Reconstruction of Fractofusus misrai, copyright Oleg Kuznetsov.

Belongs within: Amorphea.
Contains: Teretosporea, Choanoflagellata, Metazoa, Rotosphaerida, Opisthosporidia, Neocallimastigaceae, Chytridiomycota, Blastocladiales, Zoopagomycota, Mucoromycota, Glomeromycota, Ascomycota, Pucciniomycotina, Orthomycotina.

The Opisthokonta are the major clade of eukaryotes uniting the total groups of Metazoa (animals) and Fungi, and their immediate unicellular allies. They are ancestrally characterised by cells bearing a single posterior cilium without mastigonemes in at least one stage of the life cycle stage, though this has been secondarily lost in some lineages. Cells bear a pair of kinetosomes or centrioles, sometimes modified, and flat (rarely tubular) mitochondrial cristae in the unicellular stage (Adl et al. 2019). The basal clades Holozoa, comprising all taxa closer to animals than fungi, and the inverse clade Nucletmycea are established by molecular data.

The Fungi are often filamentous heterotrophic organisms with vegetative and/or spore cell walls of chitin and β-glucan and lacking phagocytosis (Cavalier-Smith 1998). Basal members of the fungi retain ciliate zoospores, but many lineages (including all macroscopic fungi) have lost cilia in all stages of the life cycle. Earlier authors divided fungi other than the Ascomycota and Basidiomycota between Chytridiomycota (ciliate at some stage in the life cycle) and Zygomycota (lacking cilia at all times), but these are not phylogenetically coherent groups in these senses, with the loss of cilia having occurred multiple times (James et al. 2006). The Ascomycota and Basidiomycota form a clade Dikarya characterised by the presence of a dikaryotic stage in the life cycle (Adl et al. 2019).

Giants of the Silurian
Published 11 April 2008
Reconstruction of Prototaxites as columnar perrenial fungus from Hueber (2001), painted by Mary Parrish.

Compared to life in the water, life on land got off to a decidedly slow start. The seas had been swarming with life for hundreds of millions of years before the first tentative colonisation of the previously barren continents by small algae and plants in the Ordovician about 470 million years ago. Even then, it wasn’t until about the middle part of the Silurian, about 430 million years ago (give or take) that the first vascular plants and anything bigger than a moss put in an appearance. These early plants were still small by modern standards—one of the larger forms, Cooksonia, was practically a giant at heights of nearly a metre. The first really large land organism made its imposing appearance in the late Silurian.

Reaching heights of up to eight metres, Prototaxites would have loomed over anything else around at the time—something that is made pretty clear in the reconstruction above from Hueber (2001) (obtained via Hans Steur’s palaeobotany page, which has a far better analysis of Prototaxites than I’m about to give you). Prototaxites is known from fossils reminiscent of the trunk of a tree, and was originally described as such in 1859. However, the internal structure of Prototaxites is very different from that of any tree. Rather than the vascular cells of a tree, Prototaxites is composed of long filamentous tubes—large unbranched tubes that probably supplied structural support, and much smaller multi-branched tubes that ran among and around the skeletal filaments and probably held the structure together. This structure was compared to the filamentous structure of a number of modern algae, and for many years Prototaxites was considered a giant alga, probably related to the modern brown algae.

However, all known algae are aquatic whereas Prototaxites was found in association with terrestrial organisms and was undoubtedly terrestrial itself. In cross-section Prototaxites also lacks the orderliness of structure found in brown algae, with filaments arranged randomly in the trunk with no clear division between pith and cortex. Hueber (2001) therefore made the suggestion that instead of being an alga, Prototaxites might be a gigantic fungus. He compared it to the living perennial bracket fungi, which have a rigid structure and can also reach notable sizes (up to a metre in diameter) through cumulative growth over many years.

The main barrier to accepting Prototaxites as a fungus lies in the size of the filaments (which would then be hyphae). The skeletal filaments reach a diameter of up to 50μm while modern fungi will rarely be more than 10μm. The other major issue, of course, is its size overall. Where did such a large saprobic fungus gain its nutrients from? Despite the plausability of Hueber’s (2001) model of perennial growth, and his comparison with the gigantic size attained by hyphal masses in other modern fungi such as a specimen of Armillaria bulbosa covering an area of 15ha, Selosse (2002) pointed out that the modern taxa live in an environment with a much larger overall biomass. A purely saprobic Prototaxites would have required far more nutrition than could be supplied by the meagre vegetation of the Silurian. Selosse (2002) therefore suggested that, rather than being purely fungal, Prototaxites might have been a photosynthetic lichen-like symbiosis. The large skeletal filaments, he suggested, might still represent an algal form, contained and protected by the surrounding fungal hyphae. The idea of an eight-metre-tall lichen is still pretty amazing, but not incredible when we consider that there would have been relatively little competition in the Silurian compared to the modern environment. Prototaxites may have been a slow and inefficient grower, but there was little to exclude it. Hueber (2001) identified supposed reproductive structures on Prototaxites that he held indicated a position for it among the basidiomycetes, which include the bracket fungi. However, as pointed out by Selosse (2002), the reproductive nature of these structures is unconvincing.

No convincing evidence has been found for branching in Prototaxites. Many authors interpreting Prototaxites as a plant or alga have suggested that the flattened fossil Nematothallus found in the same deposits, which also has a filamentous structure, might represent leaves or leaf-like appendages of Prototaxites. However, the two have never been found directly attached. Graham et al. (2004) noted a similarity between Nematothallus and semi-decayed modern liverworts, and suggested that Nematothallus might belong to the latter group.

Prototaxites was around for about 50 million years, a quite impressive amount of time, but it eventually became extinct during the Devonian. Perhaps as the terrestrial environment increased in complexity, the window of low competition that Prototaxites had occupied closed. More efficient plants supplanted Prototaxites, while increased numbers of herbivores may have eaten down growing hyphae faster than the organism could regrow them. Prototaxites‘ time had come to a close.

Prototaxites: a giant that never was?
Published 10 February 2010

Nearly two years ago, I wrote the above section on Prototaxites, a mysterious fossil of the late Silurian and the earliest truly large terrestrial organism known from the fossil record. I have discussed the possibility that Prototaxites might have represented a giant fungus but a recent publication by Graham et al. (2010a) presents a new alternative interpretation of Prototaxites. If they are correct, the Silurian may never be the same again.

Thalli of the liverwort Marchantia. Photo from here.

In Graham et al.‘s estimation, Prototaxites should not be classed with the fungi but with the liverworts. Liverworts are small, often mosslike plants of moist habitats. Members of one group of liverworts, the thallose liverworts, lack any distinction between leaves and stem but grow as a flattened thallus anchored to the ground by rhizoids (rootlets) on the lower surface. Liverworts are one of the earliest diverging groups of land plants and they or their ancestors would have certainly been part of the Silurian flora. One group of Silurian plant fossils, the nematophytes, possess a microstructure of criscrossing tubular filaments; Graham et al. (2004) demonstrated that this structure was also found in the decaying remains of modern thallose liverworts, as the upper tissue of the thallus rotted away to leave the more resistant rhizoids and connective tissue. The microstructure of Prototaxites is also similar to that of nematophytes, to the extent that some palaeontologists have regarded nematophytes as Prototaxites leaves (this interpretation is not currently supported as nematophytes have never been found actually attached to Prototaxites). But modern liverworts lack strong supporting tissue and would be pushing to reach an inch in height – how could they have produced the eight-metre columns recorded for Prototaxites?

The largest known Prototaxites fossil (at least as of 2001), photographed by Charles Meissner in Saudi Arabia. From Hueber (2001).

A transverse section of Prototaxites shows a ring structure like that found in a tree trunk. Hueber (2001), who interpreted Prototaxites as a perennial fungal fruiting body, felt that this ring structure also resembled tree rings in indicating discontinuous growth by the organism. Graham et al. (2010a) interpret the ring structure differently. They suggest that large mats of thallose liverworts covered the Silurian landscape. These mats could become detached from their substrate by agents such as wind and rain, and start to roll up as they decayed. As they rolled, they would form the large columns that, after being compressed by burial and fossilised, would eventually be identified as Prototaxites.

Reconstruction by Kandis Elliot of Silurian liverwort mats being rolled by wind, gravity and/or water movement to form ‘Prototaxites’. From Graham et al. (2010a).

