregon.edu/dist/c/8944/files/2014/09/Lecture-7-Ustilaginomycets-1w4kndx.pdf |archive-date=2020-07-30 |url-status=live|title=Lecture 7 : Ustilaginomycets|website=Cpb-us-e1.wpmucdn.com|access-date=2 March 2022}}</ref> The three classes in the Agaricomycotina are the [[Agaricomycetes]], the [[Dacrymycetes]], and the [[Tremellomycetes]].<ref>{{harvnb|Kirk|Cannon|Stalpers|2008|p=13}}</ref>

The class [[Wallemiomycetes]] is not yet placed in a subdivision, but recent genomic evidence suggests that it is a sister group of [[Agaricomycotina]].<ref>{{cite journal | last1 = Zajc | first1 = J. | last2 = Liu | first2 = Y. | last3 = Dai | first3 = W. | last4 = Yang | first4 = Z. | last5 = Hu | first5 = J. | last6 = Gostinčar | first6 = C. | last7 = Gunde-Cimerman | first7 = N. |title=Genome and transcriptome sequencing of the halophilic fungus Wallemia ichthyophaga: haloadaptations present and absent|journal=[[BMC Genomics]]|date=Sep 13, 2013|volume=14|page=617|pmid=24034603|doi=10.1186/1471-2164-14-617 |pmc=3849046 | doi-access = free }}</ref><ref>{{cite journal | last1 = Padamsee | first1 = M. | last2 = Kumar | first2 = T. K. | last3 = Riley | first3 = R. | last4 = Binder | first4 = M. | last5 = Boyd | first5 = A. | last6 = Calvo | first6 = A. M. | last7 = Furukawa | first7 = K. | last8 = Hesse | first8 = C. | last9 = Hohmann | first9 = S. | last10 = James | first10 = T. Y. | last11 = LaButti | first11 = K. | last12 = Lapidus | first12 = A. | last13 = Lindquist | first13 = E. | last14 = Lucas | first14 = S. | last15 = Miller | first15 = K. | last16 = Shantappa | first16 = S. | last17 = Grigoriev | first17 = I. V. | last18 = Hibbett | first18 = D. S. | last19 = McLaughlin | first19 = D. J. | last20 = Spatafora | first20 = J. W. | last21 = Aime | first21 = M. C. |title=The genome of the xerotolerant mold Wallemia sebi reveals adaptations to osmotic stress and suggests cryptic sexual reproduction|journal=[[Fungal Genetics and Biology]] |date=Mar 2012|volume=49|issue=3|pages=217–26|pmid=22326418|doi=10.1016/j.fgb.2012.01.007| url = http://www.escholarship.org/uc/item/7d6261dx }}</ref>

===Pucciniomycotina===
The [[Pucciniomycotina]] include the rust fungi, the insect parasitic/symbiotic genus ''[[Septobasidium]]'', a former group of smut fungi (in the [[Microbotryomycetes]], which includes mirror yeasts), and a mixture of odd, infrequently seen, or seldom recognized fungi, often parasitic on plants. The eight classes in the Pucciniomycotina are [[Agaricostilbomycetes]], [[Atractiellomycetes]], [[Classiculomycetes]], [[Cryptomycocolacomycetes]], [[Cystobasidiomycetes]], [[Microbotryomycetes]], [[Mixiomycetes]], and [[Pucciniomycetes]].<ref>{{harvnb|Kirk|Cannon|Stalpers|2008|p=581}}</ref>

===Ustilaginomycotina===
The [[Ustilaginomycotina]] are most (but not all) of the former smut fungi and the [[Exobasidiales]]. The classes of the Ustilaginomycotina are the [[Exobasidiomycetes]], the Entorrhizomycetes, and the [[Ustilaginomycetes]].<ref>{{harvnb|Kirk|Cannon|Stalpers|2008|pp=717–718}}</ref>

