Armillaria gallica

Armillaria gallica (synonymous with A. bulbosa and A. lutea) is a species of

Armillaria gallica
honey mushroom in the family Physalacriaceae of the order Agaricales. The
species is a common and ecologically important wood-decay fungus that can live
as a saprobe, or as an opportunistic parasite in weakened tree hosts to cause root
or butt rot. It is found in temperate regions of Asia, North America, and Europe.
The species forms fruit bodies singly or in groups in soil or rotting wood. The
fungus has been inadvertently introduced to South Africa. Armillaria gallica has
had a confusing taxonomy, due in part to historical difficulties encountered in
distinguishing between similar Armillaria species. The fungus received
international attention in the early 1990s when an individual colony living in a
Michigan forest was reported to cover an area of 15 hectares (37 acres), weigh at
least 9,500 kilograms (21,000 lb), and be 1,500 years old. This individual is Scientific classification
popularly known as the "humungous fungus", and is a tourist attraction and Kingdom: Fungi
inspiration for an annual mushroom-themed festival inCrystal Falls.
Division: Basidiomycota
Armillaria gallica is a largely subterranean fungus, and it produces fruit bodies
Class: Agaricomycetes
that are up to about 10 cm (3.9 in) in diameter, yellow-brown, and covered with
small scales. On the underside of the caps are gills that are white to creamy or Order: Agaricales
pale orange. The stem may be up to 10 cm (3.9 in) long, with a white cobwebby Family: Physalacriaceae
ring that divides the color of the stem into pale orange to brown above, and
Genus: Armillaria
lighter-colored below. The fungus can develop an extensive system of
underground root-like structures, called rhizomorphs, that help it to efficiently Species: A. gallica
decompose dead wood in temperate broadleaf and mixed forests. It has been the Binomial name
subject of considerable scientific research due to its importance as a plant
Armillaria gallica
pathogen, its ability to bioluminesce, its unusual life cycle, and its ability to form
large and long-lived colonies. Marxm. & Romagn.

Synonyms

Armillaria bulbosa (Barla) Kile & Watling

Contents Armillaria inflata Velen.

Armillaria lutea Gillet
Phylogeny, taxonomy and naming Armillaria mellea var. bulbosa Barla
Description Armillariella bulbosa (Barla) Romagn.
Microscopic features
Edibility
Similar species
Armillaria gallica
Metabolites
Bioluminescence Mycological characteristics
Humongous fungus gills on hymenium
Life cycle and growth
Habitat and distribution cap is convex

Ecology
hymenium is adnate
See also
References stipe has a ring
External links
spore print is white
Phylogeny, taxonomy and naming
Confusion has surrounded the ecology is saprotrophic
A. solidipes nomenclature and taxonomy of
the species now known as or parasitic
A. gemina
Armillaria gallica, paralleling edibility: edible
that surrounding the genus
A. calvescens
Armillaria.[2] The type species, Armillaria mellea, was until the 1970s believed to

