Evolution of fungi#Ongoing evolution

Evidence from DNA analysis suggests that all fungi are descended from a most recent common ancestor that lived at least 1.2 to 1.5 billion years ago. It is probable that these earliest fungi lived in water, and had flagella.

However, a 2.4-billion-year-old basalt from the Palaeoproterozoic Ongeluk Formation in South Africa containing filamentous fossils in vescicles and fractures, that form mycelium-like structures may push back the origin of the Kingdom over one billion years before.{{cite journal |last1=Bengtson |first1=Stefan |last2=Rasmussen |first2=Birger |last3=Ivarsson |first3=Magnus |last4=Muhling |first4=Janet |last5=Broman |first5=Curt |last6=Marone |first6=Federica |last7=Stampanoni |first7=Marco |last8=Bekker |first8=Andrey |title=Fungus-like mycelial fossils in 2.4-billion-year-old vesicular basalt |journal=Nature Ecology & Evolution |date=June 2017 |volume=1 |issue=6 |pages=0141 |doi=10.1038/s41559-017-0141 |pmid=28812648 |hdl=20.500.11937/67718 |s2cid=25586788 |hdl-access=free }}

The earliest terrestrial fungus fossils, or at least fungus-like fossils, have been found in South China from around 635 million years ago. The researchers who reported on these fossils suggested that these fungus-like organisms may have played a role in oxygenating Earth's atmosphere in the aftermath of the Cryogenian glaciations.

About 250 million years ago fungi became abundant in many areas, based on the fossil record, and could even have been the dominant form of life on the earth at that time.{{cite book |title=Fungi evolution |series=CK-12 Biology Flexbook |at=§8.11 |publisher=CK12-Foundation |via=flexbooks.ck12.org |url=https://flexbooks.ck12.org/cbook/ck-12-biology-flexbook-2.0/section/8.11/primary/lesson/fungi-evolution-bio |access-date=2020-05-19}}

{{See also|Paleomycology}}

A rich diversity of fungi is known from the lower Devonian Rhynie chert; an earlier record is absent. Since fungi do not biomineralise, they do not readily enter the fossil record; there are only three claims of early fungi. One from the Ordovician{{cite journal

| author1 = Redecker, D.

| author2 = Kodner, R.

| author3 = Graham, L.E.

| s2cid = 43553633

| year = 2000

| title = Glomalean Fungi from the Ordovician

| journal = Science

| volume = 289

| issue = 5486

| pages = 1920–1

| doi = 10.1126/science.289.5486.1920

| pmid = 10988069

|bibcode = 2000Sci...289.1920R}} has been dismissed on the grounds that it lacks any distinctly fungal features, and is held by many to be contamination;{{cite journal

| author = Butterfield, N.J.

| year = 2005

| title = Probable Proterozoic fungi

| journal = Paleobiology

| volume = 31

| issue = 1

| pages = 165–182

| doi = 10.1666/0094-8373(2005)031<0165:PPF>2.0.CO;2

| bibcode = 2005Pbio...31..165B

| s2cid = 86332371

}} the position of a "probable" Proterozoic fungus is still not established, and it may represent a stem group fungus. There is also a case for a fungal affinity for the enigmatic microfossil Ornatifilum. Since the fungi form a sister group to the animals, the two lineages must have diverged before the first animal lineages, which are known from fossils as early as the Ediacaran.{{Cite web

| author = Miller, A.J.

| year = 2004

| title = A Revised Morphology of Cloudina with Ecological and Phylogenetic Implications

| url = http://ajm.pioneeringprojects.org/files/CloudinaPaper_Final.pdf

| access-date = 2007-04-24

}}

In contrast to plants and animals, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal fruiting bodies, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi.{{cite book|vauthors=Donoghue MJ, Cracraft J |title=Assembling the tree of life |publisher=Oxford University Press |location=Oxford (Oxfordshire)|year=2004 |page=187 |isbn=978-0-19-517234-8}} Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy.Taylor and Taylor, p. 19. Compression fossils are studied by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.Taylor and Taylor, pp. 7–12.