===Genera included===
There are several genera classified in the Basidiomycota that are 1) poorly known, 2) have not been subjected to DNA analysis, or 3) if analysed phylogenetically do not group with as yet named or identified families, and have not been assigned to a specific family (i.e., they are ''[[incertae sedis]]'' with respect to familial placement). These include:
{{Div col}}
* ''[[Anastomyces]]'' <small>W.P.Wu, B.Sutton & Gange (1997)</small>
* ''[[Anguillomyces]]'' <small>Marvanová & Bärl. (2000)</small>
* ''[[Anthoseptobasidium]]'' <small>Rick (1943)</small>
* ''[[Arcispora]]'' <small>Marvanová & Bärl. (1998)</small>
* ''[[Arrasia]]'' <small>Bernicchia, Gorjón & Nakasone (2011)</small>
* ''[[Brevicellopsis]]'' <small>Hjortstam & Ryvarden (2008)</small>
* ''[[Celatogloea]]'' <small>P.Roberts (2005)</small>
* ''[[Cleistocybe]]'' <small>Ammirati, A.D.Parker & Matheny (2007)</small>
* ''[[Cystogloea]]'' <small>P. Roberts (2006)</small>
* ''[[Dacryomycetopsis]]'' <small>Rick (1958)</small>
* ''[[Eriocybe]]'' <small>Vellinga (2011)</small>
* ''[[Hallenbergia]]'' <small>Dhingra & Priyanka (2011)</small>
* ''[[Hymenoporus]]'' <small>Tkalčec, Mešić & Chun Y.Deng (2015)</small>
* ''[[Kryptastrina]]'' <small>Oberw. (1990)</small>
* ''[[Microstella]]'' <small>K.Ando & Tubaki (1984)</small>
* ''[[Neotyphula]]'' <small>Wakef. (1934)</small>
* ''[[Nodulospora]]'' <small>Marvanová & Bärl. (2000)</small>
* ''[[Paraphelaria]]'' <small>Corner (1966)</small>
* ''[[Punctulariopsis]]'' <small>Ghob.-Nejh. (2010)</small>
* ''[[Radulodontia]]'' <small>Hjortstam & Ryvarden (2008)</small>
* ''[[Restilago]]'' <small>Vánky (2008)</small>
* ''[[Sinofavus]]'' <small>W.Y.Zhuang (2008)</small>
* ''[[Zanchia]]'' <small>Rick (1958)</small>
* ''[[Zygodesmus]]'' <small>Corda (1837)</small>
* ''[[Zygogloea]]'' <small>P.Roberts (1994)</small>
{{div col end}}

==Typical life cycle==
[[File:Fungi Sexual reproduction cycle.png|thumb|upright=2|Sexual reproduction cycle of basidiomycetes]]
[[File:03 01 07 life cycle Basidiomycota basidium (M. Piepenbring).svg|thumb|Basidiomycota life cycle]]
[[File:Fungus cell cycle-en.svg|thumb|Cell cycle of a Dikaryotic basidiomycete]]
Unlike animals and plants which have readily recognizable male and female counterparts, Basidiomycota (except for the [[rust (fungus)|Rust]] ([[Pucciniales]])) tend to have mutually indistinguishable, compatible [[haploids]] which are usually [[mycelia]] being composed of filamentous [[hyphae]]. Typically haploid Basidiomycota mycelia fuse via [[plasmogamy]] and then the compatible nuclei migrate into each other's mycelia and pair up with the resident nuclei. [[Karyogamy]] is delayed, so that the compatible nuclei remain in pairs, called a [[dikaryon]]. The hyphae are then said to be dikaryotic. Conversely, the haploid mycelia are called [[monokaryon]]s. Often, the dikaryotic mycelium is more vigorous than the individual monokaryotic mycelia, and proceeds to take over the substrate in which they are growing. The dikaryons can be long-lived, lasting years, decades, or centuries. The monokaryons are {{Em|neither}} male nor female. They have either a {{Em|bipolar}} ({{Em|unifactorial}}) or a {{Em|tetrapolar}} ({{Em|bifactorial}}) mating system. This results in the fact that following [[meiosis]], the resulting haploid [[basidiospore]]s and resultant monokaryons, have nuclei that are compatible with 50% (if bipolar) or 25% (if tetrapolar) of their sister basidiospores (and their resultant monokaryons) because the mating genes must differ for them to be compatible. However, there are sometimes more than two possible alleles for a given locus, and in such species, depending on the specifics, over 90% of monokaryons could be compatible with each other.{{cn|date=January 2024}}

