Fomes fomentarius
Encyclopedia
Fomes fomentarius is a species of fungal plant pathogen found in Europe, Asia, Africa and North America. The species produces very large polypore
Polypore
Polypores are a group of tough, leathery poroid mushrooms similar to boletes, but typically lacking a distinct stalk. The technical distinction between the two types of mushrooms is that polypores do not have the spore-bearing tissue continuous along the entire underside of the mushroom. Many...

 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...

 which are shaped like a horse's hoof
Horse hoof
A horse hoof is a structure surrounding the distal phalanx of the 3rd digit of each of the four limbs of Equus species, which is covered by complex soft tissue and keratinised structures...

 and vary in colour from a silvery grey to almost black, though they are normally brown. It grows on the side of various species of tree, which it infects through broken bark, causing rot
Wood-decay fungus
A wood-decay fungus is a variety of fungus that digests moist wood, causing it to rot. Some wood-decay fungi attack dead wood, such as brown rot, and some, such as Armillaria , are parasitic and colonize living trees. Fungi that not only grow on wood but actually cause it to decay, are called...

. The species typically continues to live on trees long after they have died, changing from a parasite to a decomposer
Decomposer
Decomposers are organisms that break down dead or decaying organisms, and in doing so carry out the natural process of decomposition. Like herbivores and predators, decomposers are heterotrophic, meaning that they use organic substrates to get their energy, carbon and nutrients for growth and...

.

Though inedible, F. fomentarius has traditionally seen use as the main ingredient of amadou
Amadou
Amadou is a spongy, flammable substance prepared from bracket fungi. The species generally used is Fomes fomentarius which in English is also called horse's hoof fungus or tinder fungus. The amadou layer can be found on top of the fungus just below the outer skin and above the pores...

, a material used primarily as tinder
Tinder
Tinder is easily combustible material used to ignite fires by rudimentary methods. A small fire consisting of tinder is then used to ignite kindling. Anything that can be ignited by a match can be considered tinder; or by more rigorous definition, anything that begins to glow under a shower of...

, but also used to make clothing and other items. The 5,000-year-old Ötzi the Iceman
Ötzi the Iceman
Ötzi the Iceman , Similaun Man, and Man from Hauslabjoch are modern names for a well-preserved natural mummy of a man who lived about 5,300 years ago. The mummy was found in September 1991 in the Ötztal Alps, near Hauslabjoch on the border between Austria and Italy. The nickname comes from the...

 carried four pieces of F. fomentarius, concluded to be for use as tinder. It also has medicinal and other uses. The species is both a pest and useful in timber production.

Taxonomy

The first scientific description of the fungus appeared in the literature in the 1753 Species Plantarum
Species Plantarum
Species Plantarum was first published in 1753, as a two-volume work by Carl Linnaeus. Its prime importance is perhaps that it is the primary starting point of plant nomenclature as it exists today. This means that the first names to be considered validly published in botany are those that appear...

by Carl Linnaeus; he called it Boletus fomentarius. The specific epithet fomentarius is 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...

 fomentum, referring to tinder
Tinder
Tinder is easily combustible material used to ignite fires by rudimentary methods. A small fire consisting of tinder is then used to ignite kindling. Anything that can be ignited by a match can be considered tinder; or by more rigorous definition, anything that begins to glow under a shower of...

. The species has been described as a member of numerous different genera. In 1783, Jean-Baptiste Lamarck
Jean-Baptiste Lamarck
Jean-Baptiste Pierre Antoine de Monet, Chevalier de la Marck , often known simply as Lamarck, was a French naturalist...

 named the species Agaricus fomentarius in his Encyclopédie Méthodique: Botanique. In 1818, Georg Friedrich Wilhelm Meyer described Polyporus fomentarius in his Primitiae Florae Essequeboensis, and this name was 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:...

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

 in the 1821 publication of the first volume of his 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....

