Biology:Evolution of fungi

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Short description: Origin and diversification of fungi through geologic time
Different species of fungus. Clockwise from top left: Amanita muscaria, a basidiomycete; Sarcoscypha coccinea, an ascomycete; bread covered in mold; a chytrid; an Aspergillus conidiophore.

Fungi diverged from other life around 1.5 billion years ago,[1][2] with the glomaleans branching from the "higher fungi" (dikaryans) at ~570 million years ago, according to DNA analysis. (Schüssler et al., 2001; Tehler et al., 2000)[2] Fungi probably colonized the land during the Cambrian, over 500 million years ago, (Taylor & Osborn, 1996),[2] and possibly 635 million years ago during the Ediacaran,[3][4] but terrestrial fossils only become uncontroversial and common during the Devonian, 400 million years ago.[2]

Early 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.[5]

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.[6]

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.[3]

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.[5]

Fossil record

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[7] has been dismissed on the grounds that it lacks any distinctly fungal features, and is held by many to be contamination;[8] the position of a "probable" Proterozoic fungus is still not established,[8] 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.[9]

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.[10] 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.[11] Compression fossils are studied by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.[12]

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis, in which hyphal branches recombine.[6] 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.[13] 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.[14] 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.[15] 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.[16] Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.[17]

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.[18][19][20] Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants.[2] 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;[21] but these have been dismissed as contamination.[8][22] 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.[2][23][24] At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged,[25] and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).[26]

Lichen-like fossils have been found in the Doushantuo Formation in southern China dating back to 635–551 Ma.[27] Lichens were a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 400 Ma;[28] this date corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert.[29] The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian.[30] 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.[31][32]

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.[33] However, the proportion of fungal spores relative to spores formed by algal species is difficult to assess,[34] the spike did not appear worldwide,[35][36] and in many places it did not fall on the Permian-Triassic boundary.[37]

Approximately 66 million years ago, immediately after the Cretaceous-Tertiary (K-T) extinction that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi, apparently due to the death of most plant and animal species, creating a huge fungal bloom like "a massive compost heap".[38] The lack of K-T extinction in fungal evolution is also supported by molecular data, because phylogenetic comparative analyses of a tree consist of 5,284 mushroom species (Agaricomycetes) didn't show signal for a mass extinction event around the Cretaceous-Tertiary boundary.[39]


