Collybia cirrhata
Encyclopedia
Collybia cirrhata is a species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of fungus
Fungus
A fungus is a member of a large 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, Fungi, which is separate from plants, animals, and bacteria...

 in the Tricholomataceae
Tricholomataceae
The Tricholomataceae are a large family of mushrooms within the Agaricales. A classic "wastebasket taxon", the Tricholomataceae is inclusive of any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to the Amanitaceae, Lepiotaceae, Hygrophoraceae,...

 family of the Agaricales
Agaricales
The fungal order Agaricales, also known as gilled mushrooms , or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species, along with five extinct genera known only from the fossil record...

 order (gilled mushroom
Mushroom
A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or on its food source. The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus; hence the word "mushroom" is most often applied to those fungi that...

s). The species was first described
Species description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...

 in the scientific literature in 1786, but not validly named until 1803. Found in Europe, Northern Eurasia, and North America, it is known from temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

, boreal
Subarctic climate
The subarctic climate is a climate characterized by long, usually very cold winters, and short, cool to mild summers. It is found on large landmasses, away from the moderating effects of an ocean, generally at latitudes from 50° to 70°N poleward of the humid continental climates...

, and alpine
Alpine climate
Alpine climate is the average weather for a region above the tree line. This climate is also referred to as mountain climate or highland climate....

 or arctic
Climate of the Arctic
The climate of the Arctic is characterized by long, cold winters and short, cool summers. There is a large amount of variability in climate across the Arctic, but all regions experience extremes of solar radiation in both summer and winter...

 habitats. It is a saprobic species that grows in clusters on the decaying or blackened remains of other mushrooms. The fruit bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

 are small, with whitish convex to flattened caps
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...

 up to 11 mm (0.433070866141732 in) in diameter, narrow white gills, and slender whitish stems
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...

 8 – long and up to 2 mm (0.078740157480315 in) thick. C. cirrhata can be distinguished from the other two members of Collybia
Collybia
Collybia is a genus of mushrooms in the Tricholomataceae family. The genus has a widespread but rare distribution in north temperate areas, and contains three species that grow on the decomposing remains of other mushrooms....

by the absence of a sclerotium
Sclerotium
A sclerotium 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 until a favorable opportunity for growth. Other fungi that produce...

 at the base of the stem. The mushroom, although not poisonous, is considered inedible because of its insubstantial size.

Taxonomy and phylogeny

The species first appeared in the scientific literature in 1786 as Agaricus amanitae by August Johann Georg Karl Batsch; Agaricus amanitae subsp. cirrhatus, proposed by Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 in 1800, is considered synonymous. A later combination based on this name, Collybia amanitae, was published by Hanns Kreisel
Hanns Kreisel
Hanns Kreisel is a German mycologist and professor emeritus.Kreisel was a professor at the University of Greifswald. His field is the classification of fungi, where he has studied this group of organisms not only in Germany but in almost all continents, as in Brazil, Seychelles, Vietnam, Cuba and...

 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 correct name
Correct name (botany)
In botany, the correct name according to the International Code of Botanical Nomenclature is the one and only botanical name that is to be used for a particular taxon, when that taxon has a particular circumscription, position and rank. Determining whether a name is correct is a complex procedure...

 was published in 1803 by Heinrich Christian Friedrich Schumacher
Heinrich Christian Friedrich Schumacher
Heinrich Christian Friedrich Schumacher , was a Danish surgeon, botanist and professor of anatomy at the University of Copenhagen...

, who called the species Agaricus cirrhatus. French mycologist Lucien Quélet
Lucien Quélet
thumb|Lucien QuéletLucien Quélet was a French mycologist and naturalist who discovered several species and was the founder of the Société mycologique de France, a society devoted to mycological studies....

 transferred it to Collybia in 1879, resulting in the binomial by which it is currently known. The species had also been transferred to Microcollybia by Georges Métrod in 1952 and again by Lennox in 1979 (because Métrod's transfer was considered a nomen nudum
Nomen nudum
The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...

, and thus invalid according to nomenclatural rules); the genus Microcollybia has since been wrapped into Collybia.

