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Mushroom Taxonomy: The Big Picture

by Michael Kuo

I frequently receive e-mails from frantic biology students who have been asked to discover the kingdom, phylum, class, order, family, genus, and species of a certain mushroom. Here, with the student's typo included, is the most entertaining example I've received so far:

Recently in my biology class we were asked to chose an orgasm. I chose the Armillariella ostoyae. My professor wants us to find the kingdom, phylum, class, order, family, genus, species, and or variety. I know the kingdom, genus and species. I have had difficulty finding the phylum, class, order and family. Do you know of any good sites that can help me with my research. Thank you very much.

Aside from recommending that the student might want to find a new professor, I replied that the taxonomical hierarchy for Armillaria ostoyae is:

    Fungi: Basidiomycota: Basidiomycetes: Agaricales: Tricholomataceae: Armillaria: ostoyae

. . . in the traditional, and now probably outdated, system. Armillaria has been reconceived within the past few years, resulting in the elimination of what the student called "Armillariella," and placing the genus in the Marasmiaceae rather than the Tricholomataceae; also, there is debate about whether or not the kingdom and phylum distinctions should be made at some other level in the hierarchy.

    Note, 2011: This text was originally written in 2003. As a further demonstration of some of my points in this essay, the genus Armillaria has now been placed in the Physalacriaceae, and the species "Armillaria ostoyae" no longer exists, since it has been synonymized with an older species (Armillaria solidipes)!

But uncertainty is not what professors want on homework assignments. The problem is that there is no "correct" answer to the professor's question. Or, better said, the answer to the question changes constantly, and has been changing ever since Linnaeus started using Latin names to arrange organisms.

Though it is a fact usually unobserved in introductory biology classes, taxonomy does not represent organisms. Rather, taxonomy represents how we perceive and organize organisms. This is a very important difference. It is the difference, for example, between what happened at the scene of the crime, and what the witness saw happen at the scene of the crime--and anyone who has ever watched a courtroom drama knows how different these two things can be.

In my field (I am an English teacher), the rules of grammar and punctuation are seen by most teachers as unchanging and universal. Students are "wrong" if they omit the apostrophe from don't, or write "Everyday someone gets their lunch." Yet there was a time--not that long ago, from a historical perspective--when dont was perfectly correct, and the time is coming (or is already here) when this use of everyday and their is correct. People of my mother's generation physically cringe when they hear "their" used like this. People of my generation notice a problem, but use it anyway as a substitute for the sexist "his." My students don't even notice. Within my lifetime, the language has changed, as a result of a change in our culture: we became more aware of sexism, and less comfortable using masculine pronouns as universal pronouns.

With grammar and punctuation, however, the rule makers usually lag far behind the general population. This is because the rule makers (the authors and publishers of dictionaries and grammar handbooks) are conservative by nature, and often see themselves as corrective agents, holding back the masses and saving them from their mistakes. But with taxonomy, things are reversed. It is the mycologists, in the case of mushrooms, who are constantly changing things, and the general population that lags behind. Thus, I must provide the biology student above with an answer I know to be incorrect, knowing that her professor is likely working from outdated information.

Once, mushroom taxonomy was an arrangement of mushrooms based on their physical appearance. This one had gills, so it belonged in a group with other gilled mushrooms, while another mushroom, this one with pores, belonged in a different group. For well over a hundred years, advances in mushroom taxonomy simply represented more careful attention to the physical features of the mushrooms--and, importantly, the fact that more and more mushrooms from around the world were being sent to scientists in northern Europe. These scientists began to discover that closer examination revealed other groupings. Some of the gilled mushrooms had white spore prints, for example, and gills that were attached to the stem. New families and genera were named; species were placed in the hierarchy accordingly.

Then, roughly a hundred years ago, scientists began looking at mushrooms with microscopes. Some mycologists had been doing so earlier, but the hegemony of microscope mycology didn't take hold until the 20th century. As a result, new groupings emerged. These mushrooms, for example, had ornamented spores, indicating that they formed a group separate from other mushrooms that looked more or less the same to the naked eye, but had smooth spores. As microscopes got better and better, more taxonomical changes were made.