The maintenance of the dikaryotic status in dikaryons in many Basidiomycota is facilitated by the formation of [[clamp connection]]s that physically appear to help coordinate and re-establish pairs of compatible nuclei following synchronous [[mitotic]] nuclear divisions. Variations are frequent and multiple. In a typical Basidiomycota lifecycle the long lasting dikaryons periodically (seasonally or occasionally) produce [[basidia]], the specialized usually club-shaped end cells, in which a pair of compatible nuclei fuse ([[karyogamy]]) to form a [[diploid]] cell. [[Meiosis]] follows shortly with the production of 4 haploid nuclei that migrate into 4 external, usually apical basidiospores. Variations occur, however. Typically the basidiospores are [[Ballistics|ballistic]], hence they are sometimes also called [[ballistospore]]s. In most species, the basidiospores disperse and each can start a new haploid mycelium, continuing the lifecycle. Basidia are microscopic but they are often produced on or in multicelled large fructifications called [[basidiocarp]]s or basidiomes, or [[Sporocarp (fungi)|fruitbodies]], variously called mushrooms, [[puffball]]s, etc. Ballistic basidiospores are formed on [[sterigmata]] which are tapered spine-like projections on basidia, and are typically curved, like the horns of a bull. In some Basidiomycota the spores are not ballistic, and the sterigmata may be straight, reduced to stubs, or absent. The basidiospores of these non-ballistosporic basidia may either bud off, or be released via dissolution or disintegration of the basidia.{{cn|date=February 2025}}

[[Image:Basidium schematic.svg|thumb|upright=1.5|Scheme of a typical basidiocarp, the dipoid reproductive structure of a basidiomycete, showing fruiting body, [[hymenium]] and basidia.]]

In summary, meiosis takes place in a diploid basidium. Each one of the four haploid nuclei migrates into its own basidiospore. The basidiospores are ballistically discharged and start new haploid mycelia called monokaryons. There are no males or females, rather there are compatible thalli with multiple compatibility factors. Plasmogamy between compatible individuals leads to delayed karyogamy leading to establishment of a dikaryon. The dikaryon is long lasting but ultimately gives rise to either fruitbodies with basidia or directly to basidia without fruitbodies. The paired dikaryon in the basidium fuse (i.e. karyogamy takes place). The diploid basidium begins the cycle again.{{cn|date=February 2025}}

===Meiosis===
''[[Coprinopsis cinerea]]'' is a basidiomycete mushroom. It is particularly suited to the study of [[meiosis]] because meiosis progresses synchronously in about 10 million cells within the mushroom cap, and the meiotic prophase stage is prolonged. Burns et al.<ref name="pmid20885784">{{cite journal | last1 = Burns | first1 = C. | last2 = Stajich | first2 = J. E. | last3 = Rechtsteiner | first3 = A. | last4 = Casselton | first4 = L. | last5 = Hanlon | first5 = S. E. | last6 = Wilke | first6 = S. K. | last7 = Savytskyy | first7 = O. P. | last8 = Gathman | first8 = A. C. | last9 = Lilly | first9 = W. W. | last10 = Lieb | first10 = J. D. | last11 = Zolan | first11 = M. E. | last12 = Pukkila | first12 = P. J. |title=Analysis of the Basidiomycete Coprinopsis cinerea reveals conservation of the core meiotic expression program over half a billion years of evolution |journal=[[PLOS Genetics]] |volume=6 |issue=9 |pages=e1001135 |date=September 2010  |pmid=20885784 |pmc=2944786 |doi=10.1371/journal.pgen.1001135 | doi-access = free }}</ref> studied the expression of genes involved in the 15-hour meiotic process, and found that the pattern of gene expression of ''C. cinerea'' was similar to two other fungal species, the yeasts ''[[Saccharomyces cerevisiae]]'' and ''[[Schizosaccharomyces pombe]]''. These similarities in the patterns of expression led to the conclusion that the core expression program of meiosis has been conserved in these fungi for over half a billion years of evolution since these species diverged.<ref name="pmid20885784" />