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some {{Ma|2400}} (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis, in which hyphal branches recombine. Other recent studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups.{{cite journal |last1=Lücking |first1=Robert |last2=Huhndorf |first2=Sabine |last3=Pfister |first3=Donald H. |last4=Plata |first4=Eimy Rivas |last5=Lumbsch |first5=H. Thorsten |title=Fungi evolved right on track |journal=Mycologia |date=November 2009 |volume=101 |issue=6 |pages=810–822 |doi=10.3852/09-016 |pmid=19927746 |s2cid=6689439 |url=http://nrs.harvard.edu/urn-3:HUL.InstRepos:14168857 |url-access=subscription }} For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant Chytrids in having flagellum-bearing spores.{{cite journal |last1=James |first1=Timothy Y. |last2=Kauff |first2=Frank |last3=Schoch |first3=Conrad L. |last4=Matheny |first4=P. Brandon |last5=Hofstetter |first5=Valérie |last6=Cox |first6=Cymon J. |last7=Celio |first7=Gail |last8=Gueidan |first8=Cécile |last9=Fraker |first9=Emily |last10=Miadlikowska |first10=Jolanta |last11=Lumbsch |first11=H. Thorsten |last12=Rauhut |first12=Alexandra |last13=Reeb |first13=Valérie |last14=Arnold |first14=A. Elizabeth |last15=Amtoft |first15=Anja |last16=Stajich |first16=Jason E. |last17=Hosaka |first17=Kentaro |last18=Sung |first18=Gi-Ho |last19=Johnson |first19=Desiree |last20=O’Rourke |first20=Ben |last21=Crockett |first21=Michael |last22=Binder |first22=Manfred |last23=Curtis |first23=Judd M. |last24=Slot |first24=Jason C. |last25=Wang |first25=Zheng |last26=Wilson |first26=Andrew W. |last27=Schüßler |first27=Arthur |last28=Longcore |first28=Joyce E. |last29=O’Donnell |first29=Kerry |last30=Mozley-Standridge |first30=Sharon |last31=Porter |first31=David |last32=Letcher |first32=Peter M. |last33=Powell |first33=Martha J. |last34=Taylor |first34=John W. |last35=White |first35=Merlin M. |last36=Griffith |first36=Gareth W. |last37=Davies |first37=David R. |last38=Humber |first38=Richard A. |last39=Morton |first39=Joseph B. |last40=Sugiyama |first40=Junta |last41=Rossman |first41=Amy Y. |last42=Rogers |first42=Jack D. |last43=Pfister |first43=Don H. |last44=Hewitt |first44=David |last45=Hansen |first45=Karen |last46=Hambleton |first46=Sarah |last47=Shoemaker |first47=Robert A. |last48=Kohlmeyer |first48=Jan |last49=Volkmann-Kohlmeyer |first49=Brigitte |last50=Spotts |first50=Robert A. |last51=Serdani |first51=Maryna |last52=Crous |first52=Pedro W. |last53=Hughes |first53=Karen W. |last54=Matsuura |first54=Kenji |last55=Langer |first55=Ewald |last56=Langer |first56=Gitta |last57=Untereiner |first57=Wendy A. |last58=Lücking |first58=Robert |last59=Büdel |first59=Burkhard |last60=Geiser |first60=David M. |last61=Aptroot |first61=André |last62=Diederich |first62=Paul |last63=Schmitt |first63=Imke |last64=Schultz |first64=Matthias |last65=Yahr |first65=Rebecca |last66=Hibbett |first66=David S. |last67=Lutzoni |first67=François |last68=McLaughlin |first68=David J. |last69=Spatafora |first69=Joseph W. |last70=Vilgalys |first70=Rytas |title=Reconstructing the early evolution of Fungi using a six-gene phylogeny |journal=Nature |date=October 2006 |volume=443 |issue=7113 |pages=818–822 |doi=10.1038/nature05110 |pmid=17051209 |bibcode=2006Natur.443..818J |s2cid=4302864 }} Phylogenetic analyses suggest that the flagellum was lost early in the evolutionary history of the fungi, and consequently, the majority of fungal species lack a flagellum.{{cite journal |vauthors=Liu YJ, Hodson MC, Hall BD |title=Loss of the flagellum happened only once in the fungal lineage: phylogenetic structure of Kingdom Fungi inferred from RNA polymerase II subunit genes |journal=BMC Evolutionary Biology |year=2006 |volume=6 |issue=1 |page=74 |doi=10.1186/1471-2148-6-74 |pmid=17010206 |pmc=1599754 |doi-access=free }} The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization.Taylor and Taylor, pp. 84–94 and 106–107. Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.{{cite journal |last1=Schoch |first1=Conrad L. |last2=Sung |first2=Gi-Ho |last3=López-Giráldez |first3=Francesc |last4=Townsend |first4=Jeffrey P. |last5=Miadlikowska |first5=Jolanta |last6=Hofstetter |first6=Valérie |last7=Robbertse |first7=Barbara |last8=Matheny |first8=P. Brandon |last9=Kauff |first9=Frank |last10=Wang |first10=Zheng |last11=Gueidan |first11=Cécile |last12=Andrie |first12=Rachael M. |last13=Trippe |first13=Kristin |last14=Ciufetti |first14=Linda M. |last15=Wynns |first15=Anja |last16=Fraker |first16=Emily |last17=Hodkinson |first17=Brendan P. |last18=Bonito |first18=Gregory |last19=Groenewald |first19=Johannes Z. |last20=Arzanlou |first20=Mahdi |last21=Sybren de Hoog |first21=G. |last22=Crous |first22=Pedro W. |last23=Hewitt |first23=David |last24=Pfister |first24=Donald H. |last25=Peterson |first25=Kristin |last26=Gryzenhout |first26=Marieka |last27=Wingfield |first27=Michael J. |last28=Aptroot |first28=André |last29=Suh |first29=Sung-Oui |last30=Blackwell |first30=Meredith |last31=Hillis |first31=David M. |last32=Griffith |first32=Gareth W. |last33=Castlebury |first33=Lisa A. |last34=Rossman |first34=Amy Y. |last35=Lumbsch |first35=H. Thorsten |last36=Lücking |first36=Robert |last37=Büdel |first37=Burkhard |last38=Rauhut |first38=Alexandra |last39=Diederich |first39=Paul |last40=Ertz |first40=Damien |last41=Geiser |first41=David M. |last42=Hosaka |first42=Kentaro |last43=Inderbitzin |first43=Patrik |last44=Kohlmeyer |first44=Jan |last45=Volkmann-Kohlmeyer |first45=Brigitte |last46=Mostert |first46=Lizel |last47=O'Donnell |first47=Kerry |last48=Sipman |first48=Harrie |last49=Rogers |first49=Jack D. |last50=Shoemaker |first50=Robert A. |last51=Sugiyama |first51=Junta |last52=Summerbell |first52=Richard C. |last53=Untereiner |first53=Wendy |last54=Johnston |first54=Peter R. |last55=Stenroos |first55=Soili |last56=Zuccaro |first56=Alga |last57=Dyer |first57=Paul S. |last58=Crittenden |first58=Peter D. |last59=Cole |first59=Mariette S. |last60=Hansen |first60=Karen |last61=Trappe |first61=James M. |last62=Yahr |first62=Rebecca |last63=Lutzoni |first63=François |last64=Spatafora |first64=Joseph W. |title=The Ascomycota Tree of Life: A Phylum-wide Phylogeny Clarifies the Origin and Evolution of Fundamental Reproductive and Ecological Traits |journal=Systematic Biology |date=1 April 2009 |volume=58 |issue=2 |pages=224–239 |doi=10.1093/sysbio/syp020 |pmid=20525580 |doi-access=free }}