Under this interpretation of Prototaxites, the fungal hyphal structures identified by Hueber (2001) within Prototaxites sections would be those of fungi growing among the liverwort mats. Boyce et al. (2007) identified significant variations in carbon isotope ratios between Prototaxites individuals as supportive of fungal identification because they suggested heterotrophy (nutrients being obtained from the surrounding environment rather than being produced by the organism itself); however, Graham et al. (2010a) establish that thallose liverworts may grow heterotrophically when conditions encourage it. The liverwort interpretation is also more consistent with the size of most Prototaxites filaments (much larger than found in modern fungi) and also explains the occasional discovery of other land plants embedded in Prototaxites columns – these would have been growing among the mats and become swept up when the mats became rolled, like Silurian Cleopatras.

I find this new interpretation intriguing, if a little difficult to accept outright. Prototaxites is represented by a reasonable number of specimens (I don’t know the actual number, but thirteen species have been named from numerous localities around the world)—were the conditions that would have lead to mat-rolling common enough to have produced that number of fossils? I wonder if it would be worth investigating how Prototaxites specimens compare in abundance to nematophyte specimens and what that might tell us about the likelihood of ‘Prototaxites‘ formation from liverwort mats. Certainly, the only thing that could be more intriguing than the existence of these giant pillars from so early in the earth’s history would be if it turned out that they never existed at all.

Prototaxites revisited
Published 5 November 2012
Reconstruction of Prototaxites by Richard Bizley, used with permission.

Richard Bizley has been kind enough to allow me to reproduce the above painting, which he produced in response to discussion arising from the post above. It shows a ‘forest’ (for want of a better word) of the enigmatic Silurian-Devonian organism Prototaxites reconstructed as a giant fungus. Richard has asked if anyone has any comments to make on the final product. Is this environment plausible? Could Prototaxites have grown in clusters like this, or would nutrient restrictions been such as to prevent such large organisms from persisting in close proximity to each other?

Since I produced my earlier post on the possible re-interpretation of Prototaxites as representing rolled ground-cover mats (Graham et al. 2010a), the proposal has been criticised in print by Boyce and Hotton (2010) and Taylor et al. (2010), and defended by Graham et al. (2010b). Boyce and Hotton regard it as taphonomically implausible that such rolls could form, while Taylor et al. also point out that the major tubes making up Prototaxites are arranged longitudinally down the ‘trunk’, not radiating outwards. Graham et al. have pointed out how they feel this is not incompatible with their liverwort mat hypothesis.

Colour-enhanced cross-section of Prototaxites specimen, from Graham et al. (2010b). Note that the ‘growth rings’ are not regularly concentric.

Prototaxites, it should be pointed out, was just one of a number of Silurian-Devonian organisms called nematophytes. Nematophytes are united by their similar internal structure, composed of hypha-like tubes. However, other nematophytes did not have the gigantic columnar form of Prototaxites: Nematothallus, for instance, was an encrusting lichen-like form, while Nematasketum fossils are only a couple of centimetres in size. Edwards and Axe (2012) have recently published a study on Nematasketum and supported comparisons between nematophytes and fungi. In particular, they compare Nematasketum to root-like anchoring and foraging structures called rhizomorphs produced by some large modern basidiomycetes. Hillier et al. (2008) nominated Prototaxites as potentially connected to root-like casts found in the Anglo-Welsh Old Red Sandstone, but admitted that the grounds for connection were slight.

