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''Galerina marginata'', known colloquially as funeral bell, deadly skullcap, autumn skullcap or deadly galerina, is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of extremely poisonous
mushroom A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans. The standard for the name "mushroom" is t ...
-forming
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
in the family
Hymenogastraceae The Hymenogastraceae is a family of fungi in the order Agaricales with both agaric and false-truffle shaped fruitbodies. Formerly, prior to molecular analyses, the family was restricted to the false-truffle genera. The mushroom genus ''Psilocybe' ...
of the order
Agaricales The fungal order Agaricales, also known as gilled mushrooms (for their distinctive gills) or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13,000 described species, alo ...
. It contains the same deadly
amatoxin Amatoxin is the collective name of a subgroup of at least nine related toxic compounds found in three genera of poisonous mushrooms (''Amanita'', ''Galerina'' and ''Lepiota'') and one species (Conocybe filaris) of the genus ''Conocybe''. Amatoxins a ...
s found in the death cap (''
Amanita phalloides ''Amanita phalloides'' (), commonly known as the death cap, is a deadly poisonous basidiomycete fungus, one of many in the genus ''Amanita''. Widely distributed across Europe, but now sprouting in other parts of the world, ''A. phalloides ...
''). Ingestion in toxic amounts causes severe
liver damage Liver disease, or hepatic disease, is any of many diseases of the liver. If long-lasting it is termed chronic liver disease. Although the diseases differ in detail, liver diseases often have features in common. Signs and symptoms Some of the si ...
with vomiting, diarrhea,
hypothermia Hypothermia is defined as a body core temperature below in humans. Symptoms depend on the temperature. In mild hypothermia, there is shivering and mental confusion. In moderate hypothermia, shivering stops and confusion increases. In severe h ...
, and eventual death if not treated rapidly. About ten poisonings have been attributed to the species now grouped as ''G. marginata'' over the last century. ''G. marginata'' is widespread in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
, including Europe, North America, and Asia, and has also been found in Australia. It is a wood-rotting fungus that grows predominantly on decaying
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
wood. The
fruit bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
of the mushroom have brown to yellow-brown
caps Caps are flat headgear. Caps or CAPS may also refer to: Science and technology Computing * CESG Assisted Products Service, provided by the U.K. Government Communications Headquarters * Composite Application Platform Suite, by Java Caps, a Java ...
that fade in color when drying. The
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
are brownish and give a rusty
spore print 300px, Making a spore print of the mushroom ''Volvariella volvacea'' shown in composite: (photo lower half) mushroom cap laid on white and dark paper; (photo upper half) cap removed after 24 hours showing pinkish-tan spore print. A 3.5-centimeter ...
. A well-defined membranous
ring Ring may refer to: * Ring (jewellery), a round band, usually made of metal, worn as ornamental jewelry * To make a sound with a bell, and the sound made by a bell :(hence) to initiate a telephone connection Arts, entertainment and media Film and ...
is typically seen on the stems of young specimens but often disappears with age. In older fruit bodies, the caps are flatter and the gills and stems browner. The species is a classic "
little brown mushroom Mushroom hunting, mushrooming, mushroom picking, mushroom foraging, and similar terms describe the activity of gathering mushrooms in the wild, typically for culinary use. This practice is popular throughout most of Europe, Australia, Japan, ...
" – a catchall category that includes all small to medium-sized, hard-to-identify brownish mushrooms, and may be easily confused with several
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
species. Before 2001, the species ''G. autumnalis'', ''G. oregonensis'', ''G. unicolor'', and ''G. venenata'' were thought to be distinct from ''G. marginata'' due to differences in habitat and the viscidity of their caps, but
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis showed that they are all the same species.


Taxonomy and naming

What is now recognized as a single morphologically variable
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
named ''Galerina marginata'' was once split into five distinct species. Norwegian mycologist Gro Gulden and colleagues concluded that all five represented the same species after comparing the DNA sequences of the
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. I ...
region of
ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chromos ...
for various North American and European specimens in ''
Galerina ''Galerina'' is a genus of small brown-spore saprobic fungi (colloquially often ''mushrooms''), with over 300 species found throughout the world from the far north to remote Macquarie Island in the Southern Ocean. The genus is most noted for some ...
