Entoloma

{{Short description|Genus of fungi}} {{Automatic taxobox | image = 2005-09-25 Entoloma sinuatum cropped.jpg | image_caption = ''Entoloma sinuatum'' | taxon = Entoloma | authority = (Fr.) P.Kumm. (1871) | type_species = ''Entoloma sinuatum'' | type_species_authority = (Bull.) P.Kumm. (1871) | synonyms = {{Collapsible list|bullets=on |''Agaricus'' trib. ''Entoloma'' Fr. (1838) |''Alboleptonia'' Largent & R.G.Benedict (1970) |''Arenicola'' Velen. (1947) |''Calliderma'' (Romagn.) Largent (1994) |''Claudopus'' Gillet (1876) |''Clitopiloides'' (Romagn.) Largent (1994) |''Eccilia'' (Fr.) P.Kumm. (1871) |''Fibropilus'' (Noordel.) Largent (1994) |''Inocephalus'' (Noordel.) P.D.Orton (1991) |''Leptonia'' (Fr.) P.Kumm. (1871) |''Leptoniella'' Earle (1909) |''Nigropogon'' Coker & Couch (1928) |''Nolanea'' (Fr.) P.Kumm. (1871) |''Omphaliopsis'' (Noordel.) P.D.Orton (1991) |''Paraeccilia'' Largent (1994) |''Paraleptonia'' (Romagn. ex Noordel.) P.D.Orton (1991) |''Pouzarella'' Mazzer (1976) |''Richoniella'' Costantin & L.M.Dufour (1900) |''Rhodocybella'' T.J.Baroni & R.H.Petersen (1987) |''Rhodogaster'' E.Horak (1964) |''Rhodophyllus'' Quél. (1886) (nom. illegit.) |''Trichopilus'' (Romagn.) P.D.Orton (1991) }} | subdivision_ranks = Species | subdivision = 2031 species<ref name="CatofLife"/> }}

'''''Entoloma''''' is a genus of fungi in the order Agaricales, with the common name of '''pinkgills'''. The basidiocarps (fruit bodies) are typically agaricoid (gilled mushrooms), though a minority are gasteroid. All have salmon-pink basidiospores which colour the gills at maturity and are angular (polyhedral) under a microscope. The genus is large, with almost 2000 species worldwide. Most species are saprotrophic, but some are ectomycorrhizal, and a few are parasitic on other fungi. The type, ''Entoloma sinuatum'', is one of several ''Entoloma'' species that are poisonous, typically causing mild to severe gastrointestinal illness.

==Taxonomy== ===History=== In 1838 the Swedish mycologist Elias Magnus Fries classified all pink-spored, gilled fungi into "tribes" or "subtribes", placing those with a ''Tricholoma''-like shape and gills attached to the stem into tribe ''Entoloma''. The small subtribe ''Leptonia'' had convex fleshy membranaceous caps, the subtribe ''Nolanea'' were slender fungi with bell-shaped caps and hollow stems, and the subtribe ''Eccilia'' had umbilicate caps and adnate gills.<ref name="Fries 1838"/> In 1871 German mycologist Paul Kummer raised these tribes and subtribes to genera.<ref name=Kummer/> Additional genera were added by subsequent authors.<ref name="Orton1991"/><ref name="Orton1991b"/><ref name="Largent1994"/> Following this classification system, ''Entoloma'' has a restricted meaning and has sometimes been referred to as ''Entoloma'' ''sensu stricto.''<ref name="Largent1994"/>

In 1886 French mycologist Lucien Quélet united all the fungi with pinkish-red adnate or sinuate gills and angular spores into a new genus ''Rhodophyllus'' (meaning "pink gill").<ref name="Quélet 1886"/> Because his new genus included the earlier name ''Entoloma'', ''Rhodophyllus'' is illegitimate, as noted by Donk,<ref name = "Donk62"/> and ''Entoloma'' was subsequently adopted to cover all the pink-spored agarics with angular spores.<ref name=Noor92/> Following this classification system, ''Entoloma'' has a broad meaning and has sometimes been referred to as ''Entoloma'' ''sensu lato''.<ref name="Orton1991"/><ref name=Noor92/> The synonyms listed here are applicable to ''Entoloma'' sensu lato.

These two classification systems continue to co-exist, with those taxonomists favouring a broad generic concept following Quélet, and the others a narrow generic concept following Kummer.