Systematics of Opisthokonta
Opisthokonta [Choanozoa, Cristidiscoidea, Enteromycetes]
    |    |--FilastereaAS12
    |    |    |--PigoraptorAB19
    |    |    |--Capsaspora Hertel et al. 2002HO09, AS12
    |    |    |    `--C. owczarzakiHO09
    |    |    `--Ministeria Patterson et al. 1993 [Ministeriida]AS12
    |    |         |--M. marisolaC-SC03
    |    |         `--M. vibransC-SC03
    |    `--+--TeretosporeaAB19
    |       `--Choanozoa [Apoikozoa]AB19
    |            |--ChoanoflagellataAB19
    |            `--+--MetazoaBJ17
    |               `--FractofususBJ17
    |                    |--F. andersoni Gehling & Narbonne 2007G14
    |                    `--F. misrai Gehling & Narbonne 2007G14
    `--Nucletmycea [Holomycota]AS12
         |  i. s.: Amoeboradix [Sanchytriaceae]AB19
         `--Fungi (see below for synonymy)C-SCL15
              |  i. s.: Dothidiella derridisDP72
              |         Ramilina americanaAB-E04
              |         Ostracoblabe Bornet & Flahault 1891KC01
              |           `--O. implexaZB10
              |         Telohannia solenopsaeES12
              |         Myrmicinosporidium durum Hölldobler 1933ES12
              |         PaleomastigomycetesKC01
              |           |--Milleromyces Taylor, Hass & Remy 1993 [Paleochytridales]KC01
              |           |    `--M. rhyniensisT93
              |           |--Krispiromyces Taylor, Hass & Remy 1993KC01
              |           |    `--K. discoidesT93
              |           `--Lyonomyces Taylor, Hass & Remy 1993KC01
              |                `--L. pyriformisT93
              |         NephridiophagaAB19 [incl. Coelosporidium Mesnil & Marchoux 1897AB19, KC01]
              |         Nematasketum diversiforme Burgess & Edwards 1988EA12
              |         PrototaxitesEA12
              |           |--P. hefteri (Schaarschmidt) Schweitzer 1983 [=Mosellophyton hefteri Schaarschmidt 1974]S00
              |           |--P. loganiEA12
              |           `--P. psygmophylloides Kräusel & Weyland 1930S00
              |         Abeliella Mägdefrau 1937KC01
              |           `--*A. riccioides Mägdefrau 1937H75
              |         Achlyites Mesch. 1902KC01
              |         Acremonites Pia 1927KC01
              |         Actonia Dodge 1935 (n. d.)KC01
              |         Actycus Raf. 1815 (n. d.)KC01
              |         Acumispora Matsush. 1980KC01
              |         Aecidites Debey & Ettingsh. 1859KC01
              |         Agaricites Mesch. 1891KC01
              |         Aldridgea Massee 1892 (n. d.)KC01
              |         Aldrigiella Rick 1934 (n. d.)KC01
              |         Alveolinus Raf. 1815 (n. d.)KC01
              |         Alysphaeria Turpin 1827 (n. d.)KC01
              |         Alytosporium Link 1824 (n. d.)KC01
              |         Ambrosiaemyces Trotter 1934KC01
              |         Amepiospora Locq. & Sal.-Cheb. 1980KC01
              |         Amerostege Theiss. 1916 (n. d.)KC01
              |         Anatolinites Elsik, Ediger & Bati 1990KC01
              |         Annella Srivast. 1976KC01
              |         Anomothallus Stevens 1925 (n. d.)KC01
              |         Anthracomyces Renault 1898KC01
              |         Anulomyces Bydgosz 1932 (n. d.)KC01
              |         Hydrothria venosaKC01
              |         Arbusculites Paradkar 1976 non Murray 1831 (ICZN)KC01
              |         Archagaricon Hancock & Atthey 1869KC01
              |         Archecribraria Locq. 1983KC01
              |         Archeomycelites Bystrov 1959KC01
              |         ArchephomaKC01
              |         Archeplax Locq. 1985KC01
              |         Arctosporidium Thor 1930 (n. d.)KC01
              |         Arthriniites Babajan & Tasl. 1977KC01
              |         Arthroon Renault 1894KC01
              |         Artocreas Berk. & Broome 1875 (n. d.)KC01
              |         Ascochytites Babajan & Tasl. 1973KC01
              |         Ascodesmites Trivedi, Chaturv. & Verma 1973KC01
              |         Ascospermum Schulzer 1863 (n. d.)KC01
              |         Aselmotrichum Corda 1831 (n. d.)KC01
              |         Aspergillites Trivedi & Verma 1969KC01
              |         Asterinites Krassilov 1967KC01
              |         ‘Asterinites’ Doub. & Pons 1973KC01
              |         Asteroides Puntoni & Léon 1940 (n. d.)KC01
              |         Asteronema Trevis. 1845 (n. d.) [=Asterothrix Kütz. 1843 non Cass. 1827]KC01
              |         Asterothyrites Cookson 1947KC01
              |         Asyregraamspora Locq. & Sal.-Cheb. 1980KC01
              |         Atractodorus Klotzsch 1832 (n. d.)KC01
              |         Bactrodesmiites Babajan & Tasl. 1977KC01
              |         Basidiosporites Elsik 1968KC01
              |         Batistamnus Bezerra & Cavalc. 1967KC01
              |         Bicricium Sorokīn 1889KC01
              |         Biporisporites Ke & Shi 1978KC01
              |         Bireticulasporis Potonié & Sah 1960KC01
              |         Boletellites Briot, Lar.-Coll. & Locq. 1983KC01
              |         Botrytites Mesch. 1892KC01
              |         Brachysporisporites Lange & Sm. 1971KC01
              |         Brefeldiellites Dilcher 1965KC01
              |         Bretonia Bertrand & Hovel. 1892KC01
              |         Cadyexinis Stach 1957KC01
              |         Caenomyces Berry 1916KC01
              |         Callorites Fiore 1932KC01
              |         Calothyriopeltis Stevens & Ryan 1925 (n. d.) [=Calothyris Clem. & Shear 1931]KC01
              |         Campylostylus Genev. 1873 (n. d.)KC01
              |         CanceromycesKC01
              |         Cannanorosporonites Ramanujam & Rao 1979KC01
              |         Capnocybe Hughes 1966KC01
              |         Capsulasclerotes Malan 1959KC01
              |         Carlia Rabenh. 1857 (n. d.) non Gray 1845 (ICZN)KC01
              |         Carpolithes Brongn. 1822KC01
              |         Cellulasclerotes Stach & Pickh. 1957KC01
              |         Celyphus Batten 1973KC01
              |         Cenangites Mesch. 1892KC01
              |         Centonites Peppers 1964KC01
              |         Ceratonema Pers. 1822 (n. d.)KC01
              |         Cercosporites Salmon 1903KC01
              |         ‘Cercosporites’ Stopes 1913 non Salmon 1903KC01
              |         Chaetocypha Corda 1829 (n. d.)KC01
              |         Chaetomites Pampal. 1902KC01
              |         Chaetophorites Pratje 1922KC01
              |           |--*C. gomontoides Pratje 1922H75
              |           `--C. cruciatus Mägdefrau 1937H75
              |         Chaetosphaerites Félix 1894KC01
              |           `--C. pollenisimilis (Horst) Butterworth & Williams 1958P63
              |         Chaetosporium Corda 1833 (n. d.) [=Chaetotrichum Rabenh. 1844]KC01
              |         Chamaenema Kütz. 1833 (n. d.)KC01
              |         Charrinia Viala & Ravaz 1894 (n. d.)KC01
              |         Chordecystia Foster 1979KC01
              |         Weinmannodora Fr. 1849 (n. d.) [=Circinastrum Clem. 1909]KC01
              |         Cirrholus Mart. 1820 (n. d.)KC01
              |         Cladophytum Leidy 1849 (n. d.)KC01
              |         Cladosporites Félix 1894KC01
              |         Clasterosporites Pia 1927KC01
              |         Clavascina Beneš 1961KC01
              |         Clypeolaria Tratt. 1825 (n. d.)KC01
              |         Clypeomyces Kirschst. 1935 (n. d.)KC01
              |         Coccidophthora Syd. & Syd. 1913 (n. d.)KC01
              |         Coccophysium Link 1833 (n. d.)KC01
              |         Coleocarpon Stubblef., Taylor et al. 1983KC01
              |         Coleospora Cribb 1959 (n. d.)KC01
              |         Colligerites Jain & Kar 1979KC01
              |         Coniothyriites Babajan & Tasl. 1970KC01
              |         Conisphaeria Cooke 1879 (n. d.)KC01
              |         Coralloides Wulfen 1776 (n. d.)KC01
              |         Corenohydnum Lloyd 1936KC01
              |         Coronasclerotes Stach & Pickh. 1957KC01
              |         Corynelites Babajan & Tasl. 1970KC01
              |         Crandallia Ellis & Sacc. 1897KC01
              |         Crenasclerotes Stach & Pickh. 1957KC01
              |         Cryptodesma Leidy 1850 (n. d.)KC01
              |         Cyanotheca Pascher 1914 (n. d.)KC01
              |         Cyathela Raf. 1819 (n. d.)KC01
              |         Cyathella Raf. 1815 (n. d.)KC01
              |         Cylindrotaenium Thomé 1867 (n. d.)KC01
              |         Cynicus Raf 1815 (n. d.)KC01
              |         Cytosporites Babajan & Tasl. 1970KC01
              |         Dactylosporites Paradkar 1976KC01
              |         Daedaleites Mesch. 1892KC01
              |         Darwiniella Speg. 1887 (n. d.)KC01
              |         Dendromyceliates Jain & Kar 1979KC01
              |         Densospora McGee 1996KC01
              |         Depazites Geinitz 1855KC01
              |         Diamphora Mart. 1821 (n. d.)KC01
              |         Dicellaeporisporites Kalgutkar 1997KC01
              |         Dicellaesporites Elsik 1968KC01
              |         Dicteridium Raf. 1815 (n. d.)KC01
              |         Dictyochora Theiss. & Syd. 1914 (n. d.)KC01
              |         Dictyochorella Theiss. & Syd. 1915 (n. d.)KC01
              |         Dictyochorina Chardón 1932 (n. d.)KC01
              |         ‘Dictyonema’ Reinsch 1875 nec Agardh ex Kunth 1822 nec Hall 1851 (ICZN)KC01
              |         Dictyosporites Félix 1894KC01
              |         Didymocrater Mart. 1821 (n. d.)KC01
              |         Didymoporisporonites Sheffy & Dilcher 1971KC01
              |         Didymosamarospora Johnson & Gold 1957 (n. d.)KC01
              |         Didymosphaerites Pia 1927KC01
              |         ‘Didymosphaerites’ Fiore 1932 non Pia 1927KC01
              |         Didymosporonites Sheffy & Dilcher 1971KC01
              |         Didymostilbe Bres. & Sacc. 1902 (n. d.)KC01
              |         Dihyphis Locq. 1985KC01
              |         Diplodites Babajan & Tasl. 1970KC01
              |         Diplosporites Pia 1927KC01
              |         Diporicellaesporites Elsik 1968KC01
              |         Diporisporites Hammen 1954KC01
