Collybia Cirrhata
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''Collybia cirrhata'' is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the family
Tricholomataceae The Tricholomataceae are a large family of fungi within the order Agaricales. Originally a classic "wastebasket taxon", the family included any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to e.g. t ...
of the order
Agaricales The Agaricales are an order (biology), order of fungi in the division (mycology), division Basidiomycota. As originally conceived, the order contained all the agarics (gilled mushrooms), but subsequent research has shown that not all agarics are ...
(gilled
mushroom A mushroom or toadstool is the fleshy, spore-bearing Sporocarp (fungi), fruiting body of a fungus, typically produced above ground on soil or another food source. ''Toadstool'' generally refers to a poisonous mushroom. The standard for the n ...
s). The species was first described in the scientific literature in 1786, but was not validly named until 1803. Found in Europe, Northern Eurasia, and North America, it is known from
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
,
boreal Boreal, northern, of the north. Derived from the name of the god of the north wind from Ancient Greek civilisation, Boreas (god), Boreas. It may also refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernand ...
, and
alpine Alpine may refer to any mountainous region. It may also refer to: Places Europe * Alps, a European mountain range ** Alpine states, which overlap with the European range Australia * Alpine, New South Wales, a Northern Village * Alpine National P ...
or
arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
habitats. It is a
saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
species that grows in clusters on the decaying or blackened remains of other mushrooms. 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 ...
are small, with whitish convex to flattened
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 ...
up to in diameter, narrow white
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 ...
, and slender whitish stems long and up to thick. ''C. cirrhata'' can be distinguished from the other two members of ''
Collybia ''Collybia'' (in the strict sense) is a genus of mushrooms in the family Clitocybaceae. The genus has a widespread distribution in northern temperate areas, and contains well known species like the Collybia nuda, blewit, Collybia sordida, s ...
'' by the absence of a
sclerotium A sclerotium (; : sclerotia () is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant u ...
at the base of the stem. A 2023 chemical analysis found that this mushroom contains 17 to 56 mg/kg of the mycotoxin muscarine.


Taxonomy and phylogeny

The species first appeared in the scientific literature in 1786 as ''Agaricus amanitae'' by
August Johann Georg Karl 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 i ...
; ''Agaricus amanitae'' subsp. ''cirrhatus'', proposed by
Christian Hendrik Persoon Christiaan Hendrik Persoon (31 December 1761 – 16 November 1836) was a Cape Colony mycologist who is recognized as one of the founders of mycology, mycological Taxonomy (biology), taxonomy. Early life Persoon was born in Cape Colony at ...
in 1800, is considered
synonymous A synonym is a word, morpheme, or phrase that means precisely 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 a ...
. A later combination based on this name, ''Collybia amanitae'', was published by
Hanns Kreisel Hanns Kreisel (16 July 1931 – 18 January 2017) was a German mycologist and professor emeritus. He was born in Leipzig in 1931. Kreisel was a professor at the University of Greifswald. His field was the classification of fungi, where he has s ...
in 1987. However, Kreisel noted the combination to be "''ined.''", indicating that he did not believe the name to be validly published, according to article 34.1 of the rules for botanical nomenclature, which states: "A name is not validly published ... when it is not accepted by the author in the original publication." The first valid name was published in 1803 by
Heinrich Christian Friedrich Schumacher Heinrich Christian Friedrich Schumacher (15 November 1757 in Glückstadt, Holstein – 9 December 1830) was a Denmark, Danish surgeon, botanist and professor of anatomy at the University of Copenhagen. Schumacher carried out significant research ...
, who called the species ''Agaricus cirrhatus''. French mycologist
Lucien Quélet Lucien Quélet (; 14 July 1832 – 25 August 1899) was a French natural history, naturalist and mycologist. Quélet discovered several species of fungi and was the founder of the Société mycologique de France, a society devoted to mycological ...
transferred it to ''Collybia'' in 1879, resulting in the
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms *Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition * ...
by which it is currently known. The species had also been transferred to ''Microcollybia'' by
Georges Métrod Georges may refer to: Places *Georges River, New South Wales, Australia *Georges Quay (Dublin) *Georges Township, Fayette County, Pennsylvania Other uses *Georges (name) * ''Georges'' (novel), a novel by Alexandre Dumas * "Georges" (song), a 1977 ...
in 1952 and again by Lennox in 1979 (because Métrod's transfer was considered a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'', and thus invalid according to nomenclatural rules); the genus ''Microcollybia'' has since been wrapped into ''Collybia''.
Molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
have shown that ''C. cirrhata'' forms a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
with the remaining two species of ''Collybia''. Because ''C. cirrhata'' is the only one of the three ''Collybia'' species lacking
sclerotia A sclerotium (; : sclerotia () is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant u ...
, it has been suggested that this character trait is an '' anapomorphy''—that is, unique to a single, terminal species within a clade. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, meaning "curled".
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 fu ...
called it the "fringed-rooted Collybia". In the United Kingdom, it is commonly known as the "piggyback shanklet".