===Current status=== Recent molecular research, based on cladistic analysis of DNA sequences, has shown that ''Entoloma'' ''sensu lato'' is monophyletic (a natural grouping), whereas ''Entoloma'' ''sensu stricto'', as previously defined, is paraphyletic (an artificial grouping). The other genera (''Leptonia'', ''Nolanea'', etc., as previously defined) are equally artificial.<ref name=CoDavid2009/>

Accordingly, ''Entoloma'' is now broadly applied by most mycologists, pending further research. Some of the component genera are, however, currently being redefined by DNA sequencing. Thus ''Nolanea'', for example, has been redefined (by excluding some species and adding others) as a monophyletic grouping within ''Entoloma'' sensu lato and treated either as a subgenus<ref name=Reschke2022/> or as a separate genus.<ref name="Karstedt2020"/> A basal group of species has also been moved to the genus ''Entocybe'' based on DNA research.<ref name="Baroni 2011"/>

==Etymology== The name ''Entoloma'' is derived from the Greek ''entos'' (ἐντός) meaning ''inner'' and ''lóma'' (λῶμα) meaning ''fringe'' from the in-rolled margin.<ref>{{cite book | vauthors = Nilson S, Persson O | year = 1977 | title = Fungi of Northern Europe 2: Gill-Fungi | publisher = Penguin|isbn=0-14-063006-6|page=98}}</ref>

==Description== [[File:Entoloma chloropolium 699336.jpg|thumb|Angular (polyhedral) basidiospores of ''Entoloma chloropolium'']] Basidiocarps (fruit bodies) are typically agaricoid (mushroom-shaped with gills), occasionally secotioid or gasteroid (truffle-like). Agaricoid species are variously large and thick-set to small and delicate, but all have lamellae (gills) that are attached to the stem (not free) and become pinkish with age from the pink basidiospores. The stipe (stem) lacks an annulus (ring). A few species are pleurotoid with a small lateral stem. Secotioid and gasteroid species (previously referred to the genera ''Richoniella'' and ''Rhodogaster'') have irregularly globose fruit bodies, brownish with a distinct stipe in secotioid species (such as the European ''Entoloma calongei'') or whitish without a stipe in truffle-like species (such as ''Entoloma gasteromycetoides'' described from New Zealand). All are internally pinkish (from the spores) when mature. Microscopically, all ''Entoloma'' species have basidiospores that are angular in all views.

==Ecology== Most species are saprotrophic, growing on decaying plant material or (less commonly) on dead wood. A few species are ectomycorrhizal. ''Entoloma sinuatum'', for example, has been shown to form an association with willows (''Salix'' species)<ref name="Agerer1997"/> and ''Entoloma nitidum'' with hornbeams (''Carpinus'' species).<ref name="Montecchio2006"/> A similar association of ''Entoloma sepium'' with fruit trees (Rosaceae) has, however, been shown to be root parasitism,<ref name="Agerer1993"/> though other studies have suggested some kind of mycorrhizal partnership may exist.<ref name=Shishikura2021/> A very few species are parasitic on other fungi, notably ''Entoloma abortivum'' a parasite of ''Armillaria'' species,<ref name="Koch2021"/> and ''Entoloma parasiticum'' which frequently grows on fruit bodies of ''Cantharellus'' species.<ref name=Noor92/>

''Entoloma'' species are found in a wide variety of habitats, including grasslands and dunes, temperate and tropical forests and woodlands, peat-bogs and moors.<ref name=Noor92/>

==Conservation== Some European ''Entoloma'' species are restricted to waxcap grassland (nutrient-poor grassland), a declining habitat as a result of changes in agricultural practice. This decline has led to four European ''Entoloma'' species, ''Entoloma bloxamii'', ''E. griseocyaneum'', ''E. porphyrophaeum'', and ''E. prunuloides'', being assessed as globally "vulnerable" on the IUCN Red List of Threatened Species.<ref name=IUCN/>

Elsewhere, several rare and localized endemic species are assessed as globally "endangered" on the IUCN Red List of Threatened Species.<ref name=IUCN/> They include ''Entoloma chilense'' in Chile, ''E. eugenei'' in Japan, Korea, and the Russian Far East, and ''E. ravinense'' in Australia. ''Entoloma alissae'' in California and ''E. necopinatum'' in Chile are assessed as globally "vulnerable".<ref name=IUCN/>