              |         Diporopollis Dutta & Sah 1970KC01
              |         ‘Discina’ Bonord. 1851 nec Lamarck 1819 (ICZN) nec (Fr.) Fr. 1849KC01
              |         Discoascina Beneš 1961KC01
              |         Ditmaria Lühnem. 1809 (n. d.)KC01
              |         Dothideites Pat. 1893KC01
              |         Dothidites Mesch. 1892KC01
              |         Melanogramma (n. d.) [incl. Dothiora Fr. 1837 (nom. rej.) non Fr. 1849 (nom. cons.)]KC01
              |         Dremuspora Sal.-Cheb. & Locq. 1980KC01
              |         Dyadosporites Hammen 1954KC01
              |         Dyadosporonites Elsik 1968KC01
              |         Elsikisporonites Kumar 1990KC01
              |         Elytrosporangium Morais, Bat. & Massa 1966 (n. d.)KC01
              |         Endaematus Raf. 1814 (n. d.) [=Endematus Raf. 1815]KC01
              |         Endonius Raf. 1819 (n. d.)KC01
              |         Enterobotryum Preuss 1853 (n. d.)KC01
              |         Enteroramus Lichtw., White et al. 1999KC01
              |         Entopeltacites Selkirk 1972KC01
              |         Eomycetopsis Schopf 1968KC01
              |           |--*E. robusta Schopf 1968KS92
              |           |--E. filiformis Schopf 1968KS92
              |           |--E. schopfiiT84
              |           `--E. siberiensis Lo 1980KS92
              |         Eopolyporoides Rigby 1982KC01
              |         Ephebella Itzigs. 1857 (n. d.)KC01
              |         Ephedracetes Huang 1981KC01
              |         Epichloea Giard 1889 (n. d.)KC01
              |         Epixyla Raf. 1806 (n. d.)KC01
              |         ‘Erysiphites’ Pampal. 1902 non Mesch. 1902KC01
              |         Euactinomyces Langeron 1922 (n. d.)KC01
              |         Eurotites Mesch. 1892KC01
              |         Stegilla Rchb. 1828 (n. d.) (see below for synonymy)KC01
              |         Excipulites Göpp. 1836KC01
              |         Exesisporites Elsik 1969KC01
              |         Falciascina Beneš 1961KC01
              |         Favaria Raf. 1815 (n. d.)KC01
              |         Faveoletisporonites Ramanujam & Rao 1979KC01
              |         Favomicrosporon Benedek 1967 (n. d.)KC01
              |         Felixites Elsik ex Janson. & Hills 1990KC01
              |         Foliopollenites Sierotin 1961KC01
              |         Fomesporites Huang 1981KC01
              |         Fomites Locq. & Koeniguer 1982KC01
              |         Foveodiporites Varma & Rawat 1963KC01
              |         Foveoletisporonites Ramanujam & Rao 1979KC01
              |         Fractisporonites Clarke 1965KC01
              |         ‘Fuckelia’ Niessl. 1875 nec Bonord. 1864 nec (Nitschke ex Sacc.) Cooke 1869KC01
              |         Fungites Hallier 1865KC01
              |         ‘Fungites’ Casp. 1907 non Hallier 1865KC01
              |         Fusariellites Babajan & Tasl. 1977KC01
              |         Fusellites Babajan & Tasl. 1977KC01
              |         Fusicladiites Babajan & Tasl. 1973KC01
              |         Fusidites Mesch. 1902KC01
              |         Fusiformisporites Rouse 1962KC01
              |         Fusitheca Bonord. 1864 (n. d.)KC01
              |         Gelasinosporites Briot, Lar.-Coll. & Locq. 1983KC01
              |         Gelatinaria Raf. 1815 (n. d.)KC01
              |         Geleenites Dijkstra 1949KC01
              |         Geotrichites Stubblef., Mill. et al. 1985KC01
              |           `--G. glaesariusP92
              |         Gigasporites Phipps & Taylor 1996KC01
              |         Giraffachitina Locq. 1985KC01
              |         Gliotrichum Eschw. 1822 (n. d.)KC01
              |         Globoasclerotes Stach & Pickh. 1957KC01
              |         Globosasclerotes Stach & Pickh. 1957KC01
              |         Globosopyreno Lloyd 1923 (n. d.)KC01
              |         Glomites Taylor, Remy et al. 1995KC01
              |         Glossifungites Lomnicki 1886KC01 [=Glossofungites (l. c.)H75]
              |         GlukomycesKC01
              |         Glutinisporidium Thor 1930 (n. d.)KC01
              |         Lanosa Fr. 1825 (n. d.) [incl. Gnaphalomyces]KC01
              |         Gonatobotrytites Pia 1927KC01
              |         Graamspora Locq. & Sal.-Cheb. 1980KC01
              |         Granatisporites Elsik & Janson. 1974KC01
              |         Grandinia Fr. 1838 (n. d.) [incl. Tuberculinia Velen. 1922 (n. n.)]KC01
              |         Graphiolites Fritel 1910KC01
              |         Guizhounema Mu 1977KC01
              |         Gyromyces Göpp. 1844KC01
              |         Halisaria Giard 1889 (n. d.)KC01
              |         Halonia Fr. 1849 (n. d.)KC01
              |         Haplographites Félix 1894KC01
              |         Haplosporium Mont. 1843 (n. d.)KC01
              |         Helminthosporites Pia 1927KC01
              |         ‘Helminthosporites’ Chitaley & Sheikh 1971 non Pia 1927KC01
              |         Hemicyphe Corda 1831 (n. d.)KC01
              |         Hermanniasporidium Thor 1930 (n. d.)KC01
              |         Oliveonia Donk 1958 (see below for synonymy)KC01
              |         Heteroporimyces Elias 1966KC01
              |         Heteroradulum Lloyd 1917 (n. d.)KC01
              |         Himantites Debey & Ettingsh. 1859KC01
              |         Hormosporites Grüss 1928KC01
              |         Hyalasterina Speg. 1919 (n. d.)KC01
              |         Hydnites Mesch. 1892KC01
              |         Typhoderma Gray 1821 (n. d.) [incl. Hygrocrocis Agardh 1824]KC01
              |         Hyperus Stevens 1927 (n. d.)KC01
              |         Hypochnites Mesch. 1898KC01
              |         Hypospila Fr. 1825 (n. d.)KC01
              |         Hypostomum Vuill. 1896 (n. d.)KC01
              |         Hypoxylites Kirschst. 1925KC01
              |         Hypoxylonites Elsik 1990KC01
              |         Hypoxylonsporites Kumar 1990KC01
              |         Hypsilophora Berk. 1879KC01
              |         Hysteriopsis Geyl. 1887KC01
              |         Hysterites Göpp. 1836KC01
              |         ‘Hysterites’ Unger 1841 non Göpp. 1836KC01
              |         Hysteromyxa Sacc. & Ellis 1882 (n. d.)KC01
              |         Inapertisporites Hammen ex Rouse 1959KC01
              |         Incertisporites Hammen 1954KC01
              |         Inciliaria Fr. 1825 (n. d.)KC01
              |         Incolaria Herzer 1893KC01
              |         Involutisporonites Clarke 1965KC01
              |         Isomunkia Theiss. & Syd. 1915KC01
              |         Jansoniisporites Kalgutkar 1997KC01
              |         Japewiella Printzen 1999KC01
              |         Kakabekia Barghoorn 1965KC01
              |           `--*K. umbellata Barghoorn 1965G79
              |         Kellermaniites Babajan & Tasl. 1977KC01
              |         Kutchiathyrites Kar 1979KC01
              |         Lacrimasporonites Clarke 1965KC01
              |         Ladrococcus Locq. 1985KC01
              |         Laestadites Mesch. 1892KC01
              |         Langloisula Ellis & Everh. 1889KC01
              |         Lecaniascus Moniez 1887KC01
              |         Lentescospora Linder 1944 (n. d.)KC01
              |         Lenzitites Mesch. 1892KC01
              |         Leolophia (n. d.)KC01
              |         Lepiotasporites Huang 1981KC01
              |         Leprosis Neck. ex Kremp. 1869 (n. d.)KC01
              |         Leptodothis Theiss. & Syd. 1914 (n. d.) [=Wrightiella Speg. 1923 non Schmitz 1893]KC01
              |         Leptonema Sm. 1896KC01
              |         Leptosphaerites Richon 1885KC01
              |         Leptospora Raf. 1808 (n. d.)KC01
              |         Leptuberia Raf. 1808KC01
              |         Liaoningnema Zheng & Zhang 1986KC01
              |         Limacinia Neger 1895 (n. d.)KC01
              |         Limnomyces Lohammar 1953 (n. d.)KC01
              |         Scopularia Preuss 1851 (n. d.) (see below for synonymy)KC01
              |         Lirasporis Potonié & Sah 1960KC01
              |         Lymphocystidium Weiser 1943KC01
              |         Macrosporites Renault 1899KC01
              |         Magnosporites Rouse 1962KC01
              |         Maireomyces Feldmann 1941 (n. d.)KC01
              |         Malacodermum Marchand 1896 (n. d.)KC01
              |         Malacodon Bataille 1923 (n. d.)KC01
              |         Manikinipollis Krutzsch 1970KC01
              |         Marssoninites Babajan & Tasl. 1970KC01
              |         Martinella (Cooke) Sacc. 1892 (n. d.) non Jousseaume 1887 (ICZN)KC01
              |         Maurya Pat. 1898 (n. d.) [=Pluesia Nieuwl. 1916]KC01
              |         Medastina (n. d.)KC01
              |         Medusomyces Lindau 1913KC01
              |         Melanconites Göpp. 1852KC01
              |         Melanomyces Syd. & Syd. 1917 (n. d.)KC01
              |         Melanosphaerites Grüss 1928KC01
              |         Melanosporites Pampal. 1902KC01
              |         Meliolopsis (Sacc.) Sacc. 1891 (n. d.)KC01
              |         MeniscoideisporitesKC01
              |         Merosporium Corda 1831 (n. d.)KC01
              |         Metasphaerella Speg. 1924 (n. d.)KC01
              |         Microdiplodiites Babajan & Tasl. 1973KC01
              |         ‘Microdiscus’ Steinecke 1916 (n. d.) non Sacc. 1916KC01
              |         Microsporonites Jain 1968KC01
              |         Microthyrites Pampal. 1902KC01
              |         Mirandia Toro 1934 (n. d.)KC01
              |         Molgosporidium Thor 1930 (n. d.)KC01
              |         Moniliites Babajan & Tasl. 1973KC01
              |         Monilites Pampal. 1902KC01
              |         Monochaetites Babajan & Tasl. 1973KC01
              |         Monoporisporites Hammen 1954KC01
              |         Chrysomilia sitophilaKC01
              |         Mucedites Bertrand & Renault 1896KC01
              |         Muchmoria Sacc. 1906 (n. d.)KC01
              |         Mucoralites Patel 1979KC01
              |         Mucorites Mesch. 1898KC01
              |         Multicellaesporites Elsik 1968KC01
              |         Murciasporidium Thor 1930KC01
              |         Mycetosporidium Léger & Hesse 1905KC01