It is important to recall that the mushrooms themselves did not change during this brief history; what changed was the way we examined them. New technologies and methods of analysis--like studies of chemical composition, mating studies, and (especially) DNA analysis--are hegemonic these days, and they are resulting in radical changes in mushroom taxonomy. Groups that we once thought were related, based on physical appearance or microscopic features, are turning out to be unrelated. But it is likely--I would say it is a certainty--that future mycologists will decide our contemporary taxonomic arrangements are inaccurate.

I offer these comments by way of introducing the table below, which represents how mycologists currently see taxonomical relationships between mushrooms. I have culled the information from Ainsworth & Bisby's 2008 Dictionary of the Fungi (see the notes below for a complete citation), and I have included only "mushroom" taxonomy--omitting the details on rusts, yeasts, lichens, molds, and so on. The editors of the Dictionary, of course, compiled information from peer-reviewed papers published in scientific journals; it should come as no surprise that editing such a compilation involves attempting to "standardize" things that have not yet become standards, resolving taxonomical conflicts that are often hotly debated, and so on. Yet Ainsworth & Bisby's Dictionary has become more or less the definitive standard for mushroom taxonomy; for better or worse, the biology student must consult this source to get the "best" current answer to a taxonomy question.


The Taxonomic Hierarchy of Kingdom Fungi

. . . based on Ainsworth & Bisby's 2008 Dictionary of the Fungi (10th. edition)

Only genera treated at MushroomExpert.Com are included. See the notes at the bottom of the page for additional information and suggestions.

      
Phylum: Ascomycota
 Subphyllum: Pezizomycotina
  Class: Arthoniomycetes (lichens . . .)
  Class: Dothideomycetes
   (saprobes, parasites [Apiosporina morbosa], lichens, dung lovers . . . )
  Class: Eurotiomycetes (includes Penicillium . . . )
  Class: Laboulbeniomycetes (insect parasites and others . . . )
  Class: Lecanoromycetes (lichens . . . )
  Class: Leotiomycetes (inoperculate Discomycetes + powdery mildews)
   Order: Cyttariales
   Order: Erysiphales (powdery mildews)
   Order: Helotiales
    Family: Ascocorticiaceae
    Family: Dermateaceae
     Genera treated: Chlorosplenium (see C. chlora)
    Family: Heliotiaceae
     Genera treated: Hymenoscyphus (see H. fructigenus); possibly also Ascocoryne (see A. sarcoides), Bisporella (see B. citrina), Chlorociboria (see C. aeruginascens), and Ionomidotis (see I. irregularis)
    Family: Hemiphacidiaceae
     Genera treated: Chlorencoelia (see C. torta)
    Family: Hyaloscyphaceae
    Family: Loramycetaceae
    Family: Phacidiaceae
    Family: Rustroemiaceae
    Family: Sclerotiniaceae
    Family: Vibrisseaceae
   Order: Leotiales
    Family: Bulgariaceae
     Genera treated: Bulgaria (see B. inquinans)
    Family: Leotiaceae
     Genera treated: Leotia (see L. lubrica), Microglossum (see M. viride)
   Order: Rhytismatales
    Family: Ascodichaenaceae
    Family: Cudoniaceae
     Genera treated: Cudonia (see C. circinans), Spathularia (see S. flavida), Spathulariopsis (see S. velutipes)
    Family: Rhytismataceae
     Genera treated: Colpoma (see C. quercinum)
  Class: Pezizomycetes
   Order: Pezizales
    Family: Ascobolaceae
    Family: Ascodesmidaceae
    Family: Caloscyphaceae
    Family: Carbomycetaceae
    Family: Chorioactidaceae
     Genera treated: Chorioactis (see C. geaster), Wolfina (see W. aurantiopsis)
    Family: Discinaceae
     Genera treated: Gyromitra
    Family: Glaziellaceae
    Family: Helvellaceae
     Genera treated: Helvella
    Family: Karstenellaceae
    Family: Morchellaceae
     Genera treated: Disciotis (see D. venosa), Morchella, Verpa (see V. bohemica)
    Family: Pezizaceae
     Genera treated: Pachyella (see P. clypeata), Peziza (see P. repanda), Sarcosphaera (see S. coronaria)
    Family: Pyronemataceae
     Genera treated: Aleuria (see A. aurantia), Cheilymenia (see C. stercorea), Geopora (see G. cooperi), Humaria (see H. hemisphaerica), Jafnea (see J. semitosta), Otidea (see O. onotica), Scutellinia (see S. scutellata), Sphaerosporella (see S. brunnea), Tarzetta (see T. bronca)
    Family: Rhizinaceae
    Family: Sarcoscyphaceae
     Genera treated: Microstoma (see M. floccosum), Sarcoscypha
    Family: Sarcosomataceae
     Genera treated: Galiella (see G. rufa), Urnula (see U. craterium)
    Family: Tuberaceae
  Class: Sordariomycetes
   [Most "Pyrenomycetes," in 15 orders, 64 families, and over 1000 genera. Genera treated: Hypomyces, Xylaria, Cordyceps (see C. militaris), Camarops (see C. petersii)]
  Class: Uncertain
   Order: Uncertain
    Family: Geoglossaceae
     Genera treated: Geoglossum (see G. nigritum)
 Subphyllum: Saccharomycotina (yeasts . . . )
 Subphyllum: Taphrinomycotina (galls, witches' brooms, Neolecta . . . )

