Trechispora hymenocystis, copyright Urmas Ojango.

Belongs within: Orthomycotina.
Contains: Cantharellales, Polyporales, Thelephorales, Agaricomycetidae, Vuilleminiales, Hericiaceae, Russulales, Aleurodiscus, Phallomycetidae, Hyphodontia.

The mushroom tree
Published 24 May 2012

Autumn being the height of mushroom season, it is appropriate that mushrooms should be my subject today. Recent years have seen great advances in our understanding of fungal evoluton: you could almost say our knowledge has mushroomed.

The familiar field mushroom Agaricus arvensis, photographed by Fred Stevens.

The great majority of the macroscopic fungi that most people would be familiar with belong to the clade of dikaryotic fungi, so called for their production of dikaryotic (double-nucleate) hyphae. When these fungi reproduce, separate hyphae fuse to form a hypha containing nuclei from both parents, like a sperm fusing to an egg in human fertilisation. However, unlike a sperm and an egg, the parent nuclei do not immediately fuse. Instead, the resulting dikaryotic hypha grows and divides, and the nuclei divide within it while remaining separate from each other. It is the dikaryotic hyphae that produce the fruiting bodies of the fungus, so when you look at a mushroom you are not looking at the product of a single individual but of two conjoined individuals working in concert. Fusion of the nuclei for sexual reproduction will only take place when the actual spores are produced.

View of a basidium (left) and ascus (right), from here.

Within the dikaryotic fungi there are two main lineages, the ascomycetes and the basidiomycetes. The two are distinguished by how their spores are produced, as shown in the photos above: in ascomycetes, groups of spores are produced within a sac called an ascus, while in basidiomycetes they are produced on top of a ‘pedestal’ called a basidium. Ascomycetes are by far the more diverse of the two lineages, but basidiomycetes (such as mushrooms) probably include more familiar members because less ascomycetes produce large visible fruiting bodies (examples of well-known ascomycetes include morels, truffles and most lichens). Within the basidiomycetes, most macroscopic forms belong to a group called the Agaricomycotina or hymenomycetes. The other basidiomycete lineages include parasitic forms such as rusts and smuts, and a wide range of yeasts.

Chantarelles Cantharellus cibarius, photographed by Strobilomyces. Chantarelles differ from mushrooms in lacking true gills; instead, they have ‘false gills’ formed from folds of the reproductive membranes that are not divided from the main fruiting body.

These distinctions were recognised some time ago, but recent years have seen a shake-up within the Agaricomycotina. Earlier authors divided this group between the heterobasidiomycetes, in which the basidia are divided by internal septa, and the homobasidiomycetes, in which the basidia are not divided. Mushrooms belong to the homobasidiomycetes; heterobasidiomycetes are a bit more obscure, but representatives include ‘jelly fungi’ with gelatinous fruiting bodies, such as the edible wood-ear fungus Auricularia auricula-judae. Phylogenetic studies have confirmed, however, that the heterobasidiomycetes are paraphyletic with regard to the homobasidiomycetes (Hibbett 2006). What was perhaps more surprising, though, is that the homobasidiomycetes appear to be polyphyletic: one basal clade, the Cantharellales (including the chanterelles and hedgehog fungi), includes both ‘heterobasidiomycetes’ and ‘homobasidiomycetes’, and is phylogenetically distinct from the clade including the remaining ‘homobasidiomycetes’. As a result, recent authors have mostly not recognised homobasidiomycetes as a formal taxon, but instead referred to the clade of Agaricomycetes including homobasidiomycetes and a few closely related heterobasidiomycetous taxa.

The flower fungus Aseroe rubra, photographed by Hugh Smith.