              |         Mycobystrovia Goujet & Locq. 1979KC01
              |         Mycocoelium Kütz. 1843 (n. d.)KC01
              |         Mycogemma Zalessky 1915KC01
              |         Mycokidstonia Pons & Locq. 1981KC01
              |         Mycorhizonium Weiss 1904KC01
              |         Mycothamnion Kütz. 1843 (n. d.)KC01
              |         Myxomonas Brzez. 1906 (n. d.)KC01
              |         Myxonyphe Rchb. 1841 (n. d.) [=Myxonema Corda 1837 non Fr. 1825, Pyonema Nieuwl. 1916]KC01
              |         Naegelia Rabenh. 1844 (n. d.)KC01
              |         Nailisporites Huang 1981KC01
              |         Nemaclada Sm. 1896KC01
              |         Nemaplana Sm. 1896KC01
              |         Nematococcus Kütz. 1833 (n. d.)KC01
              |         Piptostoma Berk. & Broome 1875 (n. d.) [=Neopiptostoma Kuntze 1898]KC01
              |         Nesophloea Fr. 1849 (n. d.)KC01
              |         Neuroecium Kunze 1823 (n. d.)KC01
              |         Nothodiscus Sacc. 1917 (n. d.)KC01
              |         Nowellia Stevens 1924 (n. d.)KC01
              |         Nyctomyces Hartig 1833KC01
              |         Octosporonites Locq. & Sal.-Cheb. 1980KC01
              |         Oesophagomyces Manier & Ormières 1980KC01
              |         Oichitonium (n. d.)KC01
              |         Oidites Mesch. 1892KC01
              |         Oidospora Will. 1878KC01
              |         ‘Ombrophila’ Quél. 1892 non Fr. 1849KC01
              |         Omoriza Paulet 1812KC01
              |         Omphalocystis Balbiani 1889KC01
              |         Onakawananus Radforth 1958KC01
              |         Oncobyrsa Agardh 1827 (n. d.)KC01
              |         Oovorus Entz 1930 (n. d.)KC01
              |         Opethyrium Speg. 1919 (n. d.)KC01
              |         Ophryomyces Léger & Hesse 1909 (n. d.)KC01
              |         Ordovicimyces Elias 1966KC01
              |         Ornasporonites Ramanujam & Rao 1979KC01
              |         Ospriosporium Corda 1831 (n. d.)KC01
              |         Ostracodermis Locq. 1982KC01
              |         Ovularites Whitford 1916KC01
              |         Palaeodikaryomyces Dörfelt 1998KC01
              |         PalaeodiploditesKC01
              |         Palaeofibula Osborn, Taylor & White 1989KC01
              |         Palaeomycelites Bystrov 1956KC01
              |         Palaeopericonia Ibáñez & Zamuner 1996KC01
              |         Palambages Whetzel 1961KC01
              |           `--*P. morulosa Wetzel 1961H75
              |         Paleoarcyria Jacq.-Fél., Mill. & Locq. 1983KC01
              |         Paleoblastocladia Remy, Taylor & Hass 1994KC01
              |         Paleocatenaria Locq. 1983KC01
              |         Paleoguttulina Locq. & Mišík 1983KC01
              |         Paleoserenomyces Currah, Stockey & LePage 1998KC01
              |         Palmellathyrites Locq., Pons & Sal.-Cheb. 1981KC01
              |         Palynomorphites Moore 1963KC01
              |         Papulasporites Shete & Kulk. 1978KC01
              |         Papulosporonites Schmied. & Schwab 1964KC01
              |         Parapolyporites Tanai 1987KC01
              |         Xenosoma Syd. & Syd. 1921 (n. d.) (see below for synonymy)KC01
              |         Pelodiscus Clem. 1901 (n. d.)KC01
              |         Penicillites Mesch. 1892KC01
              |         Peniophorina Höhn. 1917 (n. d.)KC01
              |         Perisporites Pampal. 1902KC01
              |         Perisporium Fr. 1821 (n. d.) [Perisporiales]KC01
              |         Peronosporites Sm. 1877KC01
              |           `--P. pythiusP92
              |         Peronosporoides Sm. 1896KC01
              |         Pestalozzites Berry 1917KC01
              |         Petrosphaeria Stopes & Fujii 1910KC01
              |         Pezizasporites Huang 1981KC01
              |         Phacidiopsis Geyl. 1887KC01
              |         Phacidites Mesch. 1892KC01
              |         Phaeonema Kütz. 1843KC01
              |         Phaeoscutella Henn. 1904 (n. d.)KC01
              |         Phaeosiphonia Kütz. 1849 (n. d.)KC01
              |         Phellomycetes Renault 1896 [=Phellomycites Mesch. 1896]KC01
              |         Phelonites Fresen. 1861KC01
              |         Phiala Raf. 1815 (n. d.)KC01
              |         Phialea (Fr.) Gillet 1879 (n. d.)KC01
              |         Phomites Fritel 1910KC01
              |         Phragmidiites Babajan & Tasl. 1970KC01
              |         Phragmonaevia Rehm 1896 (n. d.)KC01
              |         ‘Phragmothyriella’ Speg. 1919 (n. d.) non Höhn. 1912KC01
              |         Phycomater Fr. 1825 (n. d.)KC01
              |         Phycomycites Ellis 1915KC01
              |         Phyllerites Mesch. 1892KC01
              |         Phyllerium Fr. 1832 (n. d.)KC01
              |         Phyllonochaeta Gonz. Frag. & Cif. 1927 (n. d.)KC01
              |         Phyllops Raf. 1817 (n. d.)KC01
              |         Phyllostictites Babajan & Tasl. 1970KC01
              |         Physalosporella Speg. 1910 (n. d.)KC01
              |         Pillulasclerotes Stach & Pickh. 1957KC01
              |         ‘Pilula’ Harker, Sarjeant & Caldwell 1990 non Massee 1910KC01
              |         Piriurella Cookson & Eisenack 1979KC01
              |         Placodium Hill 1773 (n. d.)KC01
              |         Plectosclerotes Stach & Pickh. 1957KC01
              |         Pleosporites Suzuki 1910KC01
              |         Pleosporonites Lange & Sm. 1971KC01
              |         Plochmopeltinites Cookson 1947KC01
              |         Pluricellaesporites Hammen 1954KC01
              |         Pluricellulites Hammen 1954KC01
              |         Polyadosporites Hammen 1954KC01
              |         Polycellaesporonites Chandra, Saxena & Setty 1984KC01
              |         Polycellaria Pflug 1965KC01
              |           `--*P. bonnerensis Pflug 1965G79
              |         Polyporasclerotes Stach & Pickh. 1957KC01
              |         Polyporisporites Hammen 1954KC01
              |         Polyporites Lindl. & Hutton 1833KC01
              |         ‘Polyporites’ Daugherty 1941 non Lindl. & Hutton 1833KC01
              |         Polystema Raf. 1815 (n. d.)KC01
              |         Poronea Raf. 1815 (n. d.)KC01
              |         Portalites Hemer & Nygreen 1967KC01
              |         Predaldinia Briot, Lar.-Coll. & Locq. 1983KC01
              |         Premyxomyces Locq. 1979KC01
              |         Protomycites Mesch. 1892KC01
              |         Pseudohansenula Mogi 1939 (n. d.)KC01
              |         ‘Pseudohansenula’ Novák & Zsolt 1961 non Mogi 1939 (n. d.)KC01
              |         Pseudopezizites Fiore 1932KC01
              |         Pseudophyllachora Speg. 1919 (n. d.)KC01
              |         Psiammopomopiospora Locq. & Sal.-Cheb. 1980KC01
              |         Psiamspora Locq. & Sal.-Cheb. 1980KC01
              |         Psidimobipiospora Locq. & Sal.-Cheb. 1980KC01
              |         Psilodiporites Varma & Rawat 1963KC01
              |         Psorothecium Massal. 1860 (n. d.)KC01
              |         Puccinites Ettingsh. 1853KC01
              |         Puttemansiella Henn. 1908 (n. d.)KC01
              |         Pycnodothis Stevens 1924 (n. d.)KC01
              |         Pyrenophoropsis Ramesh 1988KC01
              |         Pyrisperma Raf. 1808 (n. d.)KC01
              |         Pythites Pampal. 1902KC01
              |         Rabenhorstiites Teterevn. & Tasl. 1977KC01
              |         Ramulariites Babajan & Tasl. 1973KC01
              |         Ramularites Pia 1927KC01
              |         Reduviasporonites Wilson 1962KC01
              |         Reticellites Glass, Br. & Elsik 1987KC01
              |         Reticulatisporonites Elsik 1968KC01
              |         Reticulosporidium Thor 1930 (n. d.)KC01
              |         Retidiporites Varma & Rawat 1963KC01
              |         Reymanella Marcink. 1979KC01
              |         Rhagidiasporidium Thor 1930 (n. d.)KC01
              |         Rhizoblepharis Dang. 1900 (n. d.)KC01
              |         Rhizogaster Reinsch. 1875 (n. d.)KC01
              |         ‘Rhizomorpha’ Ach. 1809 (n. d.) non Roth 1791KC01
              |         Rhizomorphites Göpp. 1848KC01
              |         Rhizophagites Rosend. 1943KC01
              |         Rhynchotheca Kleb. 1933 (n. d.)KC01
              |         Rhytismites Mesch. 1892KC01
              |         Rhytismopsis Geyl. 1887KC01
              |         Roannaisia Taylor, Galtier & Axsmith 1994KC01
              |         Rosasporina Beneš 1956KC01
              |         Rosellinites Mesch. 1892KC01
              |         ‘Rosellinites’ Potonié 1893 non Mesch. 1892KC01
              |         Saccharomycetes Grüss 1928KC01
              |         Samarosporella Linder 1944 (n. d.)KC01
              |         Sarcanthia Raf. 1817 (n. d.)KC01
              |         Sarcomelas Raf. 1817 (n. d.)KC01
              |         Sarconemus Raf. 1815 (n. d.)KC01
              |         Scabradiporites Mathur 1966KC01
              |         Schizacrospermum Henn. 1899 (n. d.)KC01
              |         Schizographa Nyl. 1857 (n. d.)KC01
              |         Scirrhiopsis Henn. 1905 (n. d.)KC01
              |         Sclerosporis Stach & Chandra 1956KC01
              |         Sclerotites Jeffrey & Chrysler 1906KC01
              |         Scolecosporites Lange & Sm. 1971KC01
              |         Scutellum Speg. 1881 (n. d.) non Pusch 1833 (ICZN)KC01
              |         Scytopezis Clem. 1903 (n. d.)KC01
              |         Selenophomites Babajan & Tasl. 1973KC01
              |         Senegalosporites Jardiné & Magloire 1965KC01
              |         Serpentisclerotes Beneš 1959KC01
              |         Sirocrocis Kütz. 1843 (n. d.)KC01
              |         Sirodesmites Pia 1927KC01
              |         Sorosporonites Mu 1977KC01
              |         Spadonia Fr. 1829 (n. d.)KC01
              |         Spegazzinites Félix 1894KC01
              |         Sphaerialites Venkatach. & Kar 1969KC01
              |         Sphaerites Unger 1850 non Duftschmid 1805 (ICZN)KC01
              |         Sphaeropleum Link 1833 (n. d.)KC01
              |         Sphaeroporalites Hemer & Nygreen 1967KC01