Phylum: Basidiomycota
 Subphyllum: Agaricomycotina
  Class: Dacrymycetes
   Order: Dacrymycetales
    Family: Dacrymycetaceae
     Genera treated: Calocera (see C. cornea), Dacryopinax (see D. elegans)
  Class: Tremellomycetes
   Order: Cystofilobasidiales
    Family: Cystofilobasidiaceae
   Order: Filobasidiales
    Family: Filobasidiaceae
   Order: Tremellales
    Family: Carcinomycetaceae
     Genera treated: Syzygospora (see S. mycetophila)
    Family: Cuniculitremaceae
    Family: Hyaloriaceae
    Family: Phragmoxenidiaceae
    Family: Rhynchogastremataceae
    Family: Sirobasidiaceae
    Family: Tetragoniomycetaceae
    Family: Tremellaceae
     Genera treated: Tremella (see T. mesenterica)
  Class: Agaricomycetes
   Order: Agaricales
    Family: Agaricaceae
     Genera treated: Agaricus, Battarrea (see B. phalloides), Bovista (see B. longispora), Calvatia (see C. craniiformis), Chlorophyllum (see C. molybdites), Coprinus (see C. comatus), Crucibulum (see C. laeve), Cyathus (see C. striatus), Cystoderma, Cystolepiota (see C. seminuda), Floccularia, Lepiota (see L. cristata), Leucoagaricus (see L. naucinus), Leucocoprinus (see L. birnbaumii), Lycoperdon (see L. pulcherrimum), Macrolepiota (see M. procera), Morganella (see M. pyriformis), Mycenastrum (see M. corium), Nidularia (see N. pulvinata), Podaxis (see P. longii), Ripartitella (see R. brasiliensis), Tulostoma (see T. lloydii), Vascellum (see V. curtisii)
    Family: Amanitaceae
     Genera treated: Amanita, Limacella
    Family: Amylocorticiaceae
    Family: Bolbitiaceae
     Genera treated: Bolbitius (see B. vitellinus), Conocybe (see C. albipes), Galeropsis (see Gastrocybe lateritia)
    Family: Broomeiaceae
    Family: Clavariaceae
     Genera treated: Clavaria (see C. vermicularis), Clavulinopsis (see C. laeticolor), Ramariopsis (see R. kunzei)
    Family: Cortinariaceae
     Genera treated: Cortinarius
    Family: Cyphellaceae
    Family: Cystostereaceae
    Family: Entolomataceae
     Genera treated: Clitopilus (see C. prunulus), Entoloma, Rhodocybe (see R. mundula)
    Family: Fistulinaceae
     Genera treated: Fistulina (see F. hepatica)
    Family: Gigaspermaceae
    Family: Hemigasteraceae
    Family: Hydnangiaceae
     Genera treated: Laccaria
    Family: Hygrophoraceae
     Genera treated: Ampulloclitocybe (see A. clavipes), Chrysomphalina (see C. chrysophylla), Cuphophyllus (see C. pratensis), Gliophorus (see G. psittacinus), Hygrocybe (see H. conica), Hygrophorus (see H. russula)
    Family: Inocybaceae
     Genera treated: Crepidotus, Flammulaster (see F. erinaceella), Inocybe, Simocybe (see S. centunculus); possibly Panaeolus
    Family: Limnoperdaceae
    Family: Lyophyllaceae
     Genera treated: Asterophora (see A. lycoperdoides), Calocybe (see C. carnea), Hypsizygus (see H. ulmarius), Lyophyllum (see L. decastes)
    Family: Marasmiaceae
     Genera treated: Baeospora (see B. myosura), Clitocybula (see C. abundans), Connopus (see C. acervatus), Crinipellis (see C. zonata), Gerronema (see G. strombodes), Gymnopus (see G. dryophilus), Macrocystidia (see M. cucumis), Marasmiellus (see M. candidus), Marasmius (see M. rotula), Megacollybia, Micromphale (see M. perforans), Mycetinis (see M. scorodonius), Omphalotus (see O. illudens), Rhodocollybia (see R. maculata), Tetrapyrgos (see T. nigripes)
    Family: Mycenaceae
     Genera treated: Mycena, Panellus (see P. stipticus); possibly Xeromphalina (see X. kauffmanii)
    Family: Niaceae
    Family: Phelloriniaceae
    Family: Physalacriaceae
     Genera treated: Armillaria, Cyptotrama (see C. asprata), Flammulina, Rhizomarasmius, Rhodotus (see R. palmatus), Xeruloid Mushrooms (including Hymenopellis and Paraxerula)
    Family: Pleurotaceae
     Genera treated: Hohenbuehelia, Pleurotus (see P. ostreatus)
    Family: Pluteaceae
     Genera treated: Pluteus, Volvariella