As well as familiar gilled mushrooms, Agaricomycetes include such diverse forms as puffballs, stinkhorns, bracket fungi, earthstars, boletes,… Some of the more unusual forms even include false truffles, or marine taxa producing underwater fruiting bodies (Hibbett 2007). Molecular phylogenetic studies have recognised a dozen or more major lineages within the Agaricomycetes, some of which were unexpected. Many of the latter include lineages dominated by so-called resupinate forms that do not form well-developed fruiting bodies but instead generally form an undifferentiated crust; molecular studies have revealed a previously unsuspected diversity among such morphologically simple taxa. Phylogenetic studies have also confirmed the polyphyletic origins of the so-called ‘gasteromycetes’, forms such as puffballs in which spores are produced internally within the fruiting body and only released when the tissues of the fruiting body break down. Other results, however, have corroborated morphological expectations: one prominent example being the Russulales, a lineage whose members produce fruiting bodies varying from mushroom-like to truffle-like, but united by the production of latex within the fruiting bodies giving them a distinct chalky texture.

Giant puffball Calvatia gigantea, from here.
Systematics of Agaricomycetes
<==Agaricomycetes (see below for synonymy)
    |--Basidiodendron Rick 1938BH05, KC01
    |    `--B. caesiocinereumBH05
    `--+--+--Heterochaete Pat. 1892BH05, KC01 [incl. Hirneolina (Pat.) Bres. 1905KC01]
       |  `--+--Exidia Fr. 1822BH05, KC01 (see below for synonymy)
       |     |    |--E. glandulosaLK04
       |     |    `--E. thuretianaBH05
       |     `--+--+--Ceratosebacina Roberts 1993BH05, KC01
       |        |  |    `--C. calosporaBH05
       |        |  `--Exidiopsis (Bref.) Møller 1895BH05, KC01
       |        |       |--E. calceaBH05
       |        |       `--E. gloeophoraBH05
       |        `--AuriculariaceaeKC01
       |             |--Anthoseptobasidium Rick 1943KC01
       |             |--Mylittopsis Pat. 1895KC01
       |             |--Neotyphula Wakef. 1934KC01
       |             |--Paraphelaria Corner 1966KC01
       |             `--Auricularia Bull. ex Juss. 1789BH05, KC01 (see below for synonymy)
       |                  |--‘Hirneola’ affinis (Jungh.) Bres. 1910SS16
       |                  |--A. auricula-judaeBH05 [=Hirneola auricula-judaeA27]
       |                  `--A. polytrichaSL02 [=Hirneola polytrichaT-W89]
       `--+--Pseudohydnum Karst. 1868BH05, KC01 [=Tremellodon (Pers.) Fr. 1874KC01; incl. Hydnogloea Curr. 1871KC01]
          |    `--P. gelatinosumBH05
             |  `--SebacinalesAS12
             |       |  i. s.: Piriformospora Verma, Varma et al. 1998BH05, KC01
             |       |           `--P. indicaBH05
             |       `--SebacinaceaeBH05
             |            |--Sebacina Tul. & Tul. 1871 (see below for synonymy)KC01
             |            `--Serendipita Roberts 1993BH05, KC01
             |                 `--S. vermifera [=Sebacina vermifera]BH05
                |  `--+--+--ThelephoralesBH05
                |     |  `--+--+--‘Gloeophyllum’ odoratumBH05
                |     |     |  `--Heliocybe Redhead & Ginns 1985BH05, KC01
                |     |     |       `--H. sulcataBH05
                |     |     `--+--Veluticeps (Cooke) Pat. 1894BH05, KC01 [incl. Chaetocarpus Karst. 1889 non Thwaites 1849KC01]
                |     |        |    `--V. berkeleyiBH05
                |     |        `--+--Neolentinus Redhead & Ginns 1985BH05, KC01
                |     |           |    |--N. dactyloidesBH05
                |     |           |    |--N. kauffmaniiBH05
                |     |           |    `--N. lepideusBH05
                |     |           `--Gloeophyllum Karst. 1882BH05, KC01 (see below for synonymy)
                |     |                |--G. abietinumNN95
                |     |                |--G. erubescens (Berkeley) Popoff 2003 (see below for synonymy)P03
                |     |                |--G. sepiariumBH05
                |     |                |--G. striatumAHR03
                |     |                `--G. trabeumBH05
                |     `--+--AgaricomycetidaeBH05
                |        `--+--VuilleminialesBH05
                |           `--+--+--HericiaceaeBH05
                |              |  `--RussulalesBH05
                |              `--Stereaceae [Aleurodiscaceae, Stereales]KC01
                |                   |--AleurodiscusBH05
                |                   |--Acanthophysellum Parmasto 1967KC01
                |                   |--Aleurocystis Lloyd ex Cunn. 1956KC01
                |                   |--Amylohyphus Ryvarden 1978KC01
                |                   |--Amylosporomyces Rattan 1977KC01
                |                   |--Chaetoderma Parmasto 1968KC01
                |                   |--Conferticium Hallenb. 1980KC01
                |                   |--Coniophorafomes Rick 1934KC01
                |                   |--Dextrinocystis Gilb. & Blackw. 1988KC01
                |                   |--Scotoderma Jülich 1974KC01
                |                   |--Scytinostromella Parmasto 1968KC01
                |                   `--Aleurobotrys Boidin 1986KC01
                |                        `--A. botryosusLK04
                   |  `--Kavinia Pilát 1938BH05, KC01 [incl. Hydnocristella Petersen 1971KC01]
                   |       |--K. alboviridisBH05
                   |       `--K. himantiaBH05
                   `--+--+--Resinicium Parmasto 1968BH05, KC01
                      |  |    `--R. meridionaleBH05
                      |  `--Hymenochaetales [Schizoporaceae, Xanthochroales]KC01
                      |       |  i. s.: Echinoporia Ryvarden 1980 [incl. Echinodia Pat. 1919]KC01
                      |       |         Leucophellinus Bondartsev & Singer 1944 [incl. Oxyflavus Ryvarden 1973]KC01
                      |       |         Paratrichaptum Corner 1987KC01
                      |       |         Poriodontia Parmasto 1982KC01
                      |       |         Rogersella Liberta & Navas 1978KC01
                      |       |         Schizopora Velen. 1922KC01
                      |       |         Peniophorella Karst. 1889 (n. d.)JM17, KC01
                      |       |--HyphodontiaBH05
                      |       `--+--Subulicium Hjortstam & Ryvarden 1979BH05, KC01
                      |          `--Oxyporus (Bourdot & Galzin) Donk 1933BH05, KC01
                      |               |--O. latemarginatusBH05
                      |               `--O. populinusBH05
                      `--+--Sistotremastrum Erikss. 1958BH05, KC01
                         |    `--S. niveocremeum [=Paullicorticium niveocremeum]BH05
                         `--+--Tubulicium Oberw. 1965BH05, KC01 [=Tubulixenasma Parmasto 1965KC01]
                            |    |--T. vermiculareBH05
                            |    `--T. vermiferumBH05
                            `--+--Porpomyces Jülich 1982BH05, KC01
                               |    `--P. mucidusBH05
                               `--+--Subulicystidium Parmasto 1968BH05, KC01 [incl. Aegeritina Jülich 1984KC01]
                                  |    `--S. longisporumBH05
                                  `--+--‘Hyphodontia’ gossypinaBH05
                                     `--Trechispora Karst. 1890BH05, KC01 (see below for synonymy)
                                          |  i. s.: T. niveaBH05
                                          |--T. farinaceaBH05
                                          `--+--T. regularisBH05
                                             `--+--+--T. araneosaBH05
                                                |  `--+--T. confinisBH05
                                                |     `--T. subsphaerosporaBH05
                                                `--+--T. hymenocystisBH05
                                                   `--+--T. incisaBH05
                                                      `--T. kavinioidesBH05