Molecular phylogenetics have shown that C. cirrhata forms a monophyletic clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 with the remaining two species of Collybia. Because C. cirrhata is the only one of the three Collybia species lacking sclerotia, 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 is derived from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 cirrata, meaning "curled". Charles Horton Peck
Charles Horton Peck
Charles Horton Peck, born March 30, 1833 in Sand Lake, New York, died 1917 in Albany, New York, was an American mycologist of the 19th and early 20th centuries...

 called it the "fringed-rooted Collybia". In the United Kingdom, it is commonly
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 known as the "piggyback shanklet".

Description

The cap
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...

 is initially convex when young, later becoming convex to flattened or slightly depressed in the center, reaching a diameter of 3 –. 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
Hygrophanous
The adjective hygrophanous refers to the color change of mushroom tissue as it loses or absorbs water, which causes the pileipellis to become more transparent when wet and opaque when dry....

 (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
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....

 is whitish, quite thin, and has no distinctive taste or odor. The gills are adnate
Adnation
Adnation in plants is the "union of unlike parts; organically united or fused with another dissimilar part, e.g. an ovary to a calyx tube, or stamens to petals". This is in contrast to connation, the fusion of similar organs....

 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. The gill edges are even, and the same color as the gill faces.

The stem
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...

 is 8 – long and up to 2 mm (0.078740157480315 in) 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-like strands or copious whitish mycelia. The stem, unlike the other two species of Collybia, do not originate from a sclerotium
Sclerotium
A sclerotium 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 until a favorable opportunity for growth. Other fungi that produce...

. The stem becomes hollow as it matures. Although it is not considered poisonous, C. cirrhata is too small and insubstantial to be considered edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

.

Microscopic characteristics

In deposit
Spore print
thumb|300px|right|Making a spore print of the mushroom Volvariella volvacea shown in composite: mushroom cap laid on white and dark paper; cap removed after 24 hours showing pinkish-tan spore print...

, the spores appear white. Individual spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

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
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...

. They are smooth, inamyloid
Amyloid (mycology)
In mycology the term amyloid refers to a crude chemical test using iodine in either Melzer's reagent or Lugol's solution, to produce a black to blue-black positive reaction. It is called amyloid because starch gives a similar reaction, and that reaction for starch is also called an amyloid reaction...

, and acyanophilous (unreactive to staining with Melzer's reagent
Melzer's Reagent
Melzer's reagent is a chemical reagent used by mycologists to assist with the identification of fungi.-Composition:...

 and methyl blue
Methyl blue
Methyl blue, also known as Cotton blue, Helvetia blue, Acid blue 93, or C.I. 42780, is a chemical compound with the molecular formula C37H27N3Na2O9S3. It is used as a stain in histology. Methyl blue stains collagen blue in tissue sections. It is soluble in water and slightly soluble in ethanol...

, respectively). The basidia (spore-bearing cells of the hymenium
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 others some cells develop into sterile cells called cystidia or...

) are roughly club-shaped, four-spored, and measure 17.5–21 by 4.8–5.6 µm. The gills do not have cystidia. The gill tissue is made of hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

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
Pileipellis
thumb|300px||right|The cuticle of some mushrooms, such as [[Russula mustelina]] shown here, can be peeled from the cap, and may be useful as an identification feature....

 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
Alkali
In chemistry, an alkali is a basic, ionic salt of an alkali metal or alkaline earth metal element. Some authors also define an alkali as a base that dissolves in water. A solution of a soluble base has a pH greater than 7. The adjective alkaline is commonly used in English as a synonym for base,...

ne solution. They give rise to a covering of tangled and branched caulocystidia (cystidia on the stem) that have multiple septa
Septum
In anatomy, a septum is a wall, dividing a cavity or structure into smaller ones.-In human anatomy:...

. The caulocystidia are 2.8–4.8 µm in diameter, smooth, thin walled, and shaped like contorted cylinders. Clamp connection
Clamp connection
A clamp connection is a structure formed by growing hyphal cells of certain fungi. It is created to ensure each septum, or segment of hypha separated by crossed walls, receives a set of differing nuclei, which are obtained through mating of hyphae of differing sexual types...

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
Collybia is a genus of mushrooms in the Tricholomataceae family. The genus has a widespread but rare distribution in north temperate areas, and contains three species that grow on the decomposing remains of other mushrooms....