. Fries later, in his 1849 Summa vegetabilium Scandinaviae, moved the species to the genus Fomes
Fomes
Fomes is a genus of perennial woody fungi in the family Polyporaceae. Species are typically hoof-shaped . New growth each season is added to the margin, resulting in a downward extension of the hymenium. This often results in a zonate appearance of the upper surface, that is, marked by concentric...

. Subsequent attempts to change the genus of the species have been unsuccessful; the species was named Placodes fomentarius by 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....

 in 1886, Ochroporus fomentarius by Joseph Schröter
Joseph Schröter
Joseph Schröter was a noted German mycologist, doctor and scientist. During his lifetime, he wrote several books and texts, and discovered and described many species of flora and fungi...

 in 1888 and Scindalma fomentarium by Otto Kuntze
Otto Kuntze
Otto Carl Ernst Kuntze was a German botanist.-Biography:Otto Kuntze was born in Leipzig.An apothecary in his early career, he published an essay entitled Pocket Fauna of Leipzig. Between 1863 and...

 in 1898. In the twentieth century, Narcisse Théophile Patouillard
Narcisse Théophile Patouillard
Narcisse Théophile Patouillard was a French pharmacist and mycologist.He was born in Macornay, a town in the department of Jura...

 named the species Ungulina fomentaria in 1900, and William Murrill
William Murrill
William Alphonso Murrill was an American mycologist, known for his contributions to the knowledge of the Agaricales and Polyporaceae.- Education :...

 twice reallocated the species; in 1903, he named it Elfvingia fomentaria and in 1914, he named it Elfvingiella fomentaria. In 1963, Shu Chün Teng named it Pyropolyporus fomentarius. These names are considered obligate synonyms; that is, different names for the same species based on a single description or specimen. In addition to the obligate synonyms, there are a number of taxonomic synonyms, whereby names have been described as separate species, but have come to be considered synonymous. The species is commonly known
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...

 as the Tinder Fungus, Hoof Fungus, Tinder Polypore or Ice Man Fungus.

Description

Fomes fomentarius has a fruit body of between 5 and 45 cm (2 and 17.7 in) across, 3 and 25 cm (1.2 and 9.8 in) wide and 2 and 25 cm (0.78740157480315 and 9.8 in) thick, which attaches broadly to the tree on which the fungus is growing. While typically shaped like a horse's hoof
Horse hoof
A horse hoof is a structure surrounding the distal phalanx of the 3rd digit of each of the four limbs of Equus species, which is covered by complex soft tissue and keratinised structures...

, it can also be more bracket
Bracket fungus
Bracket fungi, or shelf fungi, among many groups of the fungi in the phylum Basidiomycota. Characteristically, they produce shelf- or bracket-shaped fruiting bodies called conks that lie in a close planar grouping of separate or interconnected horizontal rows...

-like with an umbonate attachment to the substrate
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...

. The species typically has broad, concentric ridges, with a blunt and rounded margin. 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 hard and fibrous, and a cinnamon
Cinnamon
Cinnamon is a spice obtained from the inner bark of several trees from the genus Cinnamomum that is used in both sweet and savoury foods...

 brown colour. The upper surface is tough, bumpy, hard and woody, varying in colour, usually a light brown or grey. The margin is whitish during periods of growth. The hard crust is from 1 to 2 mm (0.0393700787401575 to 0.078740157480315 in) thick, and covers the tough flesh. The underside has round pores of a cream colour when new, maturing to brown, though they darken when handled. The pores are circular, and there are 2–3 per millimetre. The tubes are 2 to 7 mm (0.078740157480315 to 0.275590551181102 in) long and a rusty brown colour.

The colouration and size of the fruit body can vary based on where the specimen has grown. Silvery-white, greyish and nearly black specimens have been known. The darkest fruit bodies were previously classified as Fomes nigricans, but this is now recognised as a synonym of Fomes fomentarius. The colour is typically lighter at lower latitudes and altitudes, as well as on fruit bodies in the Northern Hemisphere
Northern Hemisphere
The Northern Hemisphere is the half of a planet that is north of its equator—the word hemisphere literally means “half sphere”. It is also that half of the celestial sphere north of the celestial equator...

 that grow on the south side of trees. However, studies have concluded that there is no reliable way to differentiate varieties; instead, the phenotypic
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...

 differences can "be attributed either to different ecotypes or to interactions between the genotype and its environment".

