Armillaria gallica (synonymous with A. bulbosa and A. lutea) is a species of 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 , weigh at least , and be 1,500 years old. This individual is popularly known as the "humongous fungus", and is a tourist attraction and inspiration for an annual mushroom-themed festival in Crystal Falls. Recent studies have revised the fungus's age to 2,500 years and its size to about , four times the original estimate.
Armillaria gallica is a largely subterranean fungus, and it produces fruit bodies that are up to about in diameter, yellow-brown, and covered with small scales. On the underside of the caps are gills that are white to creamy or pale orange. The stem may be up to long, with a white cobwebby ring that divides the color of the stem into pale orange to brown above, and lighter-colored below. The fungus can develop an extensive system of underground root-like structures, called rhizomorphs, that help it to efficiently decompose dead wood in temperate broadleaf and mixed forests. It has been the subject of considerable scientific research due to its importance as a plant pathogen, its ability to bioluminesce, its unusual life cycle, and its ability to form large and long-lived colonies.
Phylogeny, taxonomy and naming
Confusion has surrounded the nomenclature and taxonomy of the species now known as Armillaria gallica, paralleling that surrounding the genus Armillaria. 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. 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. 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. 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); NABS VII was demonstrated shortly after to be the same species as EBS E. 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, as it was thought to be equivalent to Armillaria mellea var. bulbosa, first described by Jean Baptiste Barla (Joseph Barla) in 1887, and later raised to species status by Josef Velenovský in 1927. 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. Therefore, the name A. bulbosa was a misapplied name for EBS E. In 1987 Romagnesi and Marxmüller renamed EBS E to Armillaria gallica. Another synonym, A. lutea, had originally been described by Claude Casimir Gillet in 1874, and proposed as a name for EBS E. 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. A. inflata (Velenovský, 1920) may represent another synonym, but the type specimens were not preserved, so it is considered a dubious name (nomen dubium). 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.
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. These results are similar to those reported in 1992 that compared sequences of nuclear ribosomal DNA.
The specific epithet gallica is botanical Latin for "French" (from Gallia, "Gaul"), and refers to the type locality. The prior name bulbosa is Latin for "bulb-bearing, bulbous" (from bulbus and the suffix -osa). Armillaria is derived from the Latin armilla, or "bracelet".
Description
The fruit bodies of Armillaria gallica have caps that are 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 long and thick, and almost club-shaped with the base up to 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 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.
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. 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. 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.
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. One author advises to consume only a small portion initially, as some people may experience an upset stomach. The taste is described as "mild to bitter", and the odor "sweet", or reminiscent of camembert cheese.
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. 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. 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.
Metabolites Armillaria gallica can produce cyclobutane-containing metabolites such as arnamiol, a natural product that is classified as a sesquiterpenoid aryl ester. 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.
Bioluminescence The mycelia and fruiting 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. Bioluminescence is caused by the action of luciferases, enzymes that produce light by the oxidation of a luciferin (a pigment). 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, it may be a by-product of other biochemical functions, or it may help deter heterotrophs that might consume the fungus.
Humongous fungus