Mycena aurantiomarginata
Encyclopedia
Mycena aurantiomarginata, commonly known as the golden-edge bonnet, is a species 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 Mycenaceae
Mycenaceae
The Mycenaceae are a family of fungi in the order Agaricales. According to the Dictionary of the Fungi , the family contains 10 genera and 705 species. This is one of several families that were separated from the Tricholomataceae as a result of phylogenetic analyses...

 family. It has a widespread distribution, common in Europe, and North America, and also having been collected in North Africa, Asia, and Central America. The fungus is saprobic, and produces 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 that grow on the floor
Forest floor
The forest floor, also called detritus, duff and the O horizon, is one of the most distinctive features of a forest ecosystem. It mainly consists of shed vegetative parts, such as leaves, branches, bark, and stems, existing in various stages of decomposition above the soil surface...

 of coniferous forests. The mushrooms have a bell-shaped 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...

 up to 2 cm (0.78740157480315 in) in diameter, a slender 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...

 up to 6 cm (2.4 in) long with yellow to orange fibrils at the base, and characteristic bright orange color of their gill edges. A microscopic characteristic is the club-shaped cystidia covered with numerous spiky projections that resemble a mace. The edibility
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...

 of the mushroom has not been determined. A 2010 publication reported the discovery and characterization of a novel pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

 named mycenaaurin A, isolated from the fruit body
Sporocarp (fungi)
In fungi, the sporocarp is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne...

 of the mushroom. The pigment is likely responsible for the color of the mushroom, and it has antibiotic
Antibiotic
An antibacterial is a compound or substance that kills or slows down the growth of bacteria.The term is often used synonymously with the term antibiotic; today, however, with increased knowledge of the causative agents of various infectious diseases, antibiotic has come to denote a broader range of...

 activity that may function in nature to prevent certain bacteria from growing on the fruit bodies.

Taxonomy, classification, phylogeny, and naming

The species, originally named Agaricus marginatus by the Danish naturalist 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...

 in 1803, has several synonyms
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

. Elias Magnus Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

 renamed it Agaricus aurantio-marginatus in his 1821 Systema Mycologicum
Systema Mycologicum
Systema Mycologicum is a systematic classification of fungi drawn up in 1821 by the Swedish mycologist and botanist Elias Fries. It took 11 years to complete....

, while Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 called it Agaricus schumacheri in 1828. Although Schumacher had the earliest publication date, Fries's name is sanctioned
Sanctioned name
In mycology, a sanctioned name is a name that was adopted in certain works of Christiaan Hendrik Persoon or Elias Magnus Fries, which are considered major points in fungal taxonomy.-Definition and effects:...

, and so the specific epithet he used is given nomenclatural precedence. 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 the species to the genus Mycena
Mycena
Mycena is a large genus of small saprotrophic mushrooms that are rarely more than a few centimeters in width. They are characterized by a white spore print, a small conical or bell-shaped cap, and a thin fragile stem. Most are gray or brown, but a few species have brighter colors. Most have a...

in 1872. In 1930 Karel Cejp considered it to be a variety of Mycena elegans.

According to Alexander H. Smith
Alexander H. Smith
Alexander Hanchett Smith was an American mycologist known for his extensive contributions to the taxonomy and phylogeny of the higher fungi, especially the agarics.-Early life:...

's organization of the genus Mycena, M. aurantiomarginata is classified in section Calodontes, subsection Granulatae, which contains species with roughened cheilocystidia, such as M. rosella
Mycena rosella
Mycena rosella, commonly known as the pink bonnet, is a species of mushroom in the Mycenaceae family. First called Agaricus roseus by Swedish mycologist Elias Magnus Fries in 1794, it was assigned its current name in 1871 by German scientist Paul Kummer.Microscopic characteristicsThe spores are...

, M. flavescens
Mycena flavescens
Mycena flavescens is a species of Mycenaceae fungus. It was first described scientifically by the Czech mycologist Josef Velenovský in 1920, based on specimens collected in Mnichovice in 1915. The mushroom is edible....