Description

The
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. The origin of the word "cap" comes from the Old French word "chapeau" which means "head co ...
is initially convex when young, later becoming convex to flattened or slightly depressed in the center, reaching a diameter of . The cap margin starts out rolled or curved inward, but straightens out as it matures. The cap surface ranges from dry to moist, smooth to covered with fine whitish hairs, and is mostly even with translucent radial grooves at the margin. It is subhygrophanous (changing color somewhat depending on hydration), becoming a grayish-orange when watery or old, and usually is white with a very faint pinkish flush when fresh. The
flesh Flesh is any aggregation of soft tissues of an organism. Various multicellular organisms have soft tissues that may be called "flesh". In mammals, including humans, ''flesh'' encompasses muscles, fats and other loose connective tissues, ...
is whitish, quite thin, and has no distinctive taste or odor. 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
adnate Adnate may refer to: * Adnation, in botany, the fusion of two or more whorls of a flower * Adnate, in mycology, a classification of lamellae (gills) * Conjoined twins Conjoined twins, popularly referred to as Siamese twins, are twins joined '' ...
to slightly arcuate (curved into the shape of a bow) with a tooth (meaning that the gills curve up to join the stem but then, very close to the stem, the gill edge curves down again). There are between 12 and 20 gills that extend completely from the cap edge to the stem, and three to five tiers of lamellulae (shorter gills that do not extend completely from the cap edge to the stem). The gills are thin, narrow to moderately broad, and white to pinkish-
buff Buff or BUFF may refer to: People * Buff (surname), a list of people * Buff (nickname), a list of people * Johnny Buff, ring name of American world champion boxer John Lisky (1888–1955) * Buff Bagwell, a ring name of American professional wr ...
. The gill edges are even, and the same color as the gill faces. The
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
is long and up to thick, equal in width throughout to slightly enlarged downward, flexible and filamentous but not fragile. The stem surface is dry, whitish to grayish-orange, sometimes with tiny hairs on the upper portion that become coarser near the base. The stem base often has
rhizomorph Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae. Cords may look similar to plant roots, and also frequently have similar functi ...
-like strands or copious whitish
mycelia Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...
. The stem, unlike the other two species of ''Collybia'', do not originate from a
sclerotium A sclerotium (; : sclerotia () is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant u ...
. The stem becomes hollow as it matures. ''C. cirrhata'' is too small and insubstantial to be considered
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 ...
.


Microscopic characteristics

In deposit, the spores appear white. Individual
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s are ellipsoid to tear-shaped in profile, obovoid to ellipsoid or roughly cylindric in face or back view, with dimensions of 4.8–6.4 by 2–2.8 (sometimes up to 3.5)
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
. They are smooth,
inamyloid In mycology a tissue or feature is said to be amyloid if it has a positive amyloid reaction when subjected to a crude chemical test using iodine as an ingredient of either Melzer's reagent or Lugol's solution, producing a blue to blue-black staini ...
, and acyanophilous (unreactive to
staining Staining is a technique used to enhance contrast in samples, generally at the Microscope, microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissue (biology), tissues), in cytology (microscopic ...
with
Melzer's reagent Melzer's reagent (also known as Melzer's iodine reagent, Melzer's solution or informally as Melzer's) is a chemical reagent used by mycologists to assist with the identification of fungi, and by phytopathologists for fungi that are plant pathogens ...
and
methyl blue Methyl blue is a chemical compound with the molecular formula C37H27N3Na2O9S3. It is used as a stain in histology, and stains collagen blue in tissue sections. It can be used in some differential staining techniques such as Mallory's trichrome ...
, respectively). The
basidia A basidium (: basidia) is a microscopic spore-producing structure found on the hymenophore of reproductive bodies of basidiomycete fungi. The presence of basidia is one of the main characteristic features of the group. These bodies are also ...
(spore-bearing cells of the
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
) are roughly club-shaped, four-spored, and measure 17.5–21 by 4.8–5.6 μm. The gills do not have
cystidia A cystidium (: cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that are o ...
. The gill tissue is made of
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e that are interwoven to roughly parallel, and inamyloid. The hyphae are 2.8–8.4 μm in diameter and smooth. The cap tissue is made of interwoven hyphae beneath the center of the cap, but radially oriented over the gills; it too is inamyloid. These hyphae are 3.5–8.4 μm in diameter, smooth, but have irregularly thickened walls. The
cap cuticle The pileipellis is the uppermost layer of hyphae in the pileus of a fungal fruit body. It covers the trama, the fleshy tissue of the fruit body. The pileipellis is more or less synonymous with the cuticle, but the cuticle generally describes th ...
is an ixocutis—a gelatinized layer of hyphae lying parallel to the cap surface. The hyphae comprising this layer are 2.8–6.4 μm in diameter, smooth, and thin-walled. They are covered with scattered, short pouch-like outgrowths. The cuticle of the stem is a layer of parallel, vertically oriented hyphae; the hyphae measure 3.5–4.2 μm, and are smooth, slightly thick-walled, and pale yellowish-brown in
alkali In chemistry, an alkali (; from the Arabic word , ) is a basic salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as a base that dissolves in water. A solution of a soluble base has a pH greater than 7.0. The a ...
ne solution. They give rise to a covering of tangled and branched caulocystidia (cystidia on the stem) that have multiple
septa SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...
. The caulocystidia are 2.8–4.8 μm in diameter, smooth, thin walled, and shaped like contorted cylinders.
Clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of basidiomycete fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rece ...
s are present in the hyphae of all tissues.


