Exsudoporus frostii

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Exsudoporus frostii
Exsudoporus frostii 100632.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Boletales
Family: Boletaceae
Genus: Exsudoporus
Species:
E. frostii
Binomial name
Exsudoporus frostii
(J.L.Russell) Vizzini, Simonini & Gelardi (2014)
Synonyms [1]
  • Boletus frostiiJ.L.Russell (1874)
  • Suillellus frostii(J.L.Russell) Murrill (1909)
  • Tubiporus frostii(J.L.Russell) S.Imai (1968)
  • Butyriboletus frostii(J.L. Russell) G. Wu, Kuan Zhao & Zhu L. Yang (2016)
Exsudoporus frostii
Information icon.svg
Pores icon.pngPores on hymenium
Convex cap icon.svgFlat cap icon.svg Cap is convex or flat
Adnate gills icon2.svg Hymenium is adnate
Bare stipe icon.svg Stipe is bare
Transparent spore print icon.svg
Spore print is olive-brown
Mycorrhizal fungus.svgEcology is mycorrhizal
Mycomorphbox Edible.pngEdibility is edible

Exsudoporus frostii (formerly Boletus frostii), commonly known as Frost's bolete or the apple bolete, is a bolete fungus first described scientifically in 1874. A member of the family Boletaceae, the mushrooms produced by the fungus have tubes and pores instead of gills on the underside of their caps. Exsudoporus frostii is distributed in the eastern United States from Maine to Georgia, and in the southwest from Arizona extending south to Mexico and Costa Rica. A mycorrhizal species, its fruit bodies are typically found growing near hardwood trees, especially oak.

Contents

Exsudoporus frostii mushrooms can be recognized by their dark red sticky caps, the red pores, the network-like pattern of the stipe, and the bluing reaction to tissue injury. Another characteristic of young, moist fruit bodies is the amber-colored drops exuded on the pore surface. Although this mushroom is considered edible when thoroughly cooked, it is generally not recommended for consumption because of the risk of confusion with other poisonous red-pored, blue-bruising boletes. B. frostii may be distinguished from other superficially similar red-capped boletes by differences in distribution, associated tree species, bluing reaction, or morphology.

Taxonomy

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, fellow 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] 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; for this reason, the name and authority are written Boletus Frostii Russell apud Frost in some older literature. [4] Bernard Ogilvie Dodge made reference to B. frostii in 1950 during an address to the Mycological Society of America, in which he spoke about the role of the amateur 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 named a fine new bolete for his friend Russell. Boletus Frostii and Boletus Russellii 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; he concluded that it was Frost who made the original species determinations, further suggesting that "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] Tubiporus has since been synonymized with Boletus . [8] In 1945, Rolf Singer described a bolete he found in Florida; although he originally described it as a subspecies of B. frostii, [9] he later considered the differences between the taxa significant enough to warrant publishing Boletus floridanus as a unique species. [4] Following the molecular studies that outlined a new phylogenetic framework for the Boletaceae, [10] [11] the bolete was transferred to the newly circumscribed genus Exsudoporus in 2014. [12]

Due to lack of sufficient sequences, Wu and colleagues (2016) were reluctant to accept Exsudoporus and considered it a synonym of Butyriboletus , so they transferred E. frostii to the genus Butyriboletus and proposed a new combination Butyriboletus frostii. [13] However, following phylogenetic and morphological analyses clearly resolved Exsudoporus as a monophyletic, homogenous and independent genus that is sister to Butyriboletus. [14] [15]

Exsudoporus frostii is commonly known as "Frost's bolete" [16] or the "apple bolete". In Mexico, its vernacular name is panza agria, which translates to "sour belly". [17]

Description

The shape of the cap of the young fruit body ranges from a half sphere to convex, later becoming broadly convex to flat or shallowly depressed, with a diameter of 5–15 cm (2.0–5.9 in). [18] The edge of the cap is curved inward, although as it ages it can uncurl and turn upward. In moist conditions, the cap surface is sticky as a result of its cuticle, which 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). [18] Young mushrooms have a whitish bloom on the cap surface. [19]

View of the hymenium Exsudoporus frostii 15406.jpg
View of the hymenium

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. [18] 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), 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 stipe 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, though it may taper somewhat toward the top; some specimens may appear ventricose (swollen in the middle). [18] The stipe surface is mostly red, or yellowish near the base; it is reticulate—characterized by ridges arranged in the form of a net-like pattern. [18] Mycelia, visible at the base of the stipe, are yellowish white to light yellow. [4]

Young fruit bodies may secrete an amber liquid. Exsudoporus frostii 50214.jpg
Young fruit bodies may secrete an amber liquid.