              |         Sphaerosporiites Babajan & Tasl. 1977KC01
              |         Spherites Dijkstra 1949KC01
              |         Spilonemella Henssen & Tonsberg 2000KC01
              |         Spilosphaerites Massal. 1857KC01
              |         Spiroidium Saito 1949KC01
              |         Spirotremesporites Dueñas 1979KC01
              |         Spongiasclerotes Stach & Pickh. 1957KC01
              |         Sporadospora Reinsch 1875 (n. d.)KC01
              |         Sporonites Potonié 1931KC01
              |         Sporophaga Harkn. 1899 (n. d.)KC01
              |         Sporoschismatites Babajan & Tasl. 1977KC01
              |         Sporotrichites Göpp. & Berendt 1945KC01
              |         Stagonosporites Babajan & Tasl. 1970KC01
              |         Staphlosporonites Sheffy & Dilcher 1971KC01
              |         Stegites Mesch. 1892KC01
              |         Stellasclerotes Beneš 1959KC01
              |         Stemphyliites Babajan & Tasl. 1973KC01
              |         Stericium Raf. 1819 (n. d.)KC01
              |         Stigmateacites Zheng & Zhang 1986KC01
              |         Stilbites Pia 1927KC01
              |         Straggaria Reinsch 1888 (n. d.)KC01
              |         Streptotrichites Mesch. 1892KC01
              |         Striadiporites Varma & Rawat 1963KC01
              |         Striadyadosporites Dueñas 1979KC01
              |         Striatasclerotes Stach & Pickh. 1957KC01
              |         Stridiporosporites Ke & Shi 1978KC01
              |         Sulcatisclerotes Beneš 1959KC01
              |         Synaphia Nees & Nees ex Rchb. 1841 (n. d.)KC01
              |         Synchaetophagus Apstein 1911 (n. d.)KC01
              |         Tetena Raf. 1806 (n. d.)KC01
              |         Theissenula Syd. & Syd. 1914 (n. d.)KC01
              |         Thuchomyces Hallbauer & Jahns 1977KC01
              |         Thyriascus Schulzer 1877 (n. d.)KC01
              |         Tormentella Pflug 1966KC01
              |           `--*T. tubiformis Pflug 1966G79
              |         Torulites Grüss 1927KC01
              |         ‘Torulites’ Pia 1927 non Grüss 1927KC01
              |         Trametites Mesch. 1892KC01
              |         Transeptaesporites Ediger 1981KC01
              |         Trematophora Eisenack 1965KC01
              |         Trematosphaerites Mesch. 1892KC01
              |         ‘Trematosphaerites’ Grüss 1924 non Mesch. 1892KC01
              |         Tricellaesporonites Sheffy & Dilcher 1971KC01
              |         Trichobacidia Vain. 1921 (n. d.)KC01
              |         Trichochora Theiss. & Syd. 1915 (n. d.)KC01
              |         Trichosporites Félix 1894KC01
              |         ‘Trichosporum’ Fr. 1825 non Don 1822KC01
              |         Trichothyrites Rosend. 1943KC01
              |         Trihyphaecites Peppers 1970KC01
              |         Triporicellaesporites Ke & Shi 1978KC01
              |         Triporisporites Hammen 1954KC01
              |         Triporisporonites Sheffy & Dilcher 1971KC01
              |         Triposporonites Sheffy & Dilcher 1971KC01
              |         Tubercularites Arcang. 1903KC01
              |         Tympanicysta Malme 1980KC01
              |         Uncinulites Pampal. 1902KC01
              |         Uredites Babajan & Tasl. 1970KC01
              |         Urocystites Babajan & Tasl. 1970KC01
              |         Urophlyctites Magnus 1903KC01
              |         Ustilagites Babajan & Tasl. 1970KC01
              |         Ustulinites Kirschst. 1925KC01
              |         Uvasporina Beneš 1956KC01
              |         Variolaria Pers. 1794 (n. d.)KC01
              |         ‘Variolaria’ Gray 1821 (n. d.) non Pers. 1794KC01
              |         Vermiculariella Oudem. 1898KC01
              |         Warkallisporonites Ramanujam & Rao 1979KC01
              |         Winfrenatia Taylor, Hass & Kerp 1997KC01
              |         Winterina Sacc. 1891KC01
              |         ‘Xyloma’ Raf. 1837 (n. d.) non Pers. 1794KC01
              |         Xylomites Unger 1841 [=Xylomides Schimp. 1869]KC01
              |         Zercosporidium Thor 1930 (n. d.)KC01
              |         Zonilia Raf. 1815 (n. d.)KC01
              |         Zygodesmella Gonz. Frag. 1917 (n. d.)KC01
              |         Zygodesmus Corda 1837 (n. d.)KC01
              |         Zygosporites Will. 1880KC01
              |         Zygothrix Reinsch ex Rabenh. 1866 (n. d.)KC01
                 |  `--ChytridiomycotaJK06
                               `--Dikarya (see below for synonymy)JK06
                                    |  i. s.: Sclerotium Tode 1790 (see below for synonymy)KC01
                                    |           |--‘Bromicolla’ aleuticaED89
                                    |           |--S. cepivorumKC01
                                    |           |--S. delphiniiKC01
                                    |           |--S. microsclerotiumKC01
                                    |           |--S. oryzaeKC01
                                    |           |--S. rolfsiiRMS95
                                    |           `--S. tuliparumKC01
                                    `--Basidiomycota (see below for synonymy)JK06
                                         |  i. s.: Mycaureola Maire & Chemin 1922RS99
                                         |           `--*M. dilseae Maire & Chemin 1922RS99
                                         |         WallemiomycotinaAB19
                                         |           |--Geminibasidium (see below for synonymy)AB19
                                         |           `--Wallemia Johan-Olsen 1887AS12 (see below for synonymy)
                                         |                `--W. sebiRC11
                                         |         Akenomyces Arnaud ex Hornby 1984KC01
                                         |         Anixia Fr. 1819 (n. d.)KC01
                                         |         Archeterobasidium Koeniguer & Locq. 1980KC01
                                         |         Arthrodochium Castañeda & Kendr. 1990KC01
                                         |         Arualis Katz 1980KC01
                                         |         Cenangiomyces Dyko & Sutton 1979KC01
                                         |         Cerinosterus Moore 1987KC01
                                         |         Cilicia Fr. 1825KC01
                                         |         Corticomyces Romero & López 1989KC01
                                         |         Cruciger Kirschner & Oberw. 1999KC01
                                         |         Dactyloporus Herzer 1893KC01
                                         |         Dendrosporomyces Nawawi, Webster & Davey 1977KC01
                                         |         Druparia Raf. 1808 (n. d.) [=Drupasia Raf. 1809]KC01
                                         |         Ellula Nag Raj 1980KC01
                                         |         Endoconia Raf. 1819 (n. d.)KC01
                                         |         Fibulochlamys Romero & Cabral 1989KC01
                                         |         Fibulocoela Nag Raj 1978KC01
                                         |         Fibulotaeniella Marvanová & Bärl. 1988KC01
                                         |         Gastromyces Ludw. 1861KC01
                                         |         Geotrichopsis Tzean & Estey 1991KC01
                                         |         Gloeoradulum Rick 1959 (n. d.)KC01
                                         |         Gloeosynnema Seifert & Okada 1988KC01
                                         |         Glomerulomyces Romero & López 1989KC01
                                         |         Glutinoagger Sivan. & Watling 1980KC01
                                         |         Grandiniochaete Rick 1940 (n. d.)KC01
                                         |         Granulina Velen. 1947 (n. d.) non Jousseaume 1888 (ICZN)KC01
                                         |         Heterobasidium Massee 1889KC01
                                         |         Hyphobasidiofera Matsush. & Matsush. 1996KC01
                                         |         Itersonilia Derx 1948KC01
                                         |         Mycodendron Massee 1891 (n. d.)KC01
                                         |         Mycospongia Velen. 1939 (n. d.)KC01
                                         |         Myrioconium Fr. 1823 [incl. Rhacodiella Peyronel 1919]KC01
                                         |         Naiadella Marvanová & Bandoni 1987KC01
                                         |         Nostoclavus Paulet 1791 (n. d.)KC01
                                         |         Nyctalina Arnaud 1952KC01
                                         |         Odontiochaete Rick 1940 (n. d.)KC01
                                         |         Pagidospora Drechsler 1960KC01
                                         |         Phellinites Singer & Archang. 1958KC01
                                         |         Phlegmatium Fr. 1819 (n. d.)KC01
                                         |         Pilacre Fr. 1825 (n. d.)KC01
                                         |         Pneumatospora Sutton, Kuthub. & Muid 1984 (see below for synonymy)KC01
                                         |         Polygaster Fr. 1823 (n. d.)KC01
                                         |         Pseudasterodon Rick 1959 (n. d.)KC01
                                         |         Pycnovellomyces Castañeda 1987KC01
                                         |         Radulochaete Rick 1940 (n. d.)KC01
                                         |         Riessia Fresen. 1852KC01
                                         |         Riessiella Jülich 1985KC01
                                         |         Rimella Raf. 1819 (n. d.)KC01
                                         |         Spiniger Stalpers 1974KC01
                                         |         Stemastrum Raf. 1808 (n. d.)KC01
                                         |         Titaeella Arnaud ex Ando & Tubaki 1985KC01
                                         |         Volvycium Raf. 1808 (n. d.)KC01
                                         |         Xenopus Penz. & Sacc. 1901 (n. d.) non Wagler 1827 (ICZN)KC01