References

  1. ↑ Wang, D.Y.C.; Kumar, S.; Hedges, S.B. (1999). "Divergence time estimates for the early history of animal phyla and the origin of plants, animals and fungi". Proceedings of the Royal Society of London B 266 (1415): 163–171. doi:10.1098/rspb.1999.0617. PMID 10097391. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 Brundrett M.C. (2002). "Coevolution of roots and mycorrhizas of land plants". New Phytologist 154 (2): 275–304. doi:10.1046/j.1469-8137.2002.00397.x. PMID 33873429. 
  3. ↑ 3.0 3.1 Gan, Tian; Luo, Taiyi; Pang, Ke; Zhou, Chuanming; Zhou, Guanghong; Wan, Bin; Li, Gang; Yi, Qiru et al. (December 2021). "Cryptic terrestrial fungus-like fossils of the early Ediacaran Period". Nature Communications 12 (1): 641. doi:10.1038/s41467-021-20975-1. PMID 33510166. 
  4. ↑ "Paleontologists Find 635-Million-Year-Old Land Fungus-Like Fossils | Paleontology | Sci-News.com" (in en-US). http://www.sci-news.com/paleontology/ediacaran-fungi-09298.html. 
  5. ↑ 5.0 5.1 Fungi evolution. CK-12 Biology Flexbook. CK12-Foundation. §8.11. https://flexbooks.ck12.org/cbook/ck-12-biology-flexbook-2.0/section/8.11/primary/lesson/fungi-evolution-bio. Retrieved 2020-05-19. 
  6. ↑ 6.0 6.1 Bengtson, Stefan; Rasmussen, Birger; Ivarsson, Magnus; Muhling, Janet; Broman, Curt; Marone, Federica; Stampanoni, Marco; Bekker, Andrey (June 2017). "Fungus-like mycelial fossils in 2.4-billion-year-old vesicular basalt". Nature Ecology & Evolution 1 (6): 0141. doi:10.1038/s41559-017-0141. PMID 28812648. 
  7. ↑ Redecker, D.; Kodner, R.; Graham, L.E. (2000). "Glomalean Fungi from the Ordovician". Science 289 (5486): 1920–1. doi:10.1126/science.289.5486.1920. PMID 10988069. Bibcode: 2000Sci...289.1920R. 
  8. ↑ 8.0 8.1 8.2 Butterfield, N.J. (2005). "Probable Proterozoic fungi". Paleobiology 31 (1): 165–182. doi:10.1666/0094-8373(2005)031<0165:PPF>2.0.CO;2. 
  9. ↑ Miller, A.J. (2004). "A Revised Morphology of Cloudina with Ecological and Phylogenetic Implications". http://ajm.pioneeringprojects.org/files/CloudinaPaper_Final.pdf. 
  10. ↑ Assembling the tree of life. Oxford (Oxfordshire): Oxford University Press. 2004. p. 187. ISBN 978-0-19-517234-8. 
  11. ↑ Taylor and Taylor, p. 19.
  12. ↑ Taylor and Taylor, pp. 7–12.
  13. ↑ Lücking, Robert; Huhndorf, Sabine; Pfister, Donald H.; Plata, Eimy Rivas; Lumbsch, H. Thorsten (November 2009). "Fungi evolved right on track". Mycologia 101 (6): 810–822. doi:10.3852/09-016. PMID 19927746. http://nrs.harvard.edu/urn-3:HUL.InstRepos:14168857. 
  14. ↑ James, Timothy Y.; Kauff, Frank; Schoch, Conrad L.; Matheny, P. Brandon; Hofstetter, Valérie; Cox, Cymon J.; Celio, Gail; Gueidan, Cécile et al. (October 2006). "Reconstructing the early evolution of Fungi using a six-gene phylogeny". Nature 443 (7113): 818–822. doi:10.1038/nature05110. PMID 17051209. Bibcode: 2006Natur.443..818J. 
  15. ↑ "Loss of the flagellum happened only once in the fungal lineage: phylogenetic structure of Kingdom Fungi inferred from RNA polymerase II subunit genes". BMC Evolutionary Biology 6 (1): 74. 2006. doi:10.1186/1471-2148-6-74. PMID 17010206. 
  16. ↑ Taylor and Taylor, pp. 84–94 and 106–107.
  17. ↑ Schoch, Conrad L.; Sung, Gi-Ho; López-Giráldez, Francesc; Townsend, Jeffrey P.; Miadlikowska, Jolanta; Hofstetter, Valérie; Robbertse, Barbara; Matheny, P. Brandon et al. (1 April 2009). "The Ascomycota Tree of Life: A Phylum-wide Phylogeny Clarifies the Origin and Evolution of Fundamental Reproductive and Ecological Traits". Systematic Biology 58 (2): 224–239. doi:10.1093/sysbio/syp020. PMID 20525580. 
  18. ↑ Zimmer, Carl (22 May 2019). "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.". The New York Times. https://www.nytimes.com/2019/05/22/science/fungi-fossils-plants.html. 
  19. ↑ Loron, Corentin C.; François, Camille; Rainbird, Robert H.; Turner, Elizabeth C.; Borensztajn, Stephan; Javaux, Emmanuelle J. (June 2019). "Early fungi from the Proterozoic era in Arctic Canada". Nature 570 (7760): 232–235. doi:10.1038/s41586-019-1217-0. PMID 31118507. Bibcode: 2019Natur.570..232L. 