    Family: Psathyrellaceae
     Genera treated: Coprinellus (see C. disseminatus), Coprinopsis (see C. atramentaria), Lacrymaria (see L. velutina), Parasola (see P. plicatilis), Psathyrella; possibly Panaeolus
    Family: Pterulaceae
    Family: Schizophyllaceae
     Genera treated: Schizophyllum (see S. commune)
    Family: Stephanosporaceae
    Family: Strophariaceae
     Genera treated: Agrocybe, Galerina (see G. marginata), Hebeloma, Hemipholiota (see H. populnea), Hypholoma, Kuehneromyces (see K. mutabilis), Leratiomyces, Pholiota, Psilocybe; possibly Gymnopilus
    Family: Tapinellaceae
     Genera treated: Tapinella (see T. panuoides)
    Family: Tricholomataceae
     Genera treated: Callistosporium (see C. luteo-olivaceum), Catathelasma, Caulorhiza (see C. umbonata), Clitocybe, Collybia (see C. cirrhata), Dendrocollybia (see D. racemosa), Infundibulicybe, Leucopaxillus, Leucopholiota, Macrocybe (see M. titans), Melanoleuca, Omphalina (see O. epichysium), Resupinatus (see R. alboniger), Tricholoma
    Family: Typhulaceae
     Genera treated: Macrotyphula (see M. juncea).
    Family: Uncertain
     Genera treated: Phyllotopsis (see P. nidulans), Rickenella (see R. fibula; also possibly in Hymenochaetales); Tricholomopsis (see T. decora)
   Order: Atheliales
    Family: Atheliaceae
   Order: Auriculariales
    Family: Auriculariaceae
     Genera treated: Auricularia (see A. auricula), Exidia (see E. glandulosa)
    Family: Uncertain
     Genera treated: Ductifera (see D. pululahuana), Guepinia (see Phlogiotis helvelloides), Pseudohydnum (see P. gelatinosum)
   Order: Boletales
    Family: Boletaceae
     Genera treated: Austroboletus, Boletellus, Boletus, Bothia (see B. castanella), Chalciporus (see C. piperatus), Chamonixia (see C. caespitosa), Harrya (see H. chromapes), Heimioporus, Leccinum, Phylloporus, Pulveroboletus (see P. ravenelii), Retiboletus (see R. ornatipes), Rubroboletus (see R. dupainii), Strobilomyces, Tylopilus, Xanthoconium (see X. purpureum), Xerocomellus (see X. chrysenteron), Xerocomus (see X. subtomentosus)
    Family: Boletinellaceae
     Genera treated: Boletinellus (see B. merulioides)
    Family: Calostomataceae
     Genera treated: Calostoma (see C. cinnabarinum)
    Family: Coniophoraceae
    Family: Diplocystidiaceae
     Genera treated: Astraeus (see A. hygrometricus)
    Family: Gastrosporiaceae
    Family: Gomphidiaceae
     Genera treated: Chroogomphus, Gomphidius
    Family: Gyroporaceae
     Genera treated: Gyroporus
    Family: Hygrophoropsidaceae
     Genera treated: Hygrophoropsis (see H. aurantiaca)
    Family: Paxillaceae
     Genera treated: Paragyrodon (see P. sphaerosporus), Paxillus (see P. vernalis)
    Family: Protogastraceae
    Family: Rhizopogonaceae
    Family: Sclerodermataceae
     Genera treated: Pisolithus (see P. tinctorius), Scleroderma
    Family: Serpulaceae
    Family: Suillaceae
     Genera treated: Suillus
   Order: Cantharellales
    Family: Aphelariaceae
    Family: Botryobasidiaceae
    Family: Cantharellaceae
     Genera treated: Cantharellus (see C. cibarius), Craterellus (see C. fallax)
    Family: Ceratobasidiaceae
    Family: Clavulinaceae
     Genera treated: Clavulina (see C. cristata)
    Family: Hydnaceae
     Genera treated: Hydnum (see H. repandum)
    Family: Tulasnellaceae
   Order: Corticiales
    Family: Corticiaceae
   Order: Geastrales
    Family: Geastraceae
     Genera treated: Geastrum (see G. saccatum)
   Order: Gloeophyllales
    Family: Gloeophyllaceae
     Genera treated: Gloeophyllum (see G. sepiarium)
   Order: Gomphales
    Family: Clavariadelphaceae
     Genera treated: Clavariadelphus (see C. unicolor)
    Family: Gomphaceae
     Genera treated: Gomphus (see G. clavatus), Ramaria (see R. botrytis), Turbinellus (see T. floccosus)
    Family: Lentariaceae