==Toxic and edible species== Several ''Entoloma'' species are known to be poisonous, causing gastroenteric symptoms (vomiting, diarrhea, nausea, and abdominal pain).<ref name="Benjamin"/> At least one poisonous species, ''E. rhodopolium'', has been found to contain significant quantities of the mycotoxin muscarine.<ref name="Benjamin"/> The English naturalist Worthington George Smith mistakenly ate ''Entoloma sinuatum'' and was "so continually and fearfully purged, and suffered so much from headache and swimming of the brain, that I really thought that every moment would be my last."<ref name="WGSmith"/> Other species known to be poisonous include ''Entoloma mammosum'', ''E. pascuum'', ''E. strictius'', and ''E. vernum''.<ref name="Benjamin"/> Additional species reported as poisonous include ''Entoloma abortivum''<ref name="Beug2006"/> (reported as edible, below), ''E. aprile'',<ref name="Beug2006"/> ''E. bahusiense'',<ref name="Beug2006"/> ''E. grande'',<ref name="Beug2006"/> ''E. luridum'',<ref name="Beug2006"/> ''E. omiense'',<ref name="Li2020"/> and ''E. quadratum''.<ref name="Li2020"/>

Fruit bodies of a number of ''Entoloma'' species are reported as being locally consumed, including ''Entoloma abortivum'' (reported as toxic, above) and ''E. clypeatum'' in Mexico, ''E. rhodopolium'' (reported as toxic, above) and ''E. clypeatum'' in Ukraine, and ''E. argyropus'' in Tanzania.<ref name="Boa 2004"/> It seems probable that some of these ''Entoloma'' species were misidentified and "all should be regarded as potentially dangerous".<ref name="Spoerke1994"/>

<gallery widths="180px" heights="180px" perrow="6"> File:2012-08-12 Entoloma murrayi (Berk. & M.A. Curtis) Sacc 250134.jpg|{{center|''Entoloma murrayi'', USA}} File:Entoloma eugenei 16579164.jpg|{{center|''Entoloma eugenei'', Russia}} File:Entoloma gasteromycetoides Co-David & Noordel 63171.jpg|{{center|''Entoloma gasteromycetoides'', New Zealand}} File:Entoloma parasiticum (Quél.) Kreisel 647283.jpg|{{center|''Entoloma parasiticum'' on ''Cantharellus'' fruit body, USA}} File:Entoloma_occidentale_var._metallicum.jpg|{{center|''Entoloma occidentale var. metallicum'', USA}} File:Entoloma necopinatum Horak 868359.jpg|{{center|''Entoloma necopinatum'', Chile}} File:2013-10-13 Entoloma abortivum (Berk. and M.A. Curtis) Donk 389687.jpg|{{center|''Entoloma abortivum'', Canada}} </gallery>

==References== {{Reflist|colwidth=30em|refs=

<ref name="Agerer1993">{{cite journal |vauthors=Agerer R, Waller K |title= Mycorrhizae of ''Entoloma saepium'': parasitism or symbiosis? |journal=Mycorrhiza |volume=3 |pages=145–154 |year=1993|issue= 4 |doi=10.1007/BF00203608|bibcode= 1993Mycor...3..145A |s2cid= 38744989 }}</ref>

<ref name="Agerer1997">{{cite journal |vauthors=Agerer R |year=1997 |title=''Entoloma sinuatum'' (Bull.: Fr.) Kummer + ''Salix'' spec. |journal=Descriptions of Ectomycorrhizae |volume=2 |pages=13–18}}</ref>

<ref name="WGSmith">{{cite book |author=Smith WG |year=1879 |title=Mushrooms and Toadstools: How to distinguish easily the differences between the Edible and Poisonous Fungi|location =London |publisher=Hardwicke and Bogue}}</ref>

<ref name="Baroni 2011">{{cite journal |vauthors=Baroni TJ, Hofstetter V, Largent DL, Vilgalys R |title=''Entocybe'' is proposed as a new genus in the Entolomataceae (Agaricomycetes, Basidiomycota) based on morphological and molecular evidence |journal=North American Fungi |year=2011 |volume=6 |issue=12 |pages=1–19 |doi=10.2509/naf2011.006.012 |doi-broken-date=12 July 2025 |url=http://www2.cortland.edu/dotAsset/49c21026-da91-4a58-8b8a-c147d8f3619d.pdf |doi-access=free }}</ref>

<ref name="Benjamin">{{cite book |author=Benjamin, Denis R. |title=Mushrooms: Poisons and Panaceas—A Handbook for Naturalists, Mycologists and Physicians |publisher=WH Freeman and Company |location=New York |year=1995 |isbn=0-7167-2600-9}}</ref>

