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Boletus frostii, commonly known as Frost's bolete or the apple bolete, is a bolete mushroom first described scientifically in 1874. A member of the Boletaceae family, the mushrooms produced by the fungus have tubes and pores instead of gills on the underside of their caps. The fruit bodies may be recognized by their dark red sticky caps, the red pores, the network-like pattern of the stem, and the bluing reaction to tissue injury. Another characteristic of young, moist fruit bodies are the amber drops exuded on the pore surface.
Boletus frostii is a mycorrhizal species, and its fruit bodies are typically found growing near hardwood trees, especially oak. B. frostii is distributed in the eastern United States from Maine to Georgia, Mexico, and Costa Rica. A subspecies, Boletus frostii ssp. floridanus, differs from the typical species in the color of the fruit body, and texture of the cap. Boletus frostii mushrooms are edible, but generally not recommended because of the risk of confusion with other poisonous red-pored boletes.
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The species was named by the Unitarian minister John Lewis Russell of Salem, Massachusetts, based on specimens found in Brattleboro, Vermont. He named the fungus after his friend, another amateur American mycologist, Charles Christopher Frost, who published a description of the species in his 1874 survey of the boletes of New England.[2][3] In some older literature, the name is given as Boletus Frostii Russell apud Frost;[4] when the name of a species is contributed by an individual, but the name is formally published by another, the contributor's name can be cited, separated from the publishing author as apud. Bernard Ogilvie Dodge made reference to B. frostii in 1950 during as address to the Mycological Society of America, in which he spoke about the role of the amateur mycologist in discovering new species:
They would have informed us all about the man Russell, who named a fine new bolete for his friend Frost, and about the man Frost, who names a fine new bolete for his friend Russell. Boletus Frostii and Boletus Russelli are mushrooms with character, even though they were described by amateurs.[5]
However, in attempting to establish a lectotype specimen, mycologist Roy Halling examined both Russell's original material and his accompanying notes, and concluded that it was Frost who made the original species determinations, and further suggested "there is no evidence to show that Russell ever collected B. frostii or wrote a description of it."[6] William Murrill in 1909 placed the species in the genus Suillellus,[7] while Sanshi Imai transferred it to Tubiporus in 1968;[1] both of these genera have since been folded into Boletus.[8] The mushroom is commonly known as "Frost's bolete",[9] or the "apple bolete". In Mexico, its vernacular name is panza agria, which translates to "sour belly".[10]
The shape of the cap of the young fruit body ranges from a half sphere to convex, later becoming broadly convex to flat to shallowly depressed, with a diameter of 5–15 cm (2.0–5.9 in).[11] The edge of the cap is curved inwards, although as it ages it can uncurl and turn upwards. Under moist conditions, the cap surface is sticky as a result of having a cuticle that is made of gelatinized hyphae. If the fruit body has dried out after a rain, the cap is especially shiny,[4] sometimes appearing finely areolate (having a pattern of block-like areas similar to cracked dried mud).[11] Young specimens have a whitish bloom on the cap surface.[12]The color is bright red initially, but fades with age. The flesh is up to 2.5 cm (1.0 in) thick, and ranges in color from pallid to pale yellow to lemon-yellow.[11] The flesh has a variable staining reaction in response to bruising, so some specimens may turn deep blue almost immediately, while others turn blue weakly and slowly.[4]
The tubes comprising the pore surface (the hymenium) are 9–15 mm deep, yellow to olivaceous yellow (mustard yellow), turning dingy blue when bruised. The pores are small (2 to 3 per mm) and circular, and until old age a deep red color that eventually becomes paler. The pore surface is often beaded with yellowish droplets when young (a distinguishing characteristic), and readily stains blue when bruised. The stem is 4 to 12 cm (1.6 to 4.7 in) long, and 1 to 2.5 cm (0.4 to 1.0 in) thick at its apex. It is roughly equal in thickness throughout its length, or it may taper somewhat toward the top, though some specimens may appear ventricose (swollen in the middle).[11] The stem surface is mostly red, or yellowish near the base; it is reticulate—characterized by ridges arranged in the form of a net-like pattern.[11] Mycelia, visible at the base of the stem, is yellowish-white to light yellow.[4]
The spore print of B. frostii is olive-brown.[11] The spores are thick-walled, smooth, and spindle-shaped, with dimensions of 11–15 by 4–5 µm. Larger spores up to 18 µm long may also be present.[11] The cap cuticle, or pileipellis, is made of a tangled layer of gelatinized hyphae that are 3–6 µm wide.[13] The spore-bearing cells, the basidia, are four-spored and measure 26–35 by 10.5–11.5 µm. Cystidia are non-fertile cells interspersed among the basidia, and they are prevalent in the hymenial tissue of B. frostii. These hyaline (translucent) cells can range in shape from somewhat like a spindle (tapering at each end, but with one end typically rounded), to subampullaceous—shaped somewhat like a swollen bottle; they are 30–53 long by 7.5–14 µm wide.[4]