     Genera treated: Lentaria (see L. micheneri)
   Order: Hymenochaetales
    Family: Hymenochaetaceae
     Genera treated: Coltricia (see C. cinnamomea), Inonotus (see I. radiatus), Phellinus (see P. gilvus), Porodaedalea (see P. pini)
    Family: Schizoporaceae
    Family: Uncertain
     Genera treated: Rickenella (see R. fibula; also possibly in Agaricales)
   Order: Hysterangiales
    Family: Gallaceaceae
    Family: Hysterangiaceae
    Family: Mesophelliaceae
    Family: Phallogastraceae
    Family: Trappeaceae
   Order: Phallales
    Family: Clastulaceae
    Family: Phallaceae
     Genera treated: Aseröe (see A. rubra), Blumenavia (see B. angolensis), Clathrus (see C. ruber), Colus (see C. hirudinosus), Ileodictyon (see I. cibarium), Laternea (see L. pusilla), Lysurus (see L. mokusin), Mutinus (see M. elegans), Phallus (see P. impudicus), Pseudocolus (see P. fusiformis), Staheliomyces (see S. cinctus)
   Order: Polyporales
    Family: Cystostereaceae
    Family: Fomitopsidaceae
     Genera treated: Daedalea (see D. quercina), Fomitopsis (see F. pinicola), Ischnoderma (see I. resinosum), Laetiporus (see L. sulphureus), Phaeolus (see P. schweinitzii), Piptoporus (see P. betulinus), Pycnoporellus (see P. alboluteus)
    Family: Ganodermataceae
     Genera treated: Ganoderma (see G. lucidum)
    Family: Grammotheleaceae
    Family: Limnoperdaceae
    Family: Meripilaceae
     Genera treated: Abortiporus (see A. biennis), Grifola (see G. frondosa), Meripilus (see M. giganteus)
    Family: Meruliaceae
     Genera treated: Bjerkandera (see B. adusta), Gloeoporus (see G. dichrous), Irpex (see I. lacteus), Mycorrhaphium (see M. adustum), Phlebia (see P. incarnata), Podoscypha (see P. aculeata), Steccherinum (see S. ochraceum)
    Family: Panaceae
     Genera treated: Panus (see P. conchatus)
    Family: Phanerochaetaceae
     Genera treated: Climacodon (see C. septentrionale)
    Family: Polyporaceae
     Genera treated: Cerrena (see C. unicolor), Coriolopsis (see C. gallica), Cryptoporus (see C. volvatus), Daedaleopsis (see D. confragosa), Fomes (see F. fomentarius), Hapalopilus (see H. nidulans), Lenzites (see L. betulina), Neofavolus (see N. alveolaris), Nigroporus (see N. vinosus), Perenniporia (see P. ohiensis), Polyporus (see P. squamosus), Poronidulus, Pycnoporus (see P. cinnabarinus), Spongipellis (see S. pachyodon), Trametes (see T. versicolor), Trichaptum (see T. biforme), Tyromyces (see T. chioneus)
    Family: Sparassidaceae
     Genera treated: Sparassis (see S. crispa)
    Family: Tubulicrinaceae
    Family: Xenasmataceae
   Order: Russulales
    Family: Albatrellaceae
     Genera treated: Albatrellus (see A. cristatus)
    Family: Amylostereaceae
    Family: Auriscalpiaceae
     Genera treated: Artomyces (see Artomyces pyxidatus), Auriscalpium (see A. vulgare), Lentinellus
    Family: Bondarzewiaceae
     Genera treated: Bondarzewia (see B. berkeleyi), Heterobasidion (see H. annosum)
    Family: Echinodontiaceae
    Family: Hericiaceae
     Genera treated: Hericium
    Family: Hybogasteraceae
    Family: Lachnocladiaceae
    Family: Peniophoraceae
     Genera treated: Peniophora (see P. rufa)
    Family: Russulaceae
     Genera treated: Arcangeliella (see A. desjardinii), Lactarius, Russula
    Family: Stereaceae
     Genera treated: Aleurodiscus (see A. oakesii), Stereum (see S. ostrea), Xylobolus (see X. frustulatus)
   Order: Sebacinales
    Family: Sebacinaceae
     Genera treated: Tremellodendron (see T. schweinitzii)
   Order: Thelephorales
    Family: Bankeraceae
     Genera treated: Boletopsis (see B. leucomelaena), Hydnellum, Phellodon (see P. confluens), Sarcodon (see S. imbricatus)
    Family: Thelephoraceae
     Genera treated: Polyozellus (see P. multiplex), Thelephora (see T. multipartita)
   Order: Trechisporales
    Family: Hydnodontaceae
 