Similar species

''Collybia cirrhata'' is most likely to be confused with the remaining members of ''
Collybia ''Collybia'' (in the strict sense) is a genus of mushrooms in the family Clitocybaceae. The genus has a widespread distribution in northern temperate areas, and contains well known species like the Collybia nuda, blewit, Collybia sordida, s ...
'', which have a similar external appearance. '' C. tuberosa'' is distinguished by its dark reddish-brown
sclerotia A sclerotium (; : sclerotia () is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant u ...
that resemble apple seeds, while '' C. cookei'' has wrinkled, often irregularly shaped sclerotia that are pale yellow to orange. Other similar mushrooms include '' Baeospora myosura'' and species of '' Strobilurus'', but these species only grow on
pine cone A conifer cone, or in formal botanical usage a strobilus, : strobili, is a seed-bearing organ on gymnosperm plants, especially in conifers and cycads. They are usually woody and variously conic, cylindrical, ovoid, to globular, and have scal ...
s.


Habitat and distribution

Like all species remaining in the genus ''Collybia'', ''C. cirrhata'' is
saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
, and is typically found growing on the decaying or blackened remains of other mushrooms; occasionally the fruit bodies may be found growing on moss or soil without any apparent connection to decaying mushrooms, although these observations may represent instances where the remnant host tissue—possibly from a previous season—has decayed to such an extent that it remains as buried fragments in the substrate. Known hosts include ''
Lactarius ''Lactarius'' is a genus of mushroom-producing, ectomycorrhizal fungi, containing several Edible mushroom, edible species. The species of the genus, common name, commonly known as milk-caps, are characterized by the milky fluid ("latex") they ex ...
'', ''
Russula ''Russula'' is a very large genus composed of around 750 worldwide species of fungi. The genus was described by Christian Hendrik Persoon in 1796. The mushrooms are fairly large, and brightly colored – making them one of the most recognizable ...
'', ''
Meripilus giganteus ''Meripilus giganteus'' is a polypore fungus in the family Meripilaceae. It causes a white rot in various types of broadleaved trees, particularly beech (''Fagus''), but also ''Abies'', ''Picea'', ''Pinus'', ''Quercus'' and ''Ulmus'' species. T ...
'', and ''
Bovista dermoxantha ''Bovista dermoxantha'' is a small, white, nearly round puffball, recognized when young by a cottony-felty outer surface that becomes inconspicuously warted, eventually leaving fine, pallid, scales on an ochre to brown endoperidium. ''Bovista p ...
''. ''Collybia cirrhata'' is known from
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
,
boreal Boreal, northern, of the north. Derived from the name of the god of the north wind from Ancient Greek civilisation, Boreas (god), Boreas. It may also refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernand ...
, and
alpine Alpine may refer to any mountainous region. It may also refer to: Places Europe * Alps, a European mountain range ** Alpine states, which overlap with the European range Australia * Alpine, New South Wales, a Northern Village * Alpine National P ...
or
arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
habitats. The fungus is widespread in Europe, including Bulgaria, Denmark, Germany, Greece, Latvia,
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
, Slovakia, Switzerland, Turkey, and the United Kingdom. The mushroom is also common in northern
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
regions of North America. In Asia, the fungus has been reported in Korea, and in
Hokkaido is the list of islands of Japan by area, second-largest island of Japan and comprises the largest and northernmost prefectures of Japan, prefecture, making up its own list of regions of Japan, region. The Tsugaru Strait separates Hokkaidō fr ...
, northern Japan. It is also known from
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
. A 2009 publication suggested that based on the known evidence, the species' distribution may be
circumboreal The Circumboreal Region in phytogeography is a floristic region within the Holarctic Kingdom in Eurasia and North America, as delineated by such geobotanists as Josias Braun-Blanquet and Armen Takhtajan. It is the largest floristic region in ...
.


References


External links

* {{good article Fungi of Asia Fungi of Europe Fungi of North America Inedible fungi Tricholomataceae Fungi described in 1803 Fungus species