'' section ''Naucoriopsis''. The results showed no genetic differences between ''G. marginata'' and ''G. autumnalis'', ''G. oregonensis'', ''G. unicolor'', and ''G. venenata'', thus reducing all these names to
synonymy A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
. The oldest of these names are ''Agaricus marginatus'', described by
August Batsch August Johann Georg Karl Batsch (28 October 1761 – 29 September 1802) was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants. Life and career Batsch was born ...
in 1789, and ''Agaricus unicolor'', described by
Martin Vahl Martin Henrichsen Vahl (10 October 1749 – 24 December 1804) was a Danish-Norwegian botanist, herbalist and zoologist. Biography Martin Vahl was born in Bergen, Norway and attended Bergen Cathedral School. He studied botany at the University ...
in 1792. ''Agaricus autumnalis'' was described by
Charles Horton Peck Charles Horton Peck (March 30, 1833 – July 11, 1917) was an American mycologist of the 19th and early 20th centuries. He was the New York State Botanist from 1867 to 1915, a period in which he described over 2,700 species of North American fun ...
in 1873, and later moved to ''Galerina'' by A. H. Smith and
Rolf Singer Rolf Singer (June 23, 1906 – January 18, 1994) was a Germany, German-born mycologist and one of the most important Taxonomy (biology), taxonomists of gilled mushrooms (agarics) in the 20th century. After receiving his Ph.D. at the University ...
in their 1962 worldwide
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
on that genus. In the same publication they also introduced the ''G. autumnalis''
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
''robusta'' and ''angusticystis''. Another of the synonymous species, ''G. oregonensis'', was first described in that monograph. ''Galerina venenata'' was first identified as a species by Smith in 1953. Since ''Agaricus marginatus'' is the oldest validly published name, it has priority according to the rules of botanical nomenclature. Another species analysed in Gulden's 2001 study, Galerina pseudomycenopsis, also could not be distinguished from ''G. marginata'' based on ribosomal DNA sequences and restriction fragment length polymorphism analyses. Because of differences in ecology, fruit body color and spore size combined with inadequate sampling, the authors preferred to maintain ''G. pseudomycenopsis'' as a distinct species. A 2005 study again failed to separate the two species using molecular methods, but reported that the incompatibility demonstrated in
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
experiments suggests that the species are distinct. In the fourth edition (1986) of Singer's comprehensive
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of the
Agaricales The fungal order Agaricales, also known as gilled mushrooms (for their distinctive gills) or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13,000 described species, alo ...
, ''G. marginata'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of ''Galerina'' section ''Naucoriopsis'', a subdivision first defined by French mycologist
Robert Kühner Robert Kühner (15 March 1903 in Paris – 27 February 1996 in Lyon) was a French mycologist most notable for reviewing many forms of agaric (mushroom fungus) genera. He studied at the Sorbonne, afterwards from 1921 until 1932, he was working ...
in 1935. It includes small brown-spored mushrooms characterized by cap edges initially curved inwards, fruit bodies resembling ''
Pholiota ''Pholiota'' is a genus of small to medium-sized, fleshy mushrooms in the family Strophariaceae. They are saprobes that typically live on wood. The genus has a widespread distribution, especially in temperate regions, and contains about 150 spe ...
'' or '' Naucoria'' and thin-walled, obtuse or acute-ended pleurocystidia that are not rounded at the top. Within this section, ''G. autumnalis'' and ''G. oregonensis'' are in
stirps In ancient Rome, a gens ( or , ; plural: ''gentes'' ) was a family consisting of individuals who shared the same nomen and who claimed descent from a common ancestor. A branch of a gens was called a ''stirps'' (plural: ''stirpes''). The ''gen ...
''Autumnalis'', while ''G. unicolor'', ''G. marginata'', and ''G. venenata'' are in stirps ''Marginata''. ''Autumnalis'' species are characterized by having a viscid to lubricous cap surface while ''Marginata'' species lack a gelatinous cap—the surface is moist, "fatty-shining", or matte when wet. However, as Gulden explains, this characteristic is highly variable: "Viscidity is a notoriously difficult character to assess because it varies with the age of the fruitbody