The spore print of B. frostii is olive brown. [18] The spores are thick walled, smooth, and spindle shaped, with dimensions of 11–15 by 4–5  μm. Longer spores up to 18 μm long may also be present. [18] The cap cuticle, or pileipellis, is made of a tangled layer of gelatinized hyphae that are 3–6 μm wide. [20] 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 E. frostii. These hyaline (translucent) cells measure 30–53 long by 7.5–14 μm wide, and 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. [4]

Edibility and nutritional composition

This species is nonpoisonous. [21] Murrill wrote in 1910 of its edibility: "Usually viewed with suspicion because of its red hymenium, but its properties are not accurately known." [22] Since then, several authors have advised against consuming the species, due to its resemblance to other toxic red-capped boletes. [16] [18] [23] In his 100 Edible Mushrooms (2007), Michael Kuo notes that although the mushroom is apparently edible for some, it "affects others negatively". [24] David Arora mentions that it is commonly sold in rural markets in Mexico; [17] a 1997 study suggests that it is only consumed in rural areas in Querétaro state. [25] Its taste and odor have been described as "pleasant" [16] or "sweet" [21] and somewhat like citrus, [19] although the cuticle of the cap may taste acidic. [18]

Chemical analysis of fresh fruit bodies collected in Mexico showed them to have the following composition: moisture 94.53%; ash 0.323%; dietary fiber 3.024%; fat 0.368%; and protein 1.581%. The free fatty acid content of dried fruit bodies was 4.5%, slightly more than the common button mushroom (Agaricus bisporus), which had 3.5%. The majority of this total was oleic acid (1.95%), followed by linoleic acid (1.68%) and palmitic acid (1.69%). [25]

Similar species

Poisonous Boletus flammans is similar in appearance. Boletus flammans 135083.jpg
Poisonous Boletus flammans is similar in appearance.

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. [17] Often confused with Exsudoporus floridanus and E. permagnificus , but the latter species is known only from Europe and Western Asia and always grow in association with oaks and occasionally also with sweet chestnut. [26] [27] [15] Exsudoporus floridanus differs from E.  frostii in having a lighter cap color and in the texture of the cap surface: it is tomentose (covered with dense, short, soft, matted hairs) or velutinous (like velvet), compared to the relatively smooth surface of E. frostii. Singer notes that although the physical characteristics between the two taxa may be blurred and are hard to define, the area of origin can reliably distinguish them: E. floridanus is found on shaded lawns and scrubland in open oak stands in non-tropical regions of Florida, typically on grassy or sandy soil, where it fruits between May and October. [4]

The fruit bodies of young specimens of B. kermesinus , newly described from Japan in 2011, are similar in appearance to E. frostii. In addition to its distribution, B. kermesinus can be distinguished from E. frostii by having flesh that does not bruise blue and a stipe in which the reticulum is not as deep and coarse. [28] B. pseudofrostii , found in Belize, produces smaller caps that are 1.7 to 2.0 cm (0.7 to 0.8 in) in diameter. [29] Boletus russelli , found in eastern North America, has a red to reddish-brown cap and reticulate stipe, but its pore surface is yellow, and the fruit body does not bruise blue. [24]

Ecology, habitat, and distribution

Immature specimen with cap not fully expanded; bruising is evident on the stipe. Exsudoporus frostii 54090.jpg
Immature specimen with cap not fully expanded; bruising is evident on the stipe.

Exsudoporus frostii is a mycorrhizal species, [30] meaning that the fungus forms associations with the roots of various species of trees. These associations are 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 outward 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. [31] Using pure culture techniques, Exsudoporus frostii has been shown to form mycorrhizae with Virginia pine (Pinus virginiana), [30] while a field study confirms a similar association with the oak Quercus laurina . [32]

The fruit bodies grow solitarily, 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 and Michigan, and in southern Arizona. [4] [19] In Mexico, it is often found under Madrone. [17] It has also been collected in Costa Rica, where it associates with the oak species Quercus copeyensis , Q. costaricensis , Q. rapurahuensis , and Q. seemanii . [33] A 1980 publication tentatively suggested that the fungus was also present in Italy, [34] but the author later determined that the putative E. frostii was actually Boletus siculus . [27]

Fruit bodies can be parasitized by the mold-like fungus Sepedonium ampullosporum . [35] Infection results in necrosis of the mushroom tissue, and a yellow color caused by the formation of large amounts of pigmented aleurioconidia (single-celled conidia produced by extrusion from the conidiophores). [36]

See also

Related Research Articles

<span class="mw-page-title-main">Boletaceae</span> Family of fungi

The Boletaceae are a family of mushroom-forming fungi, primarily characterised by small pores on the spore-bearing hymenial surface, instead of gills as are found in most agarics. Nearly as widely distributed as the agarics, the family is renowned for hosting some prime edible species highly sought after by mushroom hunters worldwide, such as the cep or king bolete . A number of rare or threatened species are also present in the family, that have become the focus of increasing conservation concerns. As a whole, the typical members of the family are commonly known as boletes.