<ref name="Beug2006">{{cite journal |vauthors=Beug MW, Shaw M, Cochran KW |title=Thirty-Plus Years of Mushroom Poisoning: Summary of the Approximately 2,000 Reports in the NAMA Case Registry |journal=McIlvainea |volume=16 |issue=2 |pages=47–68 |year=2006 |url=https://namyco.org/docs/Poisonings30year.pdf |access-date=2023-04-25 |archive-date=2023-05-30 |archive-url=https://web.archive.org/web/20230530011452/https://namyco.org/docs/Poisonings30year.pdf |url-status=dead }}</ref>

<ref name="Boa 2004">{{cite book |author=Boa ER. |title=Wild Edible Fungi: A Global Overview of Their Use and Importance to People |url=https://books.google.com/books?id=Zd2NlcNZgvcC&pg=PA138 |year=2004 |publisher=Food & Agriculture Organization |isbn=978-92-5-105157-3 |page=138}}</ref>

<ref name="CatofLife">{{cite web |title=Catalogue of Life |url=https://www.catalogueoflife.org/data/taxon/4C62 |access-date=2025-10-19}}</ref>

<ref name=CoDavid2009>{{cite journal |vauthors=Co-David D, Langeveld D, Noordeloos ME |author3-link=Machiel Noordeloos |title=Molecular phylogeny and spore evolution of Entolomataceae |journal=Persoonia |volume=23 |pages=147–76 |year=2009 |pmid=20198166 |pmc=2802732 |doi=10.3767/003158509X480944 |url=https://docserver.ingentaconnect.com/deliver/connect/nhn/00315850/v23n1/s11.pdf?expires=1685225126&id=0000&titleid=75002420&checksum=C7FFC87FF4A1FF609415D9C8CE625BB3&host=https://www.ingentaconnect.com }}</ref>

<ref name = "Donk62">{{cite journal| last=Donk| first=M.A.| title=The generic names proposed for Agaricaceae| journal=Beihefte zur Nova Hedwigia| pages=1–320| volume=5| year=1962| issn= 0078-2238}}</ref>

<ref name="Fries 1838">{{cite book |last=Fries |first=EM |year=1838 |url=https://bibdigital.rjb.csic.es/viewer/12103/?offset=#page=157&viewer=picture&o=bookmark&n=0&q= |title=Epicrisis Systematis Mycologici: Seu Synopsis Hymenomycetum |publisher=Regiae Academiae Typographia |location=Uppsala, Sweden |volume=1–2}}</ref>

<ref name=IUCN>{{cite web |url=https://www.iucnredlist.org/search?query=entoloma&searchType=species |title=''Entoloma'': The IUCN Red List of Threatened Species |access-date=2022-05-01}}</ref>

<ref name="Karstedt2020">{{cite journal |vauthors=Karstedt F, Bergemann SE, Capelari M |year=2020 |title=Five ''Nolanea'' spp. nov. from Brazil |journal=Mycotaxon |volume=135 |issue=3 |pages=589–612 |doi=10.5248/135.589 |s2cid=226328843 |url=https://docserver.ingentaconnect.com/deliver/connect/mtax/00934666/v135n3/s11.pdf?expires=1685224785&id=0000&titleid=41000038&checksum=0C66A115F25EA0FA6955768B1CD3E784&host=https://www.ingentaconnect.com}}</ref>

<ref name="Koch2021">{{cite journal |vauthors=Koch RA, Herr JR |title=Transcriptomics Reveals the Putative Mycoparasitic Strategy of the Mushroom ''Entoloma abortivum'' on Species of the Mushroom Genus ''Armillaria'' |journal=mSystems |date=2021 |volume=6 |issue=5 |pages=e0054421 |doi=10.1128/mSystems.00544-21 |doi-access=free|pmid=34636668 |pmc=8510539 }}</ref>

<ref name=Kummer>{{Cite book |last=Kummer |first=Paul |url=https://www.biodiversitylibrary.org/item/106672#page/102/mode/1upp |title=Der Führer in die Pilzkunde : Anleitung zum methodischen, leichten und sichern Bestimmen der in Deutschland vorkommenden Pilze : mit Ausnahme der Schimmel- und allzu winzigen Schleim- und Kern-Pilzchen |date=1871 |publisher=Verlag von E. Luppe's Buchhandlung |location=Zerbst |pages=94–97}}</ref>