Agaricomycetes incertae sedis:
  Tremellodendropsis (Corner) Crawford 1954LK04, KC01 (see below for synonymy)
    `--T. tuberosaLK04
  Hyaloria Møller 1895AS12, KC01 [Hyaloriaceae]
  Palaeoclavaria Poinar & Brown 2003 [Palaeoclavariaceae]PB03
    `--*P. burmitis Poinar & Brown 2003PB03
  Protomycena Hibbett, Grimaldi & Donoghue 1997 (nom. inv.)HBW03, KC01
    `--P. electraHBW03
  Archaeomarasmius Hibbett, Grimaldi & Donoghue 1997HBW03, KC01
    `--A. leggetiHBW03
  Cystidiophora castaneaBH05
  Palaeancistrus Dennis 1970KC01
    `--P. martinii Dennis 1970T93
  Acantholichen Jørg. 1998KC01
  Ambivina Katz 1974KC01
  Amylobasidium Ginns 1988KC01
  Bourdotia (Bres.) Trotter 1925 non Dall 1901 (ICZN)KC01
  Tremellodendron Atk. 1902 [=Collodendrum Clem. 1909]KC01
  Corticirama Pilát 1957KC01
  Craterocolla Bref. 1888 [incl. Ditangium Karst. 1867, Poroidea Göttinger ex Winter 1884]KC01
  Dendrophysellum Parmasto 1968KC01
  Dextrinodontia Hjortstam & Ryvarden 1980KC01
  Ductifera Lloyd 1917 [incl. Gloeostromera Ervin 1956]KC01
  Efibulobasidium Wells 1975KC01
  Eichleriella Bres. 1903KC01
  Endoperplexa Roberts 1993KC01
  Fibulosebacea Wells & Raitv. 1987KC01
  Stypella Møller 1895 (see below for synonymy)KC01
  Tremiscus (Pers.) Lév. 1846 [incl. Gyrocephalus Bref. 1888]KC01
  Heteroscypha Oberw. & Agerer 1979KC01
  Licrostroma Lenke 1964KC01
  Melzerodontia Hjortstam & Ryvarden 1980KC01
  Merulicium Erikss. & Ryvarden 1976KC01
  Microsebacina Roberts 1993KC01
  Mutatoderma (Parmasto) Gómez 1976KC01
  Mycolindtneria Rauschert 1987KC01
  Nothocorticium Gresl. & Rajchenb. 1999KC01
  Papyrodiscus Reid 1979KC01
  Patouillardina Bres. 1906 (see below for synonymy)KC01
  Protodaedalea Imazeki 1955KC01
  Protohydnum Møller 1895KC01
  Protomerulius Møller 1895KC01
  Pseudostypella McNabb 1969KC01
  Renatobasidium Hauerslev 1993KC01
  Ripexicium Hjortstam 1995KC01
  Tremellacantha Jülich 1980KC01
  Tremelloscypha Reid 1979KC01

