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2 Description

2.1 Microscopic features

2.2 Edibility

2.3 Similar species

2.4 Metabolites

2.5 Bioluminescence

3 Humongous fungus

4 Life cycle and growth

5 Habitat and distribution

6 Ecology

7 See also

8 References

9 External links

Phylogeny, taxonomy and naming

A. solidipes

A. gemina

A. calvescens

A. gallica (ST22)

A. gallica (ST23)
A. gallica (M70)

A. sinapina

A. cepistipes


A. nabsnona
A. tabescens

A. mellea

Phylogeny and relationships of A. gallica and related North American species based on amplified
fragment length polymorphism data. SY22, ST23, and M70 are A. gallica specimens collected
from Michigan, Wisconsin, and British Columbia, respectively.[1]

Confusion has surrounded the nomenclature and taxonomy of the species now known as
Armillaria gallica, paralleling that surrounding the genus Armillaria.[2] The type species,
Armillaria mellea, was until the 1970s believed to 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 reported a technique to distinguish between Armillaria species by
growing them together as single spore isolates on petri dishes and observing changes in the
morphology of the cultures.[4] Using a similar technique, Kari Korhonen showed in 1978 that the
European Armillaria mellea species complex could be separated into five reproductively isolated
species, which he named "European Biological Species" (EBS) A through E.[5] About the same
time, the North American 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 the
same species as EBS E.[7] Because several research groups had worked with this widely
distributed species, it was assigned several different names.

The species that Korhonen called EBS B was named A. bulbosa by Helga 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 status by Josef Velenovský in 1927.[10] In 1973, the
French mycologist Henri Romagnesi, unaware of Velenovský's publication, published a
description of the species he called Armillariella bulbosa, based on specimens he had found near
Compiègne and Saint-Sauveur-le-Vicomte in France. These specimens were later demonstrated
to be the same species as the EBS E of Korhonen; EBS B was later determined to be A. cepistipes.
[11] Therefore, the name A. bulbosa was a misapplied name for EBS E. In 1987 Romagnesi and
Marxmüller renamed EBS E 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

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 nuclear ribosomal 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]


The underside of a mushroom cap showing whitish cottony tissue connecting the edge of the
brown cap with the whitish stem.

Young fruit bodies have a cottony partial veil that protects the developing gills.

The underside of a mushroom cap showing numerous closely spaced gills. A small ring of whitish
cottony tissue can be seen at the stem where it attaches the cap.

Mature gills

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.

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 to rhizomorphs, black root-
like 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


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]


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]


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

Two clusters of mushrooms growing in a bed of green moss. The mushroom caps are densely
covered with small scales and are a reddish-brown that gets deeper in the center. Some caps
appear shiny as is covered with a translucent slime. The mushroom stems are club-shaped and a
very light reddish-brown.

The fruit bodies—the visible manifestation of A. gallica—belie an extensive underground

network of mycelia.

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 restriction fragment length
polymorphism (RFLP) and random amplification 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 company U-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 fungus to adapt to unfavorable changes in environmental conditions,
like drought.[45][46][47]

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 stable genome.[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

Several clusters of light brown mushrooms growing in moss on the base of a large tree.

Young fruit bodies growing in clusters at the base of a tree

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 the Western 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. The upper
limits of its altitude vary by region. In the French Massif Central, it is 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 in organic 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, and White Fir.[63] It was found to be the
most common Armillaria species in hardwood and mixed oak forests in western Massachusetts.

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, F, H, J and L) that are similar to
the common A. gallica.[54]


An aggregation of long, thin translucent cells that are multiply branched. Some of the terminal
branches have a small circular cell at their tips.

The soil-dwelling fungal pathogen Trichoderma harzianum can parasitize A. gallica rhizomorphs.

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 from A. solidipes.[75]

Six mushrooms of various shape and either brown or whitish in color, picked and laid in a row on
a bed of moss. The two brown mushrooms have stems and caps. The smallest mushroom also
has stem and cap, but is whitish-gray. Three other whitish-gray mushrooms are irregularly
shaped and lumpy.

A. gallica may be parasitized by the fungus Entoloma abortivum, resulting in 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 five Armillaria 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 result are due to the E. abortivum hyphae
penetrating the mushroom and disrupting its normal development.[78]

See also

List of Armillaria species

List of bioluminescent fungi


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External links

Wikimedia Commons has media related to Armillaria gallica.

Armillaria gallica in Index Fungorum

Taxon identifiers

Wd: Q630885 EoL: 1031756 Fungorum: 131162 GBIF: 2536847 iNaturalist: 118401 MycoBank:
131162 NCBI: 47427

Categories: ArmillariaBioluminescent fungiFungi described in 1987Fungi of AfricaFungi of

AsiaFungi of EuropeFungi of North AmericaFungal tree pathogens and diseases

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