<ref name="Largent1994">{{cite book |author=Largent DL |title=Entolomatoid fungi of the western United States and Alaska |publisher=Mad River Press |location=USA |year=1994}}</ref>

<ref name="Li2020">{{cite journal |vauthors=Li H, Zhang H, Zhang Y, Zhang K, Zhou J, Yin Y, Jiang S, Ma P, He Q, Zhang Y, Wen K, Yuan Y, Lang N, Lu J, Sun C |title=Mushroom Poisoning Outbreaks — China, 2019[J] |journal=China CDC Weekly |year=2020 |volume=2 |issue=2) |pages=19–24 |doi=10.46234/ccdcw2020.005 |pmid=34594654 |pmc=8392910 |url=https://weekly.chinacdc.cn/fileCCDCW/journal/article/ccdcw/2020/2/PDF/190051.pdf}}</ref>

<ref name="Montecchio2006">{{cite journal |vauthors=Montecchio L, Rossi S, Courty P, Garbaye J |title=Descriptions of Ectomycorrhizae. ''Entoloma nitidum'' Quel. plus ''Carpinus betulus'' L. |year=2006 |journal=Descriptions of Ectomycorrhizae |volume=9/10 |pages=31–36 |url=https://hal.inrae.fr/hal-02820776/document}}</ref>

<ref name=Noor92>{{cite book |title=Fungi Europaei:''Entoloma'' sensu lato|author=Noordeloos ME |year=1992 |publisher=Giovanna Biella |location=Saronno, Italy |pages=760 }}</ref>

<ref name="Orton1991">{{cite journal |vauthors=Orton PD |title=A revised list of British species of ''Entoloma'' sensu lato |journal=The Mycologist |year=1991 |volume=5 |issue=3 |pages=123–138 |doi=10.1016/S0269-915X(09)80307-8}}</ref>

<ref name="Orton1991b">{{cite journal |vauthors=Orton PD |title=A revised list of British species of ''Entoloma'' sensu lato (part 2) |journal=The Mycologist |year=1991 |volume=5 |issue=4 |pages=172–176 |doi=10.1016/S0269-915X(09)80478-3}}</ref>

<ref name="Quélet 1886">{{cite book |author=Quélet L. |title=Enchiridion fungorum in Europa media et praesertim in Gallia vigentium |publisher=O. Doin |location=Switzerland |year=1886 |page=155 |url=https://www.biodiversitylibrary.org/item/101574#page/65/mode/1up}}</ref>

<ref name=Reschke2022>{{cite journal |vauthors=Reschke K, Morozova OV, Dima B, Cooper JA, Corriol G, Biketova AY, Piepenbring M, Noordeloos ME |year=2022 |title=Phylogeny, taxonomy, and character evolution in ''Entoloma'' subgenus ''Nolanea'' |journal=Persoonia |volume=49|pages=136–170 |doi=10.3767/persoonia.2022.49.04 |pmid=38234382 |pmc=10792224 |url=https://docserver.ingentaconnect.com/deliver/connect/nhn/00315850/v49n1/s4.pdf?expires=1685225026&id=0000&titleid=75002420&checksum=3CB2C27E2388459C507924467B7F515B&host=https://www.ingentaconnect.com }}</ref>

<ref name=Shishikura2021>{{cite journal |vauthors=Shishikura M, Takemura Y, Sotome K, Maekawa N, Nakagiri A, Endo N |year=2021 |title=Four mycelial strains of ''Entoloma clypeatum'' species complex form ectomycorrhiza-like roots with ''Pyrus betulifolia'' seedlings in vitro, and one develops fruiting bodies 2 months after inoculation |journal=Mycorrhiza |volume=31 |issue=1 |pages=31–42 |doi=10.1007/s00572-020-00994-4|pmid=33105488 |bibcode=2021Mycor..31...31S |s2cid=254047880 }}</ref>

<ref name="Spoerke1994">{{cite book |author=Spoerke DG, Rumack BH |year=1994 |title=Handbook of mushroom poisoning: diagnosis and treatment |publisher=CRC Press |location=Boca Raton}}</ref> }}

==External links== * [http://www.mushroomexpert.com/entoloma.html Mushroom Expert - The genus ''Entoloma''] * [http://botit.botany.wisc.edu/toms_fungi/sep2006.html Tom Volk's Fungus of the Month - ''Entoloma abortivum'']

Category:Entolomataceae Category:Agaricales genera Category:Taxa named by Elias Magnus Fries Category:Taxa described in 1838 Category:Entoloma