, M. elegans, and M. strobilinoides. In his 1992 study of Mycena, Dutch mycologist Rudolph Arnold Maas Geesteranus
Rudolph Arnold Maas Geesteranus
Rudolph Arnold Maas Geesteranus, born 20 January 1911 in The Hague, died May 18 2003 in Oegstgeest, was a Dutch mycologist.-References:...

 put M. aurantiomarginata in the section Luculentae, characterized by species with a olive to yellowish-olive and moist cap, pallid to gray-olive gills with bright orange margins, brownish to grayish-olive stem, white spore 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...

, and spiny cystidia. M. aurantiomarginata was included in a 2010 molecular analysis focused on clarifying the phylogenetic relationships between Northern European species in the section Calodontes. The results suggested that, based on similarity of nuclear large subunit ribosomal DNA
Ribosomal DNA
Ribosomal DNA codes for ribosomal RNA. The ribosome is an intracellular macromolecule that produces proteins or polypeptide chains. The ribosome itself consists of a composite of proteins and RNA. As shown in the figure, rDNA consists of a tandem repeat of a unit segment, an operon, composed of...

 sequences, the fungus is closely related to M. crocata
Mycena crocata
Mycena crocata is a species of mushroom in the Mycenaceae family. The common name Saffrondrop Bonnet refers to the red or orange latex that it exudes if the stipe is broken. Its habitat is woody debris and leaf litter in deciduous woodland, especially Beech....

and M. leaiana. This conclusion was corroborated in a 2009 publication that reported using molecular analysis to determine that several Mycena species can be mycorrhiza
Mycorrhiza
A mycorrhiza is a symbiotic association between a fungus and the roots of a vascular plant....

l partners of the orchid Gastrodia confusa.

The specific epithet aurantiomarginata is 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...

, and refers to the orange edges of its gills (aurantio, "orange"; marginata, "edge"). In the United Kingdom, the mushroom 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 "golden-edge bonnet".

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...

 of M. aurantiomarginata ranges in shape from obtusely conic to bell-shaped, and becomes flat in maturity, reaching diameters of 0.8 –. The cap color is variable, ranging from dark olive fuscous (dark brownish-gray) to yellowish-olive in the center, while the margin is orangish. Alexander H. Smith
Alexander H. Smith
Alexander Hanchett Smith was an American mycologist known for his extensive contributions to the taxonomy and phylogeny of the higher fungi, especially the agarics.-Early life:...

, in his 1947 monograph of North American Mycena species, states that the caps are not hygrophanous
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 depending on the level of hydration), while Mycena specialist Arne Aronsen says they are. The overall color fades as the mushroom ages. The surface is moist, and young individuals are covered with very fine whitish powder, but this soon sloughs off to leave a polished surface that develops radial grooves in maturity. 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 thin (about 1 mm thick in the center of the cap) and flexible. 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....

 with a decurrent tooth (where the gills curve up to join the stem but then, very close to the stem, the margin turns down again), and initially narrow but become broad when old. They are pallid to grayish-olive with bright orange edges. Smith notes that the edge color may spread to the gill faces in some specimens, because the pigment, rather than being encrusted on the walls of the cystidia, is found in the cytosol
Cytosol
The cytosol or intracellular fluid is the liquid found inside cells, that is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondrion into compartments....

 and therefore more readily diffusible. The gills are spaced close together, with between 16 to 26 gills reaching the stem, and there are up to three tiers of lamellulae (short gills that do not extend fully from the cap edge to the stem). The cylindrical 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 3 – by 0.1 –, hollow, and stiff but flexible. It has a brownish- to grayish-olive color that is sometimes tinged with shades of orange. The surface is smooth except for orange powder near the top, while the base is covered with stiff orange hairs. Smith reports the mushroom tissue to have no distinctive taste or odor, while Aronsen says the odor is "very conspicuous; sweet, fruity, often experienced as farinaceous or faintly of anise". Like many small Mycena species, the edibility
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...

 of the mushroom is unknown, as it is too insubstantial to consider collecting for the table.
The 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 elliptic, smooth, and amyloid
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...

, with dimensions of 7–9 by 4–5 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...

. 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 club-shaped, four-spored, and measure 25–32 by 5.5–7 μm. Pleurocystidia and cheilocystidia (cystidia on the gill faces and edges, respectively) are abundant and similar in morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

: club-shaped to somewhat capitate (with a head), the tops sparsely to densely with small spines (said to resemble a mace), filled with a bright orange pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

, and measuring 28–36 by 7–12 μm. The flesh of the cap is covered with a 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....