Subphyllum: Pucciniomycotina (rusts . . . see Gymnosporangium juniperi-virginianae)
 Subphyllum: Ustilaginomycotina (smuts . . . )

Phylum: Chytridiomycota (aquatic fungi . . . )
Phylum: Glomeromycota (endomycorrhizal fungi . . . )
Phylum: Microsporidia (spore-forming parasites that lack flagellae . . . )
Phylum: Zygomycota (various saprobes, parasites, and others . . . )

Notes

I have done my best to avoid typing mistakes in the table above, but I ask you to imagine typing "Hypsizygus, Syzygospora, Rhynchogastremataceae," and the like for hours on end with no recourse to a spell-checker. If you find a mistake, please drop me a line; I will appreciate knowing it.

Use Control/F or Command/F in your browser to search for taxa on this page. However, genera are only included if they are treated at MushroomExpert.Com. This is a substantial and unfortunate change since the previous version (2003) of this page, but the 2008 Dictionary no longer includes tables listing the genera included in each family, leaving the researcher no way to get this information without consulting all of the literature the editors of the Dictionary had to consult. Thus, if the genus you are searching is not on this page, it is not one that is treated at MushroomExpert.Com with at least one species page from the genus--or the genus may have been renamed, or collapsed into another genus (as has happened, for example, with the genus Stropharia, which is not listed in the Strophariaceae since it has been found to belong in Psilocybe).


Reference:

Kirk, P.M., P. F. Cannon, D. W. Minter & J. A. Stalpers, eds. (2008). Ainsworth & Bisby's dictionary of the fungi. UK: CAB International. 771 pp.



Cite this page as:

Kuo, M. (2014, March). Mushroom taxonomy: The big picture. Retrieved from the MushroomExpert.Com Web site: http://www.mushroomexpert.com/taxonomy.html

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