, which have a similar external appearance. C. tuberosa
Collybia tuberosa
Collybia tuberosa, commonly known as the lentil shanklet or the appleseed coincap, is an inedible species of fungus in the Tricholomataceae family, and the type species of the genus Collybia. Like the two other members of its genus, it lives on the decomposing remains of other fleshy mushrooms...

is distinguished by its dark reddish-brown sclerotia that resemble apple seeds, while C. cookei
Collybia cookei
Collybia cookei is a species of fungus in the Tricholomataceae family, and one of three species in the genus Collybia. It is known from Europe, Asia, and North America...

has wrinkled, often irregularly shaped sclerotia that are pale yellow to orange. Other similar mushrooms include Baeospora myosura
Baeospora myosura
Baeospora myosura is a species of fungus that produces mushrooms with long, coarse hairs. It grows on plant material and manure. It is white to cream and the spore color is white, cream, or yellowish. It is commonly found in North America and Europe. The common name of the mushroom is conifercone...

and species of Strobilurus
Strobilurus
Strobilurus is a genus of fungi in the Physalacriaceae family. The genus has a widespread distribution in temperate regions, and contains 10 species. Species of Strobilurus grow on pine cones.-Species:*S. albipilatus*S. conigenoides...

, but these species only grow on pine cones.

Habitat and distribution

Like all species remaining in the genus Collybia, C. cirrhata is saprobic, 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
Lactarius is a genus of mushroom-producing fungi. The genus, collectively known commonly as milk-caps, are characterized by the fact that they exude a milky fluid if cut or damaged...

, Russula
Russula
Around 750 worldwide species of mycorrhizal mushrooms compose the genus Russula. They are typically common, fairly large, and brightly colored - making them one of the most recognizable genera among mycologists and mushroom collectors...

, Meripilus giganteus
Meripilus giganteus
Meripilus giganteus is a polypore fungus in the Meripilaceae family. It causes a white rot in various types of broadleaved trees, particularly beech , but also Abies, Picea, Pinus, Quercus and Ulmus species...

, and Bovista dermoxantha
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 plumbea is similar, but has a smoother surface when young, and...

.

Collybia cirrhata is known from temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

, boreal
Subarctic climate
The subarctic climate is a climate characterized by long, usually very cold winters, and short, cool to mild summers. It is found on large landmasses, away from the moderating effects of an ocean, generally at latitudes from 50° to 70°N poleward of the humid continental climates...

, and alpine
Alpine climate
Alpine climate is the average weather for a region above the tree line. This climate is also referred to as mountain climate or highland climate....

 or arctic
Climate of the Arctic
The climate of the Arctic is characterized by long, cold winters and short, cool summers. There is a large amount of variability in climate across the Arctic, but all regions experience extremes of solar radiation in both summer and winter...

 habitats. The fungus is widespread in Europe, including Bulgaria, Denmark, Germany, Greece, Latvia, Scandinavia
Scandinavia
Scandinavia is a cultural, historical and ethno-linguistic region in northern Europe that includes the three kingdoms of Denmark, Norway and Sweden, characterized by their common ethno-cultural heritage and language. Modern Norway and Sweden proper are situated on the Scandinavian Peninsula,...

, Slovakia, Switzerland, Turkey, and the United Kingdom. The mushroom is also common in northern montane
Montane
In biogeography, montane is the highland area located below the subalpine zone. Montane regions generally have cooler temperatures and often have higher rainfall than the adjacent lowland regions, and are frequently home to distinct communities of plants and animals.The term "montane" means "of the...

 regions of North America. In Asia, the fungus has been reported in Korea, and in Hokkaido
Hokkaido
, formerly known as Ezo, Yezo, Yeso, or Yesso, is Japan's second largest island; it is also the largest and northernmost of Japan's 47 prefectural-level subdivisions. The Tsugaru Strait separates Hokkaido from Honshu, although the two islands are connected by the underwater railway Seikan Tunnel...

, northern Japan. It is also known from Greenland
Greenland
Greenland is an autonomous country within the Kingdom of Denmark, located between the Arctic and Atlantic Oceans, east of the Canadian Arctic Archipelago. Though physiographically a part of the continent of North America, Greenland has been politically and culturally associated with Europe for...

. A 2009 publication suggested that based on the known evidence, the species' distribution may be circumboreal
Circumboreal Region
The Circumboreal Region 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....

.
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