Basidiomycota [Basidioblastomycetes, Basidiomycetes, Basidiomycotina, Basimycetes, Heterobasidiomycetae, Sporobolomycetales, Teliomycetes]JK06

Dikarya [Blastomycetes, Blastomycota, Cryptococcales, Dangeardiomycetes, Dikaryomycota, Endomycetes, Endomycota, Neomycota]JK06

Fungi [Amastigomycota, Archemycota, Carpomycetes, Dictyomycotina, Entomycetidae, Eomycota, Eozygomycetia, Eumycetes, Eumycophyta, Eumycota, Fungales, Glomomycetes, Hysterophyta, Inophyta, Melanomycotina, Mycetalia, Mycetales, Mycetes, Mycophyta, Mycota, Neozygomycetia, Opisthomastigomycota, Pedomycetidae, Spizomycetidae, Uniflagellatae, Zoomycetes, Zygomycetes, Zygomycota, Zygomycotina]C-SCL15

Geminibasidium [Geminibasidiaceae, Geminibasidiales, Geminibasidiomycetes]AB19

Oliveonia Donk 1958 [=Heteromyces Olive 1957 non Müll. Arg. 1889; incl. Oliveorhiza Roberts 1998, Sebacinella Hauerslev 1977; Oliveoniaceae]KC01

Pneumatospora Sutton, Kuthub. & Muid 1984 [incl. Tricellulortus Matsush. 1995]KC01

Sclerotium Tode 1790 [incl. Apiosporium Kunze 1817, Astoma Gray 1821 non Oken 1815 (ICZN), Bromicolla Eichw. 1843, Coccopleum Ehrenb. 1818, Sclerotiomyces Woron. 1926, Xylochoeras Fr. 1849; Agonomycetales]KC01

Scopularia Preuss 1851 (n. d.) [=Lindavia Nieuwl. 1916 non (Schütt) De Toni & Forti 1900, Outhovia Nieuwl. 1916]KC01

Stegilla Rchb. 1828 (n. d.) [=Eustegia Fr. 1823 non Br. 1810, Sphaerothyrium Wallr. 1833, Stegia Fr. 1818 non DC. 1805]KC01

Wallemia Johan-Olsen 1887AS12 [incl. Bargellinia Borzí 1888KC01, Hemispora Vuill. 1906KC01; Wallemiaceae, Wallemiales, Wallemiomycetes]

Xenosoma Syd. & Syd. 1921 (n. d.) [=Paulia Lloyd 1916 nec Fée 1836 nec Gray 1840 (ICZN) nec Bourguignat 1882 (ICZN)]KC01

*Type species of generic name indicated


[AB19] Adl, S. M., D. Bass, C. E. Lane, J. Lukeš, C. L. Schoch, A. Smirnov, S. Agatha, C. Berney, M. W. Brown, F. Burki, P. Cárdenas, I. Čepička, L. Chistyakova, J. del Campo, M. Dunthorn, B. Edvardsen, Y. Eglit, L. Guillou, V. Hampl, A. A. Heiss, M. Hoppenrath, T. Y. James, A. Karnkowska, S. Karpov, E. Kim, M. Kolisko, A. Kudryavtsev, D. J. G. Lahr, E. Lara, L. Le Gall, D. H. Lynn, D. G. Mann, R. Massana, E. A. D. Mitchell, C. Morrow, J. S. Park, J. W. Pawlowski, M. J. Powell, D. J. Richter, S. Rueckert, L. Shadwick, S. Shimano, F. W. Spiegel, G. Torruella, N. Youssef, V. Zlatogursky & Q. Zhang. 2019. Revisions to the classification, nomenclature, and diversity of eukaryotes. Journal of Eukaryotic Microbiology 66: 4–119.

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[AB-E04] Agapow, P.-M., O. R. P. Bininda-Emonds, K. A. Crandall, J. L. Gittleman, G. M. Mace, J. C. Marshall & A. Purvis. 2004. The impact of species concept on biodiversity studies. Quarterly Review of Biology 79 (2): 161–179.

[BB03] Bauer, R., D. Begerow, F. Oberwinkler & L. Marvanová. 2003. Classicula: the teleomorph of Naiadella fluitans. Mycologia 95 (4): 756–764.

Boyce, C. K., & C. L. Hotton. 2010. Prototaxites was not a taphonomic artifact. American Journal of Botany 97 (7): 1073.