A. gallica (ST22) be a pleiomorphic species with a wide distribution, variable pathogenicity, and one
of the broadest host ranges known for the fungi.[3] In 1973, Veikko Hintikka
A. gallica (ST23) reported a technique to distinguish between Armillaria species by growing them
together as single spore isolates on petri dishes and observing changes in the
A. gallica (M70) morphology of the cultures.[4] Using a similar technique, Kari Korhonen showed
in 1978 that the European Armillaria mellea species complex could be separated
A. sinapina into five reproductively isolated species, which he named "European Biological
Species" (EBS) A through E.[5] About the same time, the North American
A. cepistipes A. mellea was shown to be ten different species (North American Biological
Species, or NABS I through X);[6] NABS VII was demonstrated shortly after to be
NABS X
the same species as EBS E.[7] Because several research groups had worked with
this widely distributed species, it was assigned several dif
ferent names.
A. nabsnona
The species that Korhonen called EBS B was named A. bulbosa by Helga
A. tabescens
Marxmüller in 1982,[8] as it was thought to be equivalent to Armillaria mellea var.
bulbosa, first described by Joseph Barla in 1887,[9] and later raised to species
A. mellea
status by Josef Velenovský in 1927.[10] In 1973, the French mycologist Henri
Romagnesi, unaware of Velenovský's publication, published a description of the
Phylogeny and relationships of
A. gallica and related North American species he called Armillariella bulbosa, based on specimens he had found near
species based on amplified fragment Compiègne and Saint-Sauveur-le-Vicomte in France. These specimens were later
length polymorphism data. SY22,
demonstrated to be the same species as the EBS E of Korhonen; EBS B was later
ST23, and M70 are A. gallica
specimens collected from Michigan, determined to be A. cepistipes.[11] Therefore, the name A. bulbosa was a
Wisconsin, and British Columbia, misapplied name for EBS E. In 1987 Romagnesi and Marxmüller renamed EBS E
respectively.[1] to Armillaria gallica.[12] Another synonym, A. lutea, had originally been described
by Claude Casimir Gillet in 1874,[13] and proposed as a name for EBS E.[14][15] Although the name had priority due to its early
publication date, it was rejected as a nomen ambiguum because of a lack of supporting evidence to identify the fungus, including a
specimen, type locality, and incomplete collection notes.[11] A. inflata (Velenovský, 1920) may represent another synonym, but the
type specimens were not preserved, so it is considered a dubious name (nomen dubium).[16] As of 2010, both the Index Fungorum
and MycoBank consider Armillaria gallica Marxm. & Romagn. to be the current name, with A. bulbosa and A. lutea as
synonyms.[17][18]

Phylogenetic analysis of North American Armillaria species based on analysis of amplified fragment length polymorphism data
suggests that A. gallica is most closely related to A. sinapina, A. cepistipes, and A. calvescens.[1] These results are similar to those
reported in 1992 that compared sequences of nuclearribosomal DNA.[19]

The specific epithet gallica is botanical Latin for "French" (from Gallia, "Gaul"),[20] and refers to the type locality.[21] The prior
name bulbosa is Latin for "bulb-bearing, bulbous" (from bulbus and the suffix -osa).[20][21] Armillaria is derived from the Latin
armilla, or "bracelet".[22]

Description
 The fruit bodies of Armillaria gallica have caps that are 2.5–9.5 cm (1.0–3.7 in) broad,
and depending on their age, may range in shape from conical to convex to flattened. The
caps are brownish-yellow to brown when moist, often with a darker-colored center; the
color tends to fade upon drying. The cap surface is covered with slender fibers (same
color as the cap) that are erect, or sloping upwards.

Young fruit bodies have a cottony
partial veil that protects the
developing gills.
When the fruit bodies are young, the underside of the caps have a cottony layer of tissue
stretching from the edge of the cap to the stem—a partial veil—which serves to protect
the developing gills. As the cap grows in size the membrane is eventually pulled away
from the cap to expose the gills. The gills have an adnate (squarely attached) to
somewhat decurrent (extending down the length of the stem) attachment to the stem.
They are initially white, but age to a creamy or pale orange covered with rust-colored
spots. The stem is 4–10 cm (1.6–3.9 in) long and 0.6–1.8 cm (0.24–0.71 in) thick, and
almost club-shaped with the base up to 1.3–2.7 cm (0.5–1.1 in) thick. Above the level of
the ring, the stem is pale orange to brown, while below it is whitish or pale pink,
becoming grayish-brown at the base. The ring is positioned about 0.4–0.9 cm (0.16–
0.35 in) below the level of the cap, and may be covered with yellowish to pale-brownish
woolly cottony mycelia. The base of the stem is attached torhizomorphs, black root-like

Mature gills structures 1–3 mm in diameter. While the primary function of the below-ground mycelia
is to absorb nutrients from the soil, the rhizomorphs serve a more exploratory function,
to locate new food bases.[23][24]