''[[Cryptococcus neoformans]]'' and ''[[Mycosarcoma maydis]]'' are examples of pathogenic basidiomycota. Such pathogens must be able to overcome the oxidative defenses of their respective hosts in order to produce a successful infection. The ability to undergo meiosis may provide a survival benefit for these fungi by promoting successful infection. A characteristic central feature of meiosis is recombination between homologous chromosomes. This process is associated with repair of DNA damage, particularly [[double-strand break]]s.  The ability of ''C. neoformans'' and ''M. maydis'' to undergo meiosis may contribute to their virulence by repairing the oxidative DNA damage caused by their host's release of [[reactive oxygen species]].<ref name="pmid15846346">{{cite journal | last1 = Lin | first1 = X. | last2 = Hull | first2 = C. M. |author-link2=Christina M. Hull | last3 = Heitman | first3 = J. | s2cid = 3195603 |title=Sexual reproduction between partners of the same mating type in Cryptococcus neoformans |journal=[[Nature (journal)|Nature]] |volume=434 |issue=7036 |pages=1017–21 |date=April 2005  |pmid=15846346 |doi=10.1038/nature03448 | bibcode = 2005Natur.434.1017L }}</ref><ref name="pmid18295550">{{cite journal | last1 = Michod | first1 = R. E. | last2 = Bernstein | first2 = H.  | last3 = Nedelcu | first3 = A. M. |title=Adaptive value of sex in microbial pathogens |journal=[[Infection, Genetics and Evolution]] |volume=8 |issue=3 |pages=267–85 |date=May 2008  |pmid=18295550 |doi=10.1016/j.meegid.2008.01.002 | bibcode = 2008InfGE...8..267M }}</ref>

==Variations in lifecycles==
Many variations occur: some variations are self-compatible and spontaneously form dikaryons without a separate compatible thallus being involved. These fungi are said to be homothallic, versus the normal heterothallic species with mating types. Others are secondarily homothallic, in that two compatible nuclei following meiosis migrate into each basidiospore, which is then dispersed as a pre-existing dikaryon. Often such species form only two spores per basidium, but that too varies. Following meiosis, mitotic divisions can occur in the basidium. Multiple numbers of basidiospores can result, including odd numbers via degeneration of nuclei, or pairing up of nuclei, or lack of migration of nuclei. For example, the chanterelle genus ''[[Craterellus]]'' often has six-spored basidia, while some corticioid ''[[Sistotrema]]'' species can have two-, four-, six-, or eight-spored basidia, and the cultivated button mushroom, ''[[Agaricus bisporus]]''. can have one-, two-, three- or four-spored basidia under some circumstances. Occasionally, monokaryons of some taxa can form morphologically fully formed basidiomes and anatomically correct basidia and ballistic basidiospores in the absence of dikaryon formation, diploid nuclei, and meiosis. A rare few number of taxa have extended diploid lifecycles, but can be common species. Examples exist in the mushroom genera ''[[Armillaria]]'' and ''[[Xerula]]'', both in the [[Physalacriaceae]]. Occasionally, basidiospores are not formed and parts of the "basidia" act as the dispersal agents, e.g. the peculiar mycoparasitic jelly fungus, ''[[Tetragoniomyces]]'' or the entire "basidium" acts as a "spore", e.g. in some false puffballs (''[[Scleroderma (genus)|Scleroderma]]''). In the human pathogenic genus ''[[Cryptococcus (fungus)|Cryptococcus]]'', four nuclei following meiosis remain in the basidium, but continually divide mitotically, each nucleus migrating into synchronously forming nonballistic basidiospores that are then pushed upwards by another set forming below them, resulting in four parallel chains of dry "basidiospores".{{cn|date=January 2024}}

Other variations occur: some as standard lifecycles (that themselves have variations within variations) within specific orders.{{cn|date=January 2024}}

===Rusts===
[[rust (fungus)|Rusts]] ([[Pucciniales]], previously known as [[Uredinales]]) at their greatest complexity, produce five different types of spores on two different host plants in two unrelated host families. Such rusts are heteroecious (requiring two hosts) and macrocyclic (producing all five spores types). Wheat [[stem rust]] is an example. By convention, the stages and spore states are numbered by [[Roman numerals]]. Typically, basidiospores infect host one, also known as the alternate or sexual host, and the mycelium forms [[pycnidia]], which are miniature, flask-shaped, hollow, submicroscopic bodies embedded in the host tissue (such as a leaf). This stage, numbered "0", produces single-celled spores that ooze out in a sweet liquid and that act as nonmotile [[spermatia]], and also protruding [[receptive hyphae]]. [[Insect]]s and probably other [[Vector (epidemiology)|vectors]] such as rain carry the spermatia from spermagonium to spermagonium, cross inoculating the mating types. Neither thallus is male or female. Once crossed, the dikaryons are established and a second spore stage is formed, numbered "I" and called [[aecia]], which form dikaryotic [[aeciospore]]s in dry chains in inverted cup-shaped bodies embedded in host tissue. These aeciospores then infect the second host, known as the primary or asexual host (in macrocyclic rusts). On the primary host a repeating spore stage is formed, numbered "II", the [[urediospore]]s in dry pustules called [[uredinia]]. Urediospores are dikaryotic and can infect the same host that produced them. They repeatedly infect this host over the growing season. At the end of the season, a fourth spore type, the [[teliospore]], is formed. It is thicker-walled and serves to overwinter or to survive other harsh conditions. It does not continue the infection process, rather it remains dormant for a period and then germinates to form basidia (stage "IV"), sometimes called a [[promycelium]]. In the Pucciniales, the basidia are cylindrical and become 3-[[septum|septate]] after meiosis, with each of the 4 cells bearing one basidiospore each. The basidiospores disperse and start the infection process on host 1 again. [[Autoecious]] rusts complete their life-cycles on one host instead of two, and microcyclic rusts cut out one or more stages.{{cn|date=January 2024}}