<i>Bothia</i> Genus of fungi

Bothia is a fungal genus in the family Boletaceae. A monotypic genus, it contains the single species Bothia castanella, a bolete mushroom first described scientifically in 1900 from collections made in New Jersey. Found in the eastern United States, Costa Rica, China, and Taiwan, it grows in a mycorrhizal association with oak trees. Its fruit body is chestnut brown, the cap is smooth and dry, and the underside of the cap has radially elongated tubes. The spore deposit is yellow-brown. The edibility of the mushroom is unknown. Historically, its unique combination of morphological features resulted in the transfer of B. castanella to six different Boletaceae genera. Molecular phylogenetic analysis, published in 2007, demonstrated that the species was genetically unique enough to warrant placement in its own genus.

<i>Butyriboletus appendiculatus</i> Species of fungus

Butyriboletus appendiculatus is an edible pored mushroom that grows under oaks and other broad leaved trees such as beech. It is commonly known as the butter bolete. It often grows in large colonies beneath the oak trees, and is frequently found cohabiting with old oaks in ancient woodland. It is relatively rare in Britain. Its stipe and pores are often bright yellow and its flesh stains bright blue when cut or bruised.

<i>Aureoboletus mirabilis</i> Species of fungus

Aureoboletus mirabilis, commonly known as the admirable bolete, the bragger's bolete, and the velvet top, is an edible species of fungus in the Boletaceae mushroom family. The fruit body has several characteristics with which it may be identified: a dark reddish-brown cap; yellow to greenish-yellow pores on the undersurface of the cap; and a reddish-brown stem with long narrow reticulations. Aureoboletus mirabilis is found in coniferous forests along the Pacific Coast of North America, and in Asia. Unusual for boletes, A. mirabilis sometimes appears to fruit on the wood or woody debris of Hemlock trees, suggesting a saprobic lifestyle. Despite the occasional appearances to the contrary, Aureoboletus mirabilis is mycorrhizal, and forms a close association with the tree's roots.

<i>Xerocomellus zelleri</i> Species of fungus

Xerocomellus zelleri, commonly known as Zeller's bolete, is an edible species of mushroom in the family Boletaceae. First described scientifically by American mycologist William Alphonso Murrill in 1912, the species has been juggled by various authors to several genera, including Boletus, Boletellus, and Xerocomus. Found solely in western North America from British Columbia south to Mexico, the fruit bodies are distinguished by their dark reddish brown to nearly black caps with uneven surfaces, the yellow pores on the underside of the caps, and the red-streaked yellow stems. The fungus grows in summer and autumn on the ground, often in Douglas fir forests or on their margins. The development of the fruit bodies is gymnocarpic, meaning that the hymenium appears and develops to maturity in an exposed state, not enclosed by any protective membrane.

<i>Suillus brevipes</i> Species of edible fungus in the family Suillaceae found throughout North America

Suillus brevipes is a species of fungus in the family Suillaceae. First described by American mycologists in the late 19th century, it is commonly known as the stubby-stalk or the short-stemmed slippery Jack. The fruit bodies (mushrooms) produced by the fungus are characterized by a chocolate to reddish-brown cap covered with a sticky layer of slime, and a short whitish stipe that has neither a partial veil nor prominent, colored glandular dots. The cap can reach a diameter of about 10 cm, while the stipe is up to 6 cm long and 2 cm thick. Like other bolete mushrooms, S. brevipes produces spores in a vertically arranged layer of spongy tubes with openings that form a layer of small yellowish pores on the underside of the cap.

<i>Suillellus amygdalinus</i> Species of fungus

Suillellus amygdalinus is a fungus of the bolete family found in western North America. The fruit bodies, or mushrooms, are characterized by their thick, red to brown caps, red pores, and the strong bluing reaction observed when the mushroom tissue is injured or cut. The cap can reach diameters of up to 12 cm (4.7 in) and the stipe 9 cm (3.5 in) long by 3 cm (1.2 in) thick at maturity. This mushroom has been found in manzanita and madrone woodlands of central California north to southern Oregon. Although the edibility of the mushroom is not known with certainty, it may be poisonous, and is not recommended for consumption. Other similar red-pored, bluing boletes from North America, including Rubroboletus eastwoodiae, Boletus luridiformis, and B. subvelutipes, can be distinguished from S. amygdalinus either by the color of the cap, the degree of reticulation on the stipe, or by location.