In 1910, Murrill wrote of this mushroom's edibility: "Usually viewed with suspicion because of its red hymenium, but its properties are not accurately known."[14] Since then, several authors have advised against consuming the species, due to its resemblance to other toxic red-capped boletes.[11][15][9] Despite this, Boletus frostii is edible, and David Arora mentions that it is commonly sold in farmers' markets in Mexico;[10] a 1997 study suggests that it is only consumed in rural areas in Querétaro state.[16] Its taste and odor have been described as "pleasant"[9] and somewhat like citrus,[12] although the cuticle of the cap may taste acidic.[11]
Chemical analysis of fresh fruit bodies collected in Mexico showed them to have the following composition: moisture 94.53%; ash 3.23 milligrams per gram of mushroom (mg/g); dietary fiber 30.24 mg/g; fat 3.68 mg/g; and protein 15.81 mg/g. The free fatty acid content of dried fruit bodies was 45 mg/g, slightly more than the common button mushroom, which had 35 mg/g. The majority of this total was oleic acid (19.5 mg/g), linoleic acid (16.8 mg/g), and palmitic acid (16.9 mg/g).[16]
In 1945, Rolf Singer reported a subspecies he found in Florida during his 1942–3 tenure of a Guggenheim Memorial Fellowship.[17] Boletus frostii ssp. floridanus differs from the typical form in the lighter color of the cap[18] and in texture of the cap surface: the subspecies is tomentose (covered with dense, short, soft, matted hairs) or velutinous (like velvet), compared to the relatively smooth surface of the typical. Singer notes that although the physical characteristics between the two taxa may be blurred and are hard to define, the area of origin is a reliable indicator of subspecies status. Subspecies floridanus is found on shaded lawns and scrubland in open oak stands in non-tropical regions of Florida, typically on grassy or sandy soil. It grows under or near several oak species, including chapman oak (Quercus chapmanii), swamp laurel oak (Q. laurifolia), and southern live oak (Q. virginiana), and it fruits between May and October.[4]
Other red-capped boletes include the poisonous B. flammans and B. rubroflammeus; the former grows most commonly under conifers, the latter in association with hardwoods in eastern North America and southern Arizona.[10] Often confused with B. frostii is B. permagnificus, but the latter species is known only known from France and Italy, and always grows in association with oaks.[19] The fruit bodies of young specimens of B. kermesinus, newly described from Japan in 2011, are similar in appearance to B. frostii. In addition to its distribution, B. kermesinus can be distinguished from B. frostii by having flesh that does not bruise blue, and a stem in which the reticulum is not as deep and coarse.[20] B. pseudofrostii, found in Belize, produces smaller caps that are 1.7 to 2.0 cm (0.7 to 0.8 in) in diameter.[21]
Boletus frostii is a mycorrizhal species,[22] meaning that the fungus forms associations with the roots of various species of trees. This association is mutualistic, because the fungus absorbs mineral nutrients from the soil and channels these into the plant, while the plant provides the fungus with sugars, a product of photosynthesis. The characteristic feature of the mycorrhiza is the presence of a sheath of fungal tissue that encases the terminal, nutrient-absorbing rootlets of the host plant. The fungus forms an extensive underground network of hyphae that radiate outwards from the surface of the root sheath, effectively increasing the surface area for nutrient absorption. The hyphae also invade between the root cortical cells to form a Hartig net.[23] Using pure culture techniques, Boletus frostii has been shown to form mycorrhizae with Virginia pine (Pinus virginiana,[22] while a field study confirms the same association with the oak Quercus laurina.[24]
The fruit bodies grow solitary, scattered, or in groups on the ground under hardwood trees; the fungus fruits in summer to early autumn. William Murrill noted its preference for growing in "thin oak woods, where the light is sufficient to enable grass to grow",[7] and Alexander H. Smith mentioned its preference for growing in "thin, sandy soil under scrub oak."[3] In the United States, it is distributed from Maine south to Georgia, extending west to Tennessee, Michigan, and southern Arizona.[4][12] In Mexico, it is often found under Madrone.[10] It has also been collected in Costa Rica, where it associates with the oak species Quercus copeyensis, Q. costaricensis, Q. rapurahuensis, and Q. seemanii.[25] A 1980 publication tentatively suggested that the fungus was also present in Italy,[26] but the author later determined that the putative B. frostii was actually Boletus siculus.[27]
Fruit bodies can be parasitized by the mold-like fungus Sepedonium ampullosporum.[28] Infection results in necrosis of the mushroom tissue, and a yellow color caused by the formation of large amounts of pigmented aleurioconidia (single cell conidia produced by extrusion from the conidiophores).[29]