===Smuts===
The characteristic part of the life-cycle of [[smut (fungus)|smuts]] is the thick-walled, often darkly pigmented, ornate, teliospore that serves to survive harsh conditions such as overwintering and also serves to help disperse the fungus as dry [[Diaspore (botany)|diaspores]]. The teliospores are initially dikaryotic but become diploid via karyogamy. Meiosis takes place at the time of germination. A promycelium is formed that consists of a short hypha (equated to a basidium). In some smuts such as ''[[Mycosarcoma maydis]]'' the nuclei migrate into the promycelium that becomes septate (i.e., divided into cellular compartments separated by cell walls called ''[[Septum#In mycology|septa]]''), and haploid yeast-like conidia/basidiospores sometimes called sporidia, bud off laterally from each cell. In various smuts, the yeast phase may proliferate, or they may fuse, or they may infect plant tissue and become hyphal. In other smuts, such as ''[[Tilletia tritici|Tilletia caries]]'', the elongated haploid basidiospores form apically, often in compatible pairs that fuse centrally resulting in H-shaped [[diaspore]]s which are by then dikaryotic. Dikaryotic conidia may then form. Eventually the host is infected by infectious hyphae. Teliospores form in host tissue. Many variations on these general themes occur.{{cn|date=January 2024}}

Smuts with both a yeast phase and an infectious hyphal state are examples of [[Polymorphism (biology)|dimorphic]] Basidiomycota.<ref name="Prillinger_2002">{{cite journal |last1=Prillinger |first1=Hansjörg |last2=Lopandic |first2=Ksenija |last3=Schweigkofler |first3=Wolfgang |last4=Deak |first4=Robert |display-authors=3 |date=February 2002 |title=Phylogeny and Systematics of the Fungi with Special Reference to the Ascomycota and Basidiomycota |url=https://www.researchgate.net/publication/11274315 |journal=Chemical Immunology |volume=81 |issue=9413 |pages=207–295 |doi=10.1159/000058868 |pmid=12102002 |access-date=2024-03-12}}</ref> In plant parasitic taxa, the saprotrophic phase is normally the yeast while the infectious stage is hyphal. However, there are examples of animal and human parasites where the species are dimorphic but it is the yeast-like state that is infectious.<ref name="Carres_2010">{{cite journal |last1=Rodriguez-Carres |first1=Marianela |last2=Findley |first2=Keisha |last3=Sun |first3=Sheng |last4=Dietrich |first4=Fred S. |last5=Heitman |first5=Joseph |date=2010-03-10 |title=Morphological and Genomic Characterization of Filobasidiella depauperata: A Homothallic Sibling Species of the Pathogenic Cryptococcus Species Complex |journal=PLOS ONE |volume=5 |issue=3  |pages=e9620 |doi=10.1371/journal.pone.0009620 |doi-access=free |pmid=20224779 |pmc=2835752 |bibcode=2010PLoSO...5.9620R }}</ref> The genus ''[[Filobasidiella]]'' forms basidia on hyphae but the main infectious stage is more commonly known by the [[anamorph]]ic yeast name ''[[Cryptococcus (fungus)|Cryptococcus]]'', e.g. ''[[Cryptococcus neoformans]]''<ref name="Erke_1976">{{cite journal |last1=Erke |first1=K. H. |date=October 1976 |title=Light microscopy of basidia, basidiospores, and nuclei in spores and hyphae of Filobasidiella neoformans (Cryptococcus neoformans) |journal=Journal of Bacteriology |volume=128 |issue=1 |pages=445–455 |doi=10.1128/jb.128.1.445-455.1976 |pmid=789347 |pmc=232872 }}</ref> and ''[[Cryptococcus gattii]]''.<ref name="Carres_2010" />