Agaricomycetes [Aphyllophorales, Auriculariales, Auromycetidae, Clavomycetidae, Corticiaceae, Exidiaceae, Ganodermatales, Gasteromyctes, Homobasidiomycetes, Homobasidiomycetidae, Hymenomycetidae, Perenniporiales, Pileomycetidae]

Auricularia Bull. ex Juss. 1789BH05, KC01 [=Auricula Battarra ex Kuntze 1891 nec Lamarck 1799 (ICZN) nec Castrac. 1873 (nom. cons.)KC01; incl. Auriculariella (Sacc.) Clem. 1909KC01, Conchites Paulet 1791KC01, Hirneola Fr. 1848 (nom. cons.)KC01, Oncomyces Klotzsch 1843KC01, Patila Adans. 1763KC01, Seismosarca Cooke 1889KC01]

Exidia Fr. 1822BH05, KC01 [=Spicularia Chevall. 1826 non Pers. 1822KC01; incl. Myxarium Wallr. 1833KC01, Tremellochaete Raitv. 1964KC01, Ulocolla Bref. 1888KC01]

Gloeophyllum Karst. 1882BH05, KC01 [incl. Anisomyces Pilát 1940 non Theiss. & Syd. 1914KC01, Ceratophora Humb. 1793KC01, Griseoporia Ginns 1984KC01, Lenzitina Karst. 1889KC01, Phaeocoriolellus Kotl. & Pouzar 1957KC01, Reisneria Velen. 1922KC01, Serda Adans. 1763KC01, Sesia Adans. 1763KC01, Stiptophyllum Ryvarden 1973P03; Gloeophyllaceae, Gloeophyllales]

Gloeophyllum erubescens (Berkeley) Popoff 2003 [=Daedalea erubescens Berkeley 1840, *Stiptophyllum erubescens (Berkeley) Ryvarden 1973, Xerotinus erubescens (Berkeley) Wright & Deschamps 1977]P03

Patouillardina Bres. 1906 [=Meliolinopsis Beeli 1920; incl. Atractobasidium Martin 1935, Protograndinia Rick 1933]KC01