Microscopic features
When the spores are seen in deposit, such as with a spore print, they appear whitish. They have an ellipsoid or oblong shape, usually
contain an oil droplet, and have dimensions of 7–8.5 by 5–6 µm. The spore-bearing cells, the basidia, are club-shaped, four-spored
(rarely two-spored), and measure 32–43 by 7–8.7 µm.[25] Other cells present in the fertile hymenium include the cheilocystidia
(cystidia present on the edge of a gill), which are club-shaped, roughly cylindrical and 15–25 by 5.0–12 µm. Cystidia are also present
on the stem (called caulocystidia), and are broadly club-shaped, measuring 20–55 by 11–23 µm.[26] The cap cuticle is made of
hyphae that are irregularly interwoven and project upward to form the scales seen on the surface. The hyphae that make up the
surface scales typically measure 26–88 µm long by 11–27 µm thick and can be covered with a crust of pigment. Clamp connections
are present in the hyphae of most tissues.[25]

Edibility
Like all Armillaria species, A. gallica is considered edible. Thorough cooking is usually recommended, as the raw mushroom tastes
acrid when fresh or undercooked.[23] One author advises to consume only a small portion initially, as some people may experience an
upset stomach.[27] The taste is described as "mild to bitter", and the odor "sweet",
[28] or reminiscent of camembert cheese.[26]

Similar species
Armillaria calvescens is rather similar in appearance, and can only be reliably distinguished from A. gallica by observing
microscopic characteristics. A. calvescens has a more northern distribution, and in North America, is rarely found south of the Great
Lakes.[28] A. mellea has a thinner stem than A. gallica, but can be more definitively distinguished by the absence of clamps at the
base of the basidia.[29] Similarly, A. cepistipes and A. gallica are virtually identical in appearance (especially older fruit bodies), and
are identified by differences in geographical distribution, host range, and microscopic characteristics. Molecular methods have been
developed to discriminate between the two species by comparing DNA sequences in the gene coding translation elongation factor 1-
alpha.[26]
Metabolites
Armillaria gallica can produce cyclobutane-containing metabolites such as arnamiol,[30] a natural product that is classified as a
sesquiterpenoid aryl ester.[31] Although the specific function of arnamiol is not definitively known, similar chemicals present in other
Armillaria species are thought to play a role in inhibiting the growth of antagonistic bacteria or fungi, or in killing cells of the host
plant prior to infection.[32]

Bioluminescence
The mycelia (but not the fruit bodies) of Armillaria gallica are known to be bioluminescent. Experiments have shown that the
intensity of the luminescence is enhanced when the mycelia are disturbed during growth or when they are exposed to fluorescent
light.[33] Bioluminescence is caused by the action of luciferases, enzymes that produce light by the oxidation of a luciferin (a
pigment).[34] The biological purpose of bioluminescence in fungi is not definitively known, although several hypotheses have been
suggested: it may help attract insects to help with spore dispersal,[35] it may be a by-product of other biochemical functions,[36] or it
may help deter heterotrophs that might consume the fungus.[35]

Humongous fungus
Researchers reported finding Armillaria gallica in the Upper Peninsula of
Michigan in the early 1990s, during an unrelated research project to study
the possible biological effects of extremely low frequency radio stations,
which were being investigated as a means to communicate with submerged
submarines. In one particular forest stand, Armillaria-infected oak trees had
been harvested, and their stumps were left to rot in the field. Later, when red
pines were planted in the same location, the seedlings were killed by the
fungus, identified as A. gallica (then known as A. bulbosa). Using molecular
genetics, they determined that the underground mycelia of one individual
fungal colony covered 15 ha (37 acres), weighing over 9,500 kilograms
(21,000 lb), with an estimated age of 1,500 years.[37][38] The analysis used The fruit bodies—the visible manifestation of
restriction fragment length polymorphism (RFLP) and random amplification A. gallica—belie an extensive underground
network of mycelia.
of polymorphic DNA (RAPD) to examine isolates collected from fruit
bodies and rhizomorphs (underground aggregations of fungal cells that
resemble plant roots) along 1-kilometer (0.6 mi) transects in the forest. The 15-hectare area yielded isolates that had identical mating
type alleles and mitochondrial DNA restriction fragment patterns; this degree of genetic similarity indicated that the samples were all
derived from a single genetic individual, or clone, that had reached its size through vegetative growth. In their conclusion the authors
noted: "This is the first report estimating the minimum size, mass, and age of an unambiguously defined fungal individual. Although
the number of observations for plants and animals is much greater, members of the fungal kingdom should now be recognized as
among the oldest and largest organisms on earth."[39] After the Nature paper was published, major media outlets from around the
world visited the site where the specimens were found; as a result of this publicity, the individual acquired the common name
"humongous fungus".[38] There was afterward some scholarly debate as to whether the fungus qualified to be considered in the same
category as other large organisms such as the blue whale or the giant redwood.[40]