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land.{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=How Did Life Arrive on Land? A Billion-Year-Old Fungus May Hold Clues - A cache of microscopic fossils from the Arctic hints that fungi reached land long before plants. |url=https://www.nytimes.com/2019/05/22/science/fungi-fossils-plants.html |date=22 May 2019 |work=The New York Times |access-date=23 May 2019 }}{{cite journal |last1=Loron |first1=Corentin C. |last2=François |first2=Camille |last3=Rainbird |first3=Robert H. |last4=Turner |first4=Elizabeth C. |last5=Borensztajn |first5=Stephan |last6=Javaux |first6=Emmanuelle J. |title=Early fungi from the Proterozoic era in Arctic Canada |journal=Nature |date=June 2019 |volume=570 |issue=7760 |pages=232–235 |doi=10.1038/s41586-019-1217-0 |pmid=31118507 |bibcode=2019Natur.570..232L |s2cid=162180486 }}{{cite web |last=Timmer |first=John |title=Billion-year-old fossils may be early fungus |website=Ars Technica |date=22 May 2019 |url=https://arstechnica.com/science/2019/05/billion-year-old-fossils-may-be-early-fungus/ |access-date=23 May 2019}} Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants;{{cite journal |author=Redecker D, Kodner R, Graham LE. |title=Glomalean fungi from the Ordovician |journal=Science |volume=289 |issue=5486 |pages=1920–21 |year=2000 |pmid=10988069 |doi= 10.1126/science.289.5486.1920|bibcode = 2000Sci...289.1920R |last2=Kodner |last3=Graham |s2cid=43553633 }} but these have been dismissed as contamination.{{Cite journal | doi = 10.1111/boj.12389| title = Cord-forming Palaeozoic fungi in terrestrial assemblages| year = 2016| last1 = Smith| first1 = Martin R.| journal = Botanical Journal of the Linnean Society| volume = 180| issue = 4| pages = 452–460| s2cid = 86883382| doi-access = free}} Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they are abundant in the Rhynie chert, mostly as Zygomycota and Chytridiomycota.{{cite journal |vauthors=Taylor TN, Taylor EL |year= 1996 |title=The distribution and interactions of some Paleozoic fungi |journal=Review of Palaeobotany and Palynology |volume=95 |issue=1–4 |pages=83–94 |doi=10.1016/S0034-6667(96)00029-2 }}{{cite journal |vauthors=Dotzler N, Walker C, Krings M, Hass H, Kerp H, Taylor TN, Agerer R |year=2009|title=Acaulosporoid glomeromycotan spores with a germination shield from the 400-million-year-old Rhynie chert|journal=Mycological Progress|volume=8 |issue=1 |pages=9–18 |doi=10.1007/s11557-008-0573-1|bibcode=2009MycPr...8....9D |hdl=1808/13680|s2cid=1746303|hdl-access=free}} At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged,{{cite journal |vauthors=Taylor JW, Berbee ML |title=Dating divergences in the Fungal Tree of Life: review and new analyses |journal=Mycologia |volume=98 |issue=6 |pages=838–49 |year=2006 |pmid=17486961 |doi= 10.3852/mycologia.98.6.838|url=https://zenodo.org/record/896945}} and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).{{cite web |url=http://tolweb.org/Fungi/2377 |title=Fungi. Eumycota: mushrooms, sac fungi, yeast, molds, rusts, smuts, etc. |vauthors=Blackwell M, Vilgalys R, James TY, Taylor JW |publisher=Tree of Life Web Project|year=2009 |access-date=2009-04-25}}