, on the surface of which are found scattered cystidia similar to those on the gills. Directly beneath the cuticle is a layer of enlarged cells, and beneath this are filamentous 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. 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.

Mycena aurantiomarginata uses a tetrapolar mating system
Mating in fungi
Mating in fungi is a complex process governed by mating types. Research on fungal mating has focused on only a few model species. Since not all of the fungi reproduce sexually and many that do are isogamous, the terms male and female do not apply to this kingdom...

, a system that prevents self-fertilization and ensures a high degree of genotypic
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...

 diversity. When the fungal mycelia is grown in culture on a petri dish
Petri dish
A Petri dish is a shallow glass or plastic cylindrical lidded dish that biologists use to culture cells or small moss plants. It was named after German bacteriologist Julius Richard Petri, who invented it when working as an assistant to Robert Koch...

, the colonies
Clonal colony
A clonal colony or genet is a group of genetically identical individuals that have grown in a given location, all originating vegetatively from a single ancestor. In plants, an individual in such a population is referred to as a ramet...

 are white, odorless, and typically have a central patch of congested aerial hyphae that grow upward from the colony surface, which abruptly become flattened to submerged, and occasionally form faint zone lines. The hyphae commonly form deposits of tiny amorphous crystals where they contact other mycelial fronts, especially where the hyphae are vegetatively
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...

 incompatible and destroy each other by lysis
Lysis
Lysis refers to the breaking down of a cell, often by viral, enzymic, or osmotic mechanisms that compromise its integrity. A fluid containing the contents of lysed cells is called a "lysate"....

.

Similar species

Mycena elegans is similar in appearance to M. aurantiomarginata, and some authors have considered them synonymous. M. elegans is larger, with a cap diameter up to 3.5 cm (1.4 in) and stem length up to 12 cm (4.7 in), darker, and has pale greenish-yellow colors on the gill edges and stems that stain dull reddish-brown in age. M. leaiana is readily distinguished from M. aurantiomarginata by the bright orange color of its fruit bodies, its growth on rotting wood and in clusters, and the presence of a gelatinous layer over its stem.

Habitat and distribution

Mycena aurantiomarginata is a saprobic fungus, deriving nutrients from decomposing organic matter found on the forest floor
Forest floor
The forest floor, also called detritus, duff and the O horizon, is one of the most distinctive features of a forest ecosystem. It mainly consists of shed vegetative parts, such as leaves, branches, bark, and stems, existing in various stages of decomposition above the soil surface...

. The fruit bodies of the fungus grow scattered, in groups, or in tufts under conifers (such as spruce
Spruce
A spruce is a tree of the genus Picea , a genus of about 35 species of coniferous evergreen trees in the Family Pinaceae, found in the northern temperate and boreal regions of the earth. Spruces are large trees, from tall when mature, and can be distinguished by their whorled branches and conical...

 and fir
Fir
Firs are a genus of 48–55 species of evergreen conifers in the family Pinaceae. They are found through much of North and Central America, Europe, Asia, and North Africa, occurring in mountains over most of the range...

) on moss
Moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...

 and needle carpets. In North America, it is found in Washington, Oregon
Oregon
Oregon is a state in the Pacific Northwest region of the United States. It is located on the Pacific coast, with Washington to the north, California to the south, Nevada on the southeast and Idaho to the east. The Columbia and Snake rivers delineate much of Oregon's northern and eastern...

, and California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

. It is also found in North Africa, and is widely distributed in western and northern Europe. In Central America, the mushroom has been collected on the summit of Cerro de la Muerte
Cerro de la Muerte
Cerro de la Muerte is the highest point in the Costa Rican section of the Inter-American Highway. Its name means "Mountain of Death", since in the past crossing the mountains from the Valle Central meant a three or four day journey, on foot or on horseback, and many ill-prepared travelers...

 in the Cordillera de Talamanca
Cordillera de Talamanca
The Cordillera de Talamanca is a mountain range that lies on the border between Costa Rica and Panama. Much of the range and the area around it is included in the La Amistad International Park, which also is shared between the two countries....