Microscopic features

The spores are lemon-yellow in colour, and oblong-ellipsoid in shape. They measure 15–20 by 5–7 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 species has a trimitic hyphal structure (meaning that it has generative, skeletal and binding hyphae), with generative hyphae (hyphae that are relatively undifferentiated and can develop reproductive structures) with clamp connections.

Similar species

Fomes fomentarius can easily be confused with Phellinus igniarius
Phellinus igniarius
Phellinus igniarius is a plant pathogen.- External links :*** by Diane Pleninger and Tom Volk-References:* Phellinus igniarius, Iqmik, used by native Americans with tobacco—by Diane Pleninger and Tom Volk...

, species from the genus Ganoderma
Ganoderma
Ganoderma is a genus of polypore mushrooms which grow on wood and include about 80 species, many from tropical regions. Because of their extensive use in traditional Asian medicines, and their potential in bioremediation, they are a very important genus economically. Ganoderma can be differentiated...

and Fomitopsis pinicola
Fomitopsis pinicola
Fomitopsis pinicola, in English sometimes known as Red Banded Polypore, is a polypore mushroom of the genus Fomitopsis. The species is common throughout the temperate Northern hemisphere. An alternative binomial name is Fomes pinicola....

. An easy way to differentiate F. fomentarius is by adding a drop of potassium hydroxide
Potassium hydroxide
Potassium hydroxide is an inorganic compound with the formula KOH, commonly called caustic potash.Along with sodium hydroxide , this colorless solid is a prototypical strong base. It has many industrial and niche applications. Most applications exploit its reactivity toward acids and its corrosive...

 onto a small piece of the fruit body from the upper surface. The solution will turn a dark blood red if the specimen is F. fomentarius, due to the presence of the chemical fomentariol.

Habitat and distribution

F. fomentarius has a circumboreal distribution, being found in both northern and southern Africa, throughout Asia and into eastern North America, and throughout Europe, and is frequently encountered. The optimal temperature for the species's growth is between 27 and 30 °C (80.6 and 86 F) and the maximum is between 34 and 38 °C (93.2 and 100.4 F). F. fomentarius typically grows alone, but multiple fruit bodies can sometimes be found upon the same host trunk. The species most typically grows upon hardwoods. In northern areas, it is most common on birch
Birch
Birch is a tree or shrub of the genus Betula , in the family Betulaceae, closely related to the beech/oak family, Fagaceae. The Betula genus contains 30–60 known taxa...

, while, in the south, beech
Beech
Beech is a genus of ten species of deciduous trees in the family Fagaceae, native to temperate Europe, Asia and North America.-Habit:...

 is more typical. In the Mediterranean, oak
Oak
An oak is a tree or shrub in the genus Quercus , of which about 600 species exist. "Oak" may also appear in the names of species in related genera, notably Lithocarpus...

 is the typical host. The species has also been known to grow upon maple
Maple
Acer is a genus of trees or shrubs commonly known as maple.Maples are variously classified in a family of their own, the Aceraceae, or together with the Hippocastanaceae included in the family Sapindaceae. Modern classifications, including the Angiosperm Phylogeny Group system, favour inclusion in...

, cherry
Cherry Tree
Cherry Tree may refer to:* A tree that produces cherries* An ornamental cherry tree that produces cherry blossomsPlaces* Cherry Tree, Pennsylvania, a borough in Indiana County, Pennsylvania, United States* Cherry Tree, Oklahoma...

, hickory
Hickory
Trees in the genus Carya are commonly known as hickory, derived from the Powhatan language of Virginia. The genus includes 17–19 species of deciduous trees with pinnately compound leaves and big nuts...