The fungus has since become a popular tourist attraction in Michigan, and has inspired a "Humongous Fungus Fest" held annually in
August in Crystal Falls.[41] The organism was the subject of a Late Show Top Ten List on Late Night with David Letterman,[42] and
an advertising campaign by the rental companyU-Haul.[38]

Life cycle and growth

The life cycle of A. gallica includes two diploidization–haploidization events. The first of these is the usual process of cell fusion
(forming a diploid) followed by meiosis during the formation of haploid basidiospores.[43] The second event is more cryptic, and
occurs before fruit body formation. In most basidiomycetous fungi, the hyphae of compatible mating types will fuse to form a two-
nucleate, or dikaryotic stage; this stage is not observed in Armillaria species, which have cells that are mostly monokaryotic and
diploid. Genetic analyses suggest that the dikaryotic mycelia undergo an extra haploidization event prior to fruit body formation to
create a genetic mosaic.[44] These regular and repeating haploidization events result in increased genetic diversity, which helps the
[45][46][47]
fungus to adapt to unfavorable changes in environmental conditions, like drought.

The growth rate of A. gallica rhizomorphs is between 0.3 and 0.6 m (1.0 and 2.0 ft) per year.[48] Population genetic studies of the
fungus conducted in the 1990s demonstrated that genetic individuals grow mitotically from a single point of origin to eventually
occupy territories that may include many adjacent root systems over large areas (several hectares) of forest floor.[39][49][50] Based on
the low mutation rates observed in large, long-lived individuals,A. gallica appears to have an especially stablegenome.[51] It has also
been hypothesized that genetic stability may result from self-renewing mycelial repositories of nuclei with stem cell-like
properties.[52]

Habitat and distribution

Armillaria gallica can normally be found on the ground, but sometimes on stumps
and logs.[53] Mushrooms that appear to be terrestrial are attached to plant roots
underneath the surface.[28] It is widely distributed and has been collected in North
America, Europe,[27] and Asia (China,[54] Iran,[55] and Japan[56] ). The species has
also been found in theWestern Cape Province of South Africa, where it is thought to
have been introduced from potted plants imported from Europe during the early
colonization of Cape Town.[57] In Scandinavia, it is absent in areas with very cold
climates, like Finland or Norway, but it is found in southern Sweden. It is thought to
be the most prevalent low altitude species of Armillaria in Great Britain and France.
Young fruit bodies growing in clusters
The upper limits of its altitude vary by region. In the French Massif Central, it is
at the base of a tree
found up to 1,100 m (3,600 ft), while in Bavaria, which has a more continental
climate, the upper limit of distribution reaches 600 m (2,000 ft).[58] In Serbian
forests, it is the most common Armillaria between elevations of 70 to 1,450 m (230 to 4,760 ft).[59] Field studies suggest that
A. gallica prefers sites that are low inorganic matter and have high soil pHs.[60][61]