The dimorphic Basidiomycota with yeast stages and the pleiomorphic rusts are examples of fungi with [[anamorph]]s, which are the asexual stages. Some Basidiomycota are only known as anamorphs. Many are called basidiomycetous yeasts, which differentiates them from ascomycetous yeasts in the [[Ascomycota]]. Aside from yeast anamorphs and uredinia, aecia, and pycnidia, some Basidiomycota form other distinctive anamorphs as parts of their life cycles. Examples are ''[[Collybia tuberosa]]''<ref>{{cite web|url=http://www.bio.utk.edu/mycology/ATBI/Photos/12000to12099/12074.jpg |title=Archived copy |access-date=2007-09-13 |url-status=dead |archive-url=https://web.archive.org/web/20071025221100/http://www.bio.utk.edu/mycology/ATBI/Photos/12000to12099/12074.jpg |archive-date=2007-10-25 }}</ref> with its apple-seed-shaped and coloured [[sclerotium]], ''[[Dendrocollybia racemosa]]''<ref>{{Cite web|url=http://www.pnwfungi.org/pdf%20files/manuscripts%20volume%201/pnwf20069.pdf|archiveurl=https://web.archive.org/web/20070927203256/http://www.pnwfungi.org/pdf%20files/manuscripts%20volume%201/pnwf20069.pdf|url-status=dead|title=Microsoft Word – Machnicki revised for pdf final august 24<!-- Bot generated title -->|archive-date=27 September 2007|access-date=3 March 2022}}</ref> with its sclerotium and its ''[[Tilachlidiopsis racemosa]]'' conidia, ''[[Armillaria]]'' with their [[mycelial cord|rhizomorphs]],<ref name="autogenerated1">{{Cite web |url=http://www.bioart.co.uk/lux/mycit.html |title=LUXGENE.COM: The glow-in-the-dark website |access-date=2007-09-13 |archive-date=2007-09-28 |archive-url=https://web.archive.org/web/20070928040055/http://www.bioart.co.uk/lux/mycit.html |url-status=dead }}</ref> ''[[Hohenbuehelia]]''<ref>{{cite web |url=http://www.uoguelph.ca/~gbarron/ZBiodiversity/hohenbue.htm |title=Hohenbue<!-- Bot generated title --> |access-date=2007-04-17 |archive-url=https://web.archive.org/web/20061221075703/http://www.uoguelph.ca/~gbarron/ZBiodiversity/hohenbue.htm |archive-date=2006-12-21 |url-status=dead }}</ref> with their ''[[Nematoctonus]]'' [[nematode]] infectious, state<ref>{{cite web |url=http://www.uoguelph.ca/~gbarron/ZBiodiversity/8knobs.htm |title=8knobs<!-- Bot generated title --> |access-date=2007-04-17 |archive-url=https://web.archive.org/web/20061221075333/http://www.uoguelph.ca/~gbarron/ZBiodiversity/8knobs.htm |archive-date=2006-12-21 |url-status=dead }}</ref> and the coffee leaf parasite, ''[[Mycena citricolor]]'',<ref name="autogenerated1" /> and its ''[[Decapitatus flavidus]]'' [[propagule]]s called gemmae.

==See also==
* [[Forest pathology]]
* [[List of Basidiomycota families]]
* [[Mating in fungi]]

==References==
{{Reflist|30em}}

===Sources===
* {{Cite book | last1 = Kirk | first1 = P. M. | first2 = P. F. | last2 = Cannon | first3 = J. A. | last3 = Stalpers | date = 2008 | title = Dictionary of the Fungi | edition = 10th | publisher = CABI }}

==External links==
{{Commons category|Basidiomycota}}
{{Wikispecies}}
* [http://tolweb.org/Basidiomycota/ Basidiomycota] {{Webarchive|url=https://web.archive.org/web/20201126202611/http://tolweb.org/Basidiomycota |date=2020-11-26 }} at the Tree of Life Web Project

{{Life on Earth}}
{{Eukaryota classification}}
{{Fungi classification}}
{{Taxonbar|from=Q174698}}
{{Authority control}}

[[Category:Basidiomycota| ]]
[[Category:Fungus phyla]]
[[Category:Fungi by classification]]
[[Category:Mycology]]
[[Category:Taxa named by Royall T. Moore]]
[[Category:Taxa described in 1980]]