, lime tree, poplar
Poplar
Populus is a genus of 25–35 species of deciduous flowering plants in the family Salicaceae, native to most of the Northern Hemisphere. English names variously applied to different species include poplar , aspen, and cottonwood....

, willow
Willow
Willows, sallows, and osiers form the genus Salix, around 400 species of deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions of the Northern Hemisphere...

, alder
Alder
Alder is the common name of a genus of flowering plants belonging to the birch family . The genus comprises about 30 species of monoecious trees and shrubs, few reaching large size, distributed throughout the North Temperate Zone and in the Americas along the Andes southwards to...

, hornbeam
Hornbeam
Hornbeams are relatively small hardwood trees in the genus Carpinus . Though some botanists grouped them with the hazels and hop-hornbeams in a segregate family, Corylaceae, modern botanists place the hornbeams in the birch subfamily Coryloideae...

, sycamore, and even, exceptionally, softwoods, such as conifers.

Ecology

Fomes fomentarius is a stem decay
Wood-decay fungus
A wood-decay fungus is a variety of fungus that digests moist wood, causing it to rot. Some wood-decay fungi attack dead wood, such as brown rot, and some, such as Armillaria , are parasitic and colonize living trees. Fungi that not only grow on wood but actually cause it to decay, are called...

 plant pathogen. The species' mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 penetrates the wood of trees through damaged bark or broken branches, causing rot in the host. It can grow on the bark wound, or even directly onto the bark of older or dead trees. The decayed wood shows black lines in the lightly coloured decayed areas; these are known as pseudosclerotic layers or demarcation lines. The lines are caused by enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

s called phenoloxidases, converting either fungus or plant matter into melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...

. The lines are not an absolute identifier, as they can also occur in plants infected by Kretzschmaria deusta
Kretzschmaria deusta
Kretzschmaria deusta, commonly known as brittle cinder, is a fungus and plant pathogen found in temperate regions of the Northern Hemisphere. It is common on a wide range of broadleaved trees including beech , oak , lime , and maple...

and some Armillaria species. Despite beginning as a parasite, the species is able to survive for a time (hastening decomposition
Decomposition
Decomposition is the process by which organic material is broken down into simpler forms of matter. The process is essential for recycling the finite matter that occupies physical space in the biome. Bodies of living organisms begin to decompose shortly after death...

) on fallen or felled trees as a saprotrophic feeder, and typically lives there for years, until the log is completely destroyed. It is also capable of colonising and breaking down pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 grains, giving it a second food source which is particular high in nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

. Infected trees become very brittle, and cracks can occur in the affected tree due to wind. F. fomentarius is particularly adept at moving between cracks on the tree without interruption. However, in addition to the obviously infected damaged trees, F. fomentarius is known to be an endophyte
Endophyte
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life without causing apparent disease. Endophytes are ubiquitous and have been found in all the species of plants studied to date; however, most of these endophyte/plant relationships...

, meaning that healthy trees which are not sporting F. fomentarius fruit bodies could still be infected.

The fruit bodies are perennial, surviving for up to thirty years. The strongest growth period is between early summer and autumn. The yearly growth always occurs on the bottom of the fungus, meaning that the lowest layer is the youngest. This occurs even if the host tree has been laid on the forest floor, which can happen because of the white rot induced by the fungus. This is a process known as positive gravitropism
Gravitropism
Gravitropism is a turning or growth movement by a plant or fungus in response to gravity. Charles Darwin was one of the first to scientifically document that roots show positive gravitropism and stems show negative gravitropism. That is, roots grow in the direction of gravitational pull and stems...

. Very large numbers of spores are produced, particularly in spring, with up to 887 million basidiospores an hour being produced by some fruit bodies. Spore production also takes place in autumn, though not nearly as heavily. The spores are released at comparatively low temperatures. In dry weather, the spores are visible as a white powder.

Uses and importance

The species is not 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...