Lichen-like fossils have been found in the Doushantuo Formation in southern China dating back to 635–551 Ma.{{cite journal |author=Yuan X, Xiao S, Taylor TN. |title=Lichen-like symbiosis 600 million years ago |journal=Science |volume=308 |issue=5724 |pages=1017–20 |year=2005 |pmid=15890881 |doi=10.1126/science.1111347 |bibcode = 2005Sci...308.1017Y |last2=Xiao |last3=Taylor |s2cid=27083645 }} Lichens were a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 400 Ma;{{cite journal |vauthors=Karatygin IV, Snigirevskaya NS, Vikulin SV |year=2009 |title=The most ancient terrestrial lichen Winfrenatia reticulata: A new find and new interpretation |journal=Paleontological Journal |volume=43 |issue=1 |pages=107–14|doi=10.1134/S0031030109010110|bibcode=2009PalJ...43..107K |s2cid=85262818 }} this date corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert.{{cite journal |vauthors=Taylor TN, Hass H, Kerp H, Krings M, Hanlin RT |title=Perithecial Ascomycetes from the 400 million year old Rhynie chert: an example of ancestral polymorphism |journal=Mycologia |volume=97 |issue=1 |pages=269–85 |year=2005 |pmid=16389979 |doi= 10.3852/mycologia.97.1.269|hdl=1808/16786 |hdl-access=free }} The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian.{{cite journal |author= Dennis RL.|year=1970 |title=A Middle Pennsylvanian basidiomycete mycelium with clamp connections |jstor= 3757529 |journal=Mycologia |volume=62 |issue= 3|pages=578–84 |doi= 10.2307/3757529}} Rare in the fossil record are the homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius legletti) appeared during the mid-Cretaceous, 90 Ma.{{cite journal |author=Hibbett DS, Grimaldi D, Donoghue MJ. |year=1995|title=Cretaceous mushrooms in amber |journal=Nature |volume=377 |issue=6549|page=487 |doi=10.1038/377487a0 |bibcode=1995Natur.377..487H|last2=Grimaldi |last3=Donoghue |doi-access=free }}{{cite journal |vauthors=Hibbett DS, Grimaldi D, Donoghue MJ |s2cid=22011469 |year=1997 |title=Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of homobasidiomycetes |jstor= 2446289|journal=American Journal of Botany |volume=84 |issue= 7|pages=981–91 |doi= 10.2307/2446289|pmid=21708653 |doi-access=free }}