  20. ↑ Timmer, John (22 May 2019). "Billion-year-old fossils may be early fungus". https://arstechnica.com/science/2019/05/billion-year-old-fossils-may-be-early-fungus/. 
  21. ↑ Redecker D, Kodner R, Graham LE.; Kodner; Graham (2000). "Glomalean fungi from the Ordovician". Science 289 (5486): 1920–21. doi:10.1126/science.289.5486.1920. PMID 10988069. Bibcode: 2000Sci...289.1920R. 
  22. ↑ Smith, Martin R. (2016). "Cord-forming Palaeozoic fungi in terrestrial assemblages". Botanical Journal of the Linnean Society 180 (4): 452–460. doi:10.1111/boj.12389. 
  23. ↑ "The distribution and interactions of some Paleozoic fungi". Review of Palaeobotany and Palynology 95 (1–4): 83–94. 1996. doi:10.1016/S0034-6667(96)00029-2. 
  24. ↑ "Acaulosporoid glomeromycotan spores with a germination shield from the 400-million-year-old Rhynie chert". Mycological Progress 8 (1): 9–18. 2009. doi:10.1007/s11557-008-0573-1. 
  25. ↑ "Dating divergences in the Fungal Tree of Life: review and new analyses". Mycologia 98 (6): 838–49. 2006. doi:10.3852/mycologia.98.6.838. PMID 17486961. https://zenodo.org/record/896945. 
  26. ↑ "Fungi. Eumycota: mushrooms, sac fungi, yeast, molds, rusts, smuts, etc.". Tree of Life Web Project. 2009. http://tolweb.org/Fungi/2377. 
  27. ↑ Yuan X, Xiao S, Taylor TN.; Xiao; Taylor (2005). "Lichen-like symbiosis 600 million years ago". Science 308 (5724): 1017–20. doi:10.1126/science.1111347. PMID 15890881. Bibcode: 2005Sci...308.1017Y. 
  28. ↑ "The most ancient terrestrial lichen Winfrenatia reticulata: A new find and new interpretation". Paleontological Journal 43 (1): 107–14. 2009. doi:10.1134/S0031030109010110. 
  29. ↑ "Perithecial Ascomycetes from the 400 million year old Rhynie chert: an example of ancestral polymorphism". Mycologia 97 (1): 269–85. 2005. doi:10.3852/mycologia.97.1.269. PMID 16389979. 
  30. ↑ Dennis RL. (1970). "A Middle Pennsylvanian basidiomycete mycelium with clamp connections". Mycologia 62 (3): 578–84. doi:10.2307/3757529. 
  31. ↑ Hibbett DS, Grimaldi D, Donoghue MJ.; Grimaldi; Donoghue (1995). "Cretaceous mushrooms in amber". Nature 377 (6549): 487. doi:10.1038/377487a0. Bibcode: 1995Natur.377..487H. 
  32. ↑ "Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of homobasidiomycetes". American Journal of Botany 84 (7): 981–91. 1997. doi:10.2307/2446289. PMID 21708653. 
  33. ↑ Eshet Y, Rampino MR, Visscher H.; Rampino; Visscher (1995). "Fungal event and palynological record of ecological crisis and recovery across the Permian-Triassic boundary". Geology 23 (1): 967–70. doi:10.1130/0091-7613(1995)023<0967:FEAPRO>2.3.CO;2. Bibcode: 1995Geo....23..967E. 
  34. ↑ "A revision of Reduviasporonites Wilson 1962: description, illustration, comparison and biological affinities". Palynology 26 (1): 35–58. 2002. doi:10.2113/0260035. 
  35. ↑ "Permian-Triassic transition in Spain: a multidisciplinary approach". Palaeogeography, Palaeoclimatology, Palaeoecology 229 (1–2): 1–2. 2005. doi:10.1016/j.palaeo.2005.06.028. 
  36. ↑ Looy CV, Twitchett RJ, Dilcher DL, Van Konijnenburg-van Cittert JHA, Visscher H.; Twitchett; Dilcher; Van Konijnenburg-Van Cittert; Visscher (2005). "Life in the end-Permian dead zone". Proceedings of the National Academy of Sciences USA 98 (14): 7879–83. doi:10.1073/pnas.131218098. PMID 11427710. Bibcode: 2001PNAS...98.7879L. "See image 2". 
  37. ↑ Ward PD, Botha J, Buick R, De Kock MO, Erwin DH, Garrison GH, Kirschvink JL, Smith R.; Botha; Buick; De Kock; Erwin; Garrison; Kirschvink; Smith (2005). "Abrupt and gradual extinction among late Permian land vertebrates in the Karoo Basin, South Africa". Science 307 (5710): 709–14. doi:10.1126/science.1107068. PMID 15661973. Bibcode: 2005Sci...307..709W. 
  38. ↑ 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.
  39. ↑ Varga, Torda; Krizsán, Krisztina; Földi, Csenge; Dima, Bálint; Sánchez-García, Marisol; Sánchez-Ramírez, Santiago; Szöllősi, Gergely J.; Szarkándi, János G. et al. (April 2019). "Megaphylogeny resolves global patterns of mushroom evolution". Nature Ecology & Evolution 3 (4): 668–678. doi:10.1038/s41559-019-0834-1. PMID 30886374. 



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