In North America, it is common east of the Rocky Mountains, but rare in the Pacific Northwest.[62] In California, where it is widely
distributed, the fungus is found in a variety of plant communities, including aspen, coastal oak woodland, Douglas Fir, Klamath
mixed conifer, montane hardwood, montane hardwood-conifer, montane riparian, Redwood, Sierran mixed conifer, valley oak
woodland, valley-foothill riparian, andWhite Fir.[63] It was found to be the most common Armillaria species in hardwood and mixed
oak forests in westernMassachusetts.[64]

A Chinese study published in 2001 used the molecular biological technique restriction fragment length polymorphism to analyze the
differences in DNA sequence between 23 A. gallica specimens collected from the Northern Hemisphere. The results suggest that
based on the restriction fragment length polymorphism patterns observed, there are four global A. gallica subpopulations: the
Chinese, European, North American–Chinese, and North American–European geographical lineages.[65] A 2007 study on the
northeastern and southwestern Chinese distribution of Armillaria, using fruit body and pure culture morphology, concluded that there
are several unnamed species (Chinese biological species C, ,FH, J and L) that are similar to the commonA. gallica.[54]

Ecology
Armillaria gallica is a weaker pathogen than the related A. Mellea or A. Solidipes, and is considered a secondary parasite—typically
initiating infection only after the host's defenses have been weakened by insect defoliation, drought, or infection by another
fungus.[66] Fungal infection can lead to root rot or butt rot.[67] As the diseased trees die, the wood dries, increasing the chance of
catching fire after being struck by lightning. The resulting forest fire may in turn kill the species that killed the trees.[68] Plants that
are under water stress caused by dry soils or waterlogging are more susceptible to infection by A. gallica.[69] It has been shown to be
one of several Armillaria species responsible for widespread mortality of oak trees in the Arkansas Ozarks.[70] The fungus has also
been shown to infect Daylily in South Carolina,[71] Northern highbush blueberry (Vaccinium corymbosum) in Italy[72][73] and
vineyards (Vitis species) of Rías Baixas in northwestern Spain. The latter infestation
"may be related to the fact that the vineyards from which they were isolated were
located on cleared forestry sites".[74] When A. solidipes and A. gallica co-occur in
the same forest, infection of root systems by A. gallica may reduce damage or
prevent infection fromA. solidipes.[75]

The soil-dwelling fungal pathogen

A. gallica may be parasitized by the Trichoderma harzianum can
fungus Entoloma abortivum, resulting in parasitize A. gallica rhizomorphs.
grayish-white, malformed fruit bodies.

Armillaria gallica can develop an extensive subterranean system of

rhizomorphs, which helps it to compete with other fungi for resources or to attack trees weakened by other fungi. A field study in an
ancient broadleaved woodland in England showed that of fiveArmillaria species present in the woods,A. gallica was consistently the
first to colonize tree stumps that had been coppiced the previous year.[48] Fractal geometry has been used to model the branching
patterns of the hyphae of various Armillaria species. Compared to a strongly pathogenic species like A. solidipes, A. gallica has a
relatively sparse branching pattern that is thought to be "consistent with a foraging strategy in which acceptable food bases may be
encountered at any distance, and which favours broad and divisive distribution of potential inoculum."[24] Because the rhizomorphs
form regular networks, mathematical concepts of graph theory have been employed to describe fungal growth and interpret
ecological strategies, suggesting that the specific patterns of network attachments allow the fungus "to respond opportunistically to
spatially and temporally changing environments".[76]

Armillaria gallica may itself be parasitized by other soil flora. Several species of the fungus Trichoderma, including Trichoderma
polysporum, T. harzianum and T. viride, are able to attack and penetrate and the outer tissue of A. gallica rhizomorphs and parasitize
the internal hyphae. The infected rhizomorphs become devoid of living hyphae about one week after the initial infection.[77]
Entoloma abortivum is another fungus that can live parasitically upon A. gallica. The whitish-gray malformed fruit bodies that may
[78]
result are due to the E. abortivum hyphae penetrating the mushroom and disrupting its normal development.

See also
List of Armillaria species
List of bioluminescent fungi

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External links
Armillaria gallica in Index Fungorum

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