Some time after the Permian-Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period.{{cite journal |author=Eshet Y, Rampino MR, Visscher H.|year=1995|title=Fungal event and palynological record of ecological crisis and recovery across the Permian-Triassic boundary |journal=Geology |volume=23 |issue=1 |pages=967–70 |doi=10.1130/0091-7613(1995)023<0967:FEAPRO>2.3.CO;2 |bibcode=1995Geo....23..967E|last2=Rampino|last3=Visscher|s2cid=58937537}} However, the proportion of fungal spores relative to spores formed by algal species is difficult to assess,{{cite journal |vauthors=Foster CB, Stephenson MH, Marshall C, Logan GA, Greenwood PF | year = 2002 | title = A revision of Reduviasporonites Wilson 1962: description, illustration, comparison and biological affinities| journal = Palynology | volume = 26 | issue = 1 | pages = 35–58 | doi = 10.2113/0260035 | bibcode = 2002Paly...26...35F }} the spike did not appear worldwide,{{cite journal | title=Permian-Triassic transition in Spain: a multidisciplinary approach | journal=Palaeogeography, Palaeoclimatology, Palaeoecology |

volume=229 | issue=1–2 | year=2005 | pages=1–2 | doi=10.1016/j.palaeo.2005.06.028 |vauthors=López-Gómez J, Taylor EL }}{{cite journal|author = Looy CV, Twitchett RJ, Dilcher DL, Van Konijnenburg-van Cittert JHA, Visscher H.|year=2005|title = Life in the end-Permian dead zone |journal = Proceedings of the National Academy of Sciences USA|volume = 98 |issue = 14| pages = 7879–83 |doi = 10.1073/pnas.131218098| quote = See image 2 | pmid = 11427710|pmc = 35436|bibcode = 2001PNAS...98.7879L |last2 = Twitchett|last3 = Dilcher|last4 = Van Konijnenburg-Van Cittert|last5 = Visscher|doi-access=free}} and in many places it did not fall on the Permian-Triassic boundary.{{cite journal|author=Ward PD, Botha J, Buick R, De Kock MO, Erwin DH, Garrison GH, Kirschvink JL, Smith R.|year=2005|title=Abrupt and gradual extinction among late Permian land vertebrates in the Karoo Basin, South Africa|journal=Science|volume=307|issue=5710|pages=709–14|doi=10.1126/science.1107068|pmid=15661973|bibcode = 2005Sci...307..709W |last2=Botha|last3=Buick|last4=De Kock|last5=Erwin|last6=Garrison|last7=Kirschvink|last8=Smith|citeseerx=10.1.1.503.2065|s2cid=46198018}}