; the flesh has an acrid taste, with a slightly fruity smell. The fungus has economic significance as it removes any timber
Timber
Timber may refer to:* Timber, a term common in the United Kingdom and Australia for wood materials * Timber, Oregon, an unincorporated community in the U.S...

 value of infected trees. As Fomes fomentarius infects trees through damaged bark, it will often infect trees already weakened from beech bark disease
Beech Bark Disease
Beech bark disease is a disease that causes mortality and defects in beech trees in the eastern United States and Europe. In North America, the disease results when the beech scale insect, Cryptococcus fagisuga, attacks the bark, creating a wound. Later, two different fungi Beech bark disease is a...

. However, it is too weakly parasitic to infect healthy trees, and so can be regarded merely as an aspect of the ecosystem, with the important and useful role of decomposing unusable timber.

Amadou

The species is well known for its uses in making fire
Making fire
Fire was an essential tool in early human cultural development and still important today. Many different techniques for making fire exist...

. This species, as well as others, such as Phellinus igniarius
Phellinus igniarius
Phellinus igniarius is a plant pathogen.- External links :*** by Diane Pleninger and Tom Volk-References:* Phellinus igniarius, Iqmik, used by native Americans with tobacco—by Diane Pleninger and Tom Volk...

, can be used to make amadou
Amadou
Amadou is a spongy, flammable substance prepared from bracket fungi. The species generally used is Fomes fomentarius which in English is also called horse's hoof fungus or tinder fungus. The amadou layer can be found on top of the fungus just below the outer skin and above the pores...

, a tinder
Tinder
Tinder is easily combustible material used to ignite fires by rudimentary methods. A small fire consisting of tinder is then used to ignite kindling. Anything that can be ignited by a match can be considered tinder; or by more rigorous definition, anything that begins to glow under a shower of...

. Amadou is produced from the flesh of the fruit bodies. The young fruit bodies are soaked in water before being cut into strips, and are then beaten and stretched, separating the fibres. The resulting material is referred to as "red amadou". The addition of gunpowder
Gunpowder
Gunpowder, also known since in the late 19th century as black powder, was the first chemical explosive and the only one known until the mid 1800s. It is a mixture of sulfur, charcoal, and potassium nitrate - with the sulfur and charcoal acting as fuels, while the saltpeter works as an oxidizer...

 or nitre produced an even more potent tinder. The flesh was further used to produce clothing, including caps, gloves and breeches. Amadou was used medicinally by dentists, who used it to dry teeth, and surgeons, who used it as a styptic. It is still used today in fly fishing
Fly fishing
Fly fishing is an angling method in which an artificial 'fly' is used to catch fish. The fly is cast using a fly rod, reel, and specialized weighted line. Casting a nearly weightless fly or 'lure' requires casting techniques significantly different from other forms of casting...

 for drying the flies
Artificial fly
An artificial fly or fly lure is a type of fishing lure, usually used in the sport of fly fishing . In general, artificial flies are the bait which fly fishers present to their target species of fish while fly fishing...

. Other items of clothing and even picture frames and ornaments have been known to be made from the fungus in Europe, particularly Bohemia
Bohemia
Bohemia is a historical region in central Europe, occupying the western two-thirds of the traditional Czech Lands. It is located in the contemporary Czech Republic with its capital in Prague...

. The fungus is known to have been used as a firestarter in Hedeby
Hedeby
Hedeby |heath]]land, and býr = yard, thus "heath yard"), mentioned by Alfred the Great as aet Haethe , in German Haddeby and Haithabu, a modern spelling of the runic Heiðabý was an important trading settlement in the Danish-northern German borderland during the Viking Age...

, and it is known that the fungus was used as early as 3000 BCE. When found, the 5,000-year-old Ötzi the Iceman
Ötzi the Iceman
Ötzi the Iceman , Similaun Man, and Man from Hauslabjoch are modern names for a well-preserved natural mummy of a man who lived about 5,300 years ago. The mummy was found in September 1991 in the Ötztal Alps, near Hauslabjoch on the border between Austria and Italy. The nickname comes from the...

was carrying four pieces of F. fomentarius fruit body. Chemical tests led to the conclusion that he carried it for use as tinder.
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