Boyce, C. K., C. L. Hotton, M. L. Fogel, G. D. Cody, R. M. Hazen, A. H. Knoll & F. M. Hueber. 2007. Devonian landscape heterogeneity recorded by a giant fungus. Geology 35: 399–402.

[BJ17] Budd, G. E., & S. Jensen. 2017. The origin of the animals and a ‘savannah’ hypothesis for early bilaterian evolution. Biological Reviews 92 (1): 446–473.

[C-S98] Cavalier-Smith, T. 1998. A revised six-kingdom system of life. Biological Reviews 73: 203–266.

[C-SC03] Cavalier-Smith, T., & E. E.-Y. Chao. 2003. Phylogeny of Choanozoa, Apusozoa, and other Protozoa and early eukaryote megaevolution. Journal of Molecular Evolution 56: 540–563.

[C-SCL15] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2015. Multiple origins of Heliozoa from flagellate ancestors: new cryptist subphylum Corbihelia, superclass Corbistoma, and monophyly of Haptista, Cryptista, Hacrobia and Chromista. Molecular Phylogenetics and Evolution 93: 331–362.

[DP72] Deighton, F. C., & K. A. Pirozynski. 1972. Microfungi. V. More hyperparasitic hyphomycetes. Mycological Papers 128: 1–110.

[ED89] Edgeworth David, T. W. 1889. Note on the origin of “kerosene shale”. Proceedings of the Linnean Society of New South Wales, series 2, 4 (2): 483–500, pl. 18.

[EA12] Edwards, D., & L. Axe. 2012. Evidence for a fungal affinity for Nematasketum, a close ally of Prototaxites. Botanical Journal of the Linnean Society 168: 1–18.

[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.

[G79] Glaessner, M. F. 1979. Precambrian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A79–A118. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Graham, L. E., M. E. Cook, D. T. Hanson, K. B. Pigg & J. M. Graham. 2010a. Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats. American Journal of Botany 97 (2): 268–275.

Graham, L. E., M. E. Cook, D. T. Hanson, K. B. Pigg & J. M. Graham. 2010b. Rolled liverwort mats explain major Prototaxites features: response to commentaries. American Journal of Botany 97 (7): 1079–1086.

Graham, L. E., L. W. Wilcox, M. E. Cook & P. G. Gensel. 2004. Resistant tissues of modern marchantioid liverworts resemble enigmatic Early Paleozoic microfossils. Proceedings of the National Academy of Sciences of the USA 101 (30): 11025–11029.

[G14] Grazhdankin, D. 2014. Patterns of evolution of the Ediacaran soft-bodied biota. Journal of Paleontology 88 (2): 269–283.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[HO09] Hejnol, A., M. Obst, A. Stamatakis, M. Ott, G. W. Rouse, G. D. Edgecombe, P. Martinez, J. Baguña, X. Bailly, U. Jondelius, M. Wiens, W. E. G. Müller, E. Seaver, W. C. Wheeler, M. Q. Martindale, G. Giribet & C. W. Dunn. 2009. Assessing the root of bilaterian animals with scalable phylogenomic methods. Proceedings of the Royal Society of London Series B—Biological Sciences 276 (1677): 4261–4270.

Hillier, R. D., D. Edwards & L. B. Morrissey. 2008. Sedimentological evidence for rooting structures in the Early Devonian Anglo-Welsh Basin (UK), with speculation on their producers. Palaeogeography, Palaeoclimatology, Palaeoecology 270 (3–4): 366–380.

Hueber, F. M. 2001. Rotted wood–alga–fungus: the history and life of Prototaxites Dawson 1859. Review of Palaeobotany and Palynology 116 (1–2): 123–158.

[JK06] James, T. Y., F. Kauff, C. L. Schoch, P. B. Matheny, V. Hofstetter, C. J. Cox, G. Celio, C. Gueidan, E. Fraker, J. Miadlikowska, H. T. Lumbsch, A. Rauhut, V. Reeb, A. E. Arnold, A. Amtoft, J. E. Stajich, K. Hosaka, G.-H. Sung, D. Johnson, B. O’Rourke, M. Crockett, M. Binder, J. M. Curtis, J. C. Slot, Z. Wang, A. W. Wilson, A. Schüßler, J. E. Longcore, K. O’Donnell, S. Mozley-Standridge, D. Porter, P. M. Letcher, M. J. Powell, J. W. Taylor, M. M. White, G. W. Griffith, D. R. Davies, R. A. Humber, J. B. Morton, J. Sugiyama, A. Y. Rossman, J. D. Rogers, D. H. Pfister, D. Hewitt, K. Hansen, S. Hambleton, R. A. Shoemaker, J. Kohlmeyer, B. Volkmann-Kohlmeyer, R. A. Spotts, M. Serdani, P. W. Crous, K. W. Hughes, K. Matsuura, E. Langer, G. Langer, W. A. Untereiner, R. Lücking, B. Büdel, D. M. Geiser, A. Aptroot, P. Diederich, I. Schmitt, M. Schultz, R. Yahr, D. S. Hibbett, F. Lutzoni, D. J. McLaughlin, J. W. Spatafora & R. Vilgalys. 2006. Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443: 818–822.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[KS92] Kumar, S., & P. Srivastava. 1992. Middle to late Proterozoic microbiota from the Deoban Limestone, Garhwal Himalaya, India. Precambrian Research 56: 291–318.

[LK04] Lutzoni, F., F. Kauff, C. J. Cox, D. McLaughlin, G. Celio, B. Dentinger, M. Padamsee, D. Hibbett, T. Y. James, E. Baloch, M. Grube, V. Reeb, V. Hofstetter, C. Schoch, A. E. Arnold, J. Miadlikowska, J. Spatafora, D. Johnson, S. Hambleton, M. Crockett, R. Shoemaker, G.-H. Sung, R. Lücking, T. Lumbsch, K. O’Donnell, M. Binder, P. Diederich, D. Ertz, C. Gueidan, K. Hansen, R. C. Harris, K. Hosaka, Y.-W. Lim, B. Matheny, H. Nishida, D. Pfister, J. Rogers, A. Rossman, I. Schmitt, H. Sipman, J. Stone, J. Sugiyama, R. Yahr & R. Vilgalys. 2004. Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. American Journal of Botany 91 (10): 1446–1480.

[P63] Playford, G. 1963. Lower Carboniferous microfloras of Spitsbergen. Part two. Palaeontology 5 (4): 619–678.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[RMS95] Ravishankar, J. P., V. Muruganandam & T. S. Suryanarayanan. 1995. Isolation and characterization of melanin from a marine fungus. Botanica Marina 38: 413–416.

[RC11] Rosling, A., F. Cox, K. Cruz-Martinez, K. Ihrmark, G.-A. Grelet, B. D. Lindahl, A. Menkis & T. Y. James. 2011. Archaeorhizomycetes: unearthing an ancient class of ubiquitous soil fungi. Science 333: 876–879.

[RS99] Rossman, A. Y., G. J. Samuels, C. T. Rogerson & R. Lowen. 1999. Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1–248.

[S00] Schweitzer, H. J. 2000. Neue Pflanzenfunde im Unterdevon der Eifel (Deutschland). Senckenbergiana Lethaea 80 (2): 371–395.

Selosse, M.-A. 2002. Prototaxites: a 400 myr old giant fossil, a saprophytic holobasidiomycete, or a lichen? Mycological Research 106 (6): 642–644.

[T93] Taylor, T. N. 1993. Fungi. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 9–13. Chapman & Hall: London.

Taylor, T. N., E. L. Taylor, A.-L. Decombeix, A. Schwendemann, R. Serbet, I. Escapa & M. Krings. 2010. The enigmatic Devonian fossil Prototaxites is not a rolled-up liverwort mat: comment on the paper by Graham et al. (AJB 97: 268–275). American Journal of Botany 97 (7): 1074–1078.

[T84] Tewari, V. C. 1984. Stromatolites and Precambrian-Lower Cambrian biostratigraphy of the Lesser Himalaya. Proc. Vth India Geophytol. Conf., Lucknow (1983), Spec. Publ. 1984: 71–97.

[ZB10] Zvereva, L. V., & O. G. Borzykh. 2010. Filamentous fungi in the epigrowth of the Pacific oyster Crassostrea gigas (Bivalvia) in Peter the Great Bay, Sea of Japan. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of “Comparison on Marine Biodiversity in the Northwest Pacific Ocean”, 10–11 October 2010, Qingdao (China) pp. 215–219. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

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