Sebacina Tul. & Tul. 1871 [incl. Atkinsonia Lloyd 1916 non Muell. 1865, Cristella Pat. 1887, Flahaultia Arnaud 1952, Opadorhiza Andersen & Moore 1996, Soppittiella Massee 1892, Sulphurina Pilát 1953]KC01

Stypella Møller 1895 [incl. Gloeosebacina Neuhoff 1924, Heterochaetella (Bourdot) Bourdot & Galzin 1928, Protodontia Höhn. 1907]KC01

Trechispora Karst. 1890BH05, KC01 [incl. Cristelloporia Johans. & Ryvarden 1979KC01, Echinotrema Park.-Rhodes 1955KC01, Febriciellum Erikss. & Ryvarden 1975KC01, Fibuloporia Bondartsev & Singer 1944KC01, Tomentella Karst. 1889 non Pat. 1887KC01; Trechisporales]

Tremellodendropsis (Corner) Crawford 1954LK04, KC01 [incl. Polyozus Karst. 1881KC01, Pseudotremellodendron Reid 1957KC01; Tremellodendropsidaceae, Tremellodendropsidales]

*Type species of generic name indicated


[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.

[AHR03] Aime, M. C., T. W. Henkel & L. Ryvarden. 2003. Studies in neotropical polypores 15: New and interesting species from Guyana. Mycologia 95 (4): 614–619.

[A27] Andersen, J. C. 1927. Popular names of New Zealand plants. Part 2. Transactions and Proceedings of the New Zealand Institute 57: 905–977.

[BH05] Binder, M., D. S. Hibbett, K.-H. Larsson, E. Larsson, E. Langer & G. Langer. 2005. The phylogenetic distribution of resupinate forms across the major clades of mushroom-forming fungi (Homobasidiomycetes). Systematics and Biodiversity 3 (2): 113–157.

Hibbett, D. S. 2006. A phylogenetic overview of the Agaricomycotina. Mycologia 98 (6): 917–925.

Hibbett, D. S. 2007. After the gold rush, or before the flood? Evolutionary morphology of mushroom-forming fungi (Agaricomycetes) in the early 21st century. Mycological Research 111: 1001–1018.

[HBW03] Hibbett, D. S., M. Binder & Z. Wang. 2003. Another fossil agaric from Dominican amber. Mycologia 95 (4): 685–687.

[JM17] Justo, A., O. Miettinen, D. Floudas, B. Ortiz-Santana, E. Sjökvist, D. Lindner, K. Nakasone, T. Niemelä, K.-H. Larsson, L. Ryvarden & D. S. Hibbett. 2017. A revised family-level classification of the Polyporales (Basidiomycota). Fungal Biology 121: 798–824.

[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).

[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.

[NN95] Neubert, H., W. Nowotny, K. Baumann & H. Marx. 1995. Die Myxomyceten: Deutschlands und des angrenzenden Alpenraumes unter besonderer Berücksichtigung Österreichs vol. 2. Physarales. Karlheinz Baumann Verlag: Gomaringen.

[PB03] Poinar, G. O., Jr & A. E. Brown. 2003. A non-gilled hymenomycete in Cretaceous amber. Mycological Research 107 (6): 763–768.

[P03] Popoff, O. F. 2003. Notes on Daedalea erubescens, Hexagonia decipiens and the Phaeotrametaceae. Mycotaxon 87: 103–108.

[SL02] Schweigkofler, W., K. Lopandic, O. Molnár & H. Prillinger. 2002. Analysis of phylogenetic relationships among Ascomycota with yeast phases using ribosomal DNA sequences and cell wall sugars. Organisms Diversity & Evolution 2: 1–17.

[SS16] Sydow, H., & P. Sydow. 1916. Fungi papuani. Die von C. Ledermann in Neu-Guinea gesammelten Pilze. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 54: 246–261.

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

[T-W89] Tenison-Woods, J. E. 1889. On the vegetation of Malaysia. Proceedings of the Linnean Society of New South Wales, series 2, 4 (1): 9–106, pls 1–9.

Leave a comment

Your email address will not be published. Required fields are marked *