, Costa Rica, on leaf litter
Plant litter
Plant litter, leaf litter or tree litter is dead plant material, such as leaves, bark, needles, and twigs, that has fallen to the ground. Litter provides habitat for small animals, fungi, and plants, and the material may be used to construct nests. As litter decomposes, nutrients are released to...

 of Comarostaphylis arbutoides (a highly branched evergreen shrub or tree in the heath family). In 2010, it was reported from 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...

 in northern Japan, where it was found growing on Picea glehnii
Picea glehnii
Picea glehnii is a species of conifer in the Pinaceae family.It is found in Japan and Russia. It was named after Russian botanist Peter von Glehn.-References:* Conifer Specialist Group 1998. . Downloaded on 10 July 2007....

forest litter in early winter.

Bioactive compounds

In 2010, a pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

 compound was isolated and characterized from fruit bodies of Mycena aurantiomarginata and reported as new to science by Robert Jaeger and Peter Spiteller in the Journal of Natural Products
Journal of Natural Products
The Journal of Natural Products is a peer-reviewed scientific journal, co-published by the American Society of Pharmacognosy and the American Chemical Society.. The editor in chief is A. Douglas Kinghorn .- History :...

publication. The chemical, mycenaaurin A, is a polyene
Polyene
Polyenes are poly-unsaturated organic compounds that contain one or more sequences of alternating double and single carbon-carbon bonds. These double carbon-carbon bonds interact in a process known as conjugation, which results in an overall lower energy state of the molecule.Organic compounds with...

 compound that consists of a tridecaketide (i.e., 13 adjacent methylene
Methylene
Methylene is a chemical species in which a carbon atom is bonded to two hydrogen atoms. Three different possibilities present themselves:* the -CH2- substituent group: e.g., dichloromethane ....

 and carbonyl
Carbonyl
In organic chemistry, a carbonyl group is a functional group composed of a carbon atom double-bonded to an oxygen atom: C=O. It is common to several classes of organic compounds, as part of many larger functional groups....

 functional group
Functional group
In organic chemistry, functional groups are specific groups of atoms within molecules that are responsible for the characteristic chemical reactions of those molecules. The same functional group will undergo the same or similar chemical reaction regardless of the size of the molecule it is a part of...

s with two amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

 moieties on either end of the molecule). The authors posit that the flanking amino acid groups are probably biosynthetically derived from S-adenosylmethionine. The tridecaketide itself contains an alpha-pyrone
Pyrone
Pyrones or pyranones are a class of cyclic chemical compounds. They contain an unsaturated six membered ring containing one oxygen atom and a ketone functional group. There are two isomers denoted as 2-pyrone and 4-pyrone. The 2-pyrone structure is found in nature as part of the coumarin ring...

, a conjugated
Conjugated system
In chemistry, a conjugated system is a system of connected p-orbitals with delocalized electrons in compounds with alternating single and multiple bonds, which in general may lower the overall energy of the molecule and increase stability. Lone pairs, radicals or carbenium ions may be part of the...

 hexaene, and a single alkenyl
Alkene
In organic chemistry, an alkene, olefin, or olefine is an unsaturated chemical compound containing at least one carbon-to-carbon double bond...

 moiety. Jaeger and Spiteller suggest that mycenaaurin A might function as a defense compound, since it has antibacterial activity against the Gram-positive
Gram-positive
Gram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...

 bacterium Bacillus pumilus
Bacillus pumilus
Bacillus pumilus is a Gram-positive aerobic spore-forming bacillus commonly found in soil. B. pumilus spores - with the exception of mutant strain ATCC 7061 - generally show high resistance to environmental stresses including UV light exposure, dessication, and the presence of oxidizers such as...

. The chemical is only present in the fruit bodies, and not in the colorless mycelia. An earlier screening for antimicrobial activity revealed a weak ability to inhibit the growth of the fungi Candida albicans
Candida albicans
Candida albicans is a diploid fungus that grows both as yeast and filamentous cells and a causal agent of opportunistic oral and genital infections in humans. Systemic fungal infections including those by C...

and Aspergillus fumigatus
Aspergillus fumigatus
Aspergillus fumigatus is a fungus of the genus Aspergillus, and is one of the most common Aspergillus species to cause disease in individuals with an immunodeficiency....

.
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