Approximately 66 million years ago, immediately after the Cretaceous-Tertiary (K-T) extinction, there was a dramatic increase in evidence of fungi. Fungi appear to have had the chance to flourish due to the extinction of most plant and animal species, and the resultant fungal bloom has been described as like "a massive compost heap".[http://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1002808 Fungi and the Rise of Mammals]
That ecological calamity was accompanied by massive deforestation, an event followed by a fungal bloom, as the earth became a massive compost. The lack of K-T extinction in fungal evolution is also supported by molecular data. Phylogenetic comparative analyses of a tree consisting of 5,284 agaricomycete species do not show signal for a mass extinction event around the Cretaceous-Tertiary boundary.{{cite journal |last1=Varga |first1=Torda |last2=Krizsán |first2=Krisztina |last3=Földi |first3=Csenge |last4=Dima |first4=Bálint |last5=Sánchez-García |first5=Marisol |last6=Sánchez-Ramírez |first6=Santiago |last7=Szöllősi |first7=Gergely J. |last8=Szarkándi |first8=János G. |last9=Papp |first9=Viktor |last10=Albert |first10=László |last11=Andreopoulos |first11=William |last12=Angelini |first12=Claudio |last13=Antonín |first13=Vladimír |last14=Barry |first14=Kerrie W. |last15=Bougher |first15=Neale L. |last16=Buchanan |first16=Peter |last17=Buyck |first17=Bart |last18=Bense |first18=Viktória |last19=Catcheside |first19=Pam |last20=Chovatia |first20=Mansi |last21=Cooper |first21=Jerry |last22=Dämon |first22=Wolfgang |last23=Desjardin |first23=Dennis |last24=Finy |first24=Péter |last25=Geml |first25=József |last26=Haridas |first26=Sajeet |last27=Hughes |first27=Karen |last28=Justo |first28=Alfredo |last29=Karasiński |first29=Dariusz |last30=Kautmanova |first30=Ivona |last31=Kiss |first31=Brigitta |last32=Kocsubé |first32=Sándor |last33=Kotiranta |first33=Heikki |last34=LaButti |first34=Kurt M. |last35=Lechner |first35=Bernardo E. |last36=Liimatainen |first36=Kare |last37=Lipzen |first37=Anna |last38=Lukács |first38=Zoltán |last39=Mihaltcheva |first39=Sirma |last40=Morgado |first40=Louis N. |last41=Niskanen |first41=Tuula |last42=Noordeloos |first42=Machiel E. |last43=Ohm |first43=Robin A. |last44=Ortiz-Santana |first44=Beatriz |last45=Ovrebo |first45=Clark |last46=Rácz |first46=Nikolett |last47=Riley |first47=Robert |last48=Savchenko |first48=Anton |last49=Shiryaev |first49=Anton |last50=Soop |first50=Karl |last51=Spirin |first51=Viacheslav |last52=Szebenyi |first52=Csilla |last53=Tomšovský |first53=Michal |last54=Tulloss |first54=Rodham E. |last55=Uehling |first55=Jessie |last56=Grigoriev |first56=Igor V. |last57=Vágvölgyi |first57=Csaba |last58=Papp |first58=Tamás |last59=Martin |first59=Francis M. |last60=Miettinen |first60=Otto |last61=Hibbett |first61=David S. |last62=Nagy |first62=László G. |title=Megaphylogeny resolves global patterns of mushroom evolution |journal=Nature Ecology & Evolution |date=April 2019 |volume=3 |issue=4 |pages=668–678 |doi=10.1038/s41559-019-0834-1 |pmid=30886374 |pmc=6443077 |bibcode=2019NatEE...3..668V }}

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