<i>Tylopilus tabacinus</i> Species of fungus

Tylopilus tabacinus is a species of bolete fungus in the family Boletaceae. It is characterized by a tawny-brown cap measuring up to 17.5 cm (6.9 in) in diameter, and a reticulated stem up to 16.5 cm (6.5 in) long by 6 cm (2.4 in) thick. A characteristic microscopic feature is the distinctive crystalline substance encrusted on the hyphae in the surface of the cap. The species is known from the eastern United States from Florida north to Rhode Island, and west to Mississippi, and from eastern Mexico. It is a mycorrhizal species, and associates with oak and beech trees.

<i>Exsudoporus floridanus</i> Species of fungus

Exsudoporus floridanus is a species of edible bolete mushroom in the family Boletaceae. In 1945, American mycologist Rolf Singer described a species he found in Florida during his 1942–3 tenure of a Guggenheim Memorial Fellowship. He originally described it as a subspecies of the eastern North American species Boletus frostii, but later considered it worthy of distinct species status in a 1947 publication. Based on morphological and phylogenetic data, Vizzini and colleagues transferred this species to a newly described genus Exsudoporus in 2014. Due to lack of sufficient sequences, Wu et al. (2016) were reluctant to accept Exsudoporus and considered it a synonym of Butyriboletus, so they proposed a new combination Butyriboletus floridanus. However, following phylogenetic and morphological analyses clearly resolved Exsudoporus as a monophyletic, homogenous and independent genus that is sister to Butyriboletus.

<i>Boletus curtisii</i> Species of fungus

Boletus curtisii is a species of fungus in the family Boletaceae. It produces small- to medium-sized fruit bodies (mushrooms) with a convex cap up to 9.5 cm (3.7 in) wide atop a slender stem that can reach a length of 12 cm (4.7 in). In young specimens, the cap and stem are bright golden yellow, although the color dulls to brownish when old. Both the stem and cap are slimy or sticky when young. On the underside of the cap are small circular to angular pores. The mushroom is edible, but not appealing. It is found in eastern and southern North America, where it grows in a mycorrhizal association with hardwood and conifer trees. Once classified as a species of Pulveroboletus, the yellow color of B. curtisii is a result of pigments chemically distinct from those responsible for the yellow coloring of Pulveroboletus.

<i>Boletus auripes</i> Species of fungus

Boletus auripes, commonly known as the butter-foot bolete, is a species of bolete fungus in the family Boletaceae. First described from New York in 1898, the fungus is found in eastern Asia, Central America, and eastern North America from Canada to Florida. It is a mycorrhizal species and typically grows in association with oak and beech trees.

<i>Aureoboletus auriflammeus</i> Species of fungus

Aureoboletus auriflammeus, commonly known as the flaming gold bolete, is a species of bolete fungus in the family Boletaceae. Described as new to science in 1872, it is found in eastern North America, where it grows in a mycorrhizal association with oaks. The caps of the fruit bodies are golden orange, with a yellow pore surface on the underside, and a reticulated (network-like) stem. The edibility of the mushroom is not known.

<i>Boletus miniato-olivaceus</i> Species of fungus

Boletus miniato-olivaceus is a species of bolete fungus in the family Boletaceae. Described as new to science in 1874, it is found in eastern North America and northeast Mexico.

<i>Aureoboletus projectellus</i> Species of fungus

Aureoboletus projectellus is a species of bolete fungus in the family Boletaceae. Found in North America, and recently in Europe, it grows in a mycorrhizal association with pine trees.

<i>Boletinellus merulioides</i> Species of fungus

Boletinellus merulioides, commonly known as the ash-tree bolete, is a species of bolete fungus in the family Boletinellaceae. Described as new to science in 1832, it is found in Asia and eastern North America, where it grows on the ground near ash trees.

<i>Pulveroboletus ravenelii</i> Species of fungus

Pulveroboletus ravenelii, commonly known as Ravenel's bolete or the powdery sulfur bolete, is a species of bolete fungus in the family Boletaceae. Described as new to science in 1853, the widely distributed species is known from Asia, Australia, North America, Central America, and South America. Mycorrhizal with oak, the fungus fruits on the ground singly, scattered, or in groups in woods. Fruit bodies (mushrooms) have convex to flat, yellowish to brownish-red caps up to 10 cm (4 in) in diameter. On the cap underside, the pore surface is bright yellow before turning dingy yellow to grayish brown with age; it stains greenish blue then grayish brown after injury. A cottony and powdery partial veil remains as a ring on the stipe. The mushrooms are edible, and have been used in traditional Chinese medicine and for mushroom dyeing.

<i>Porphyrellus indecisus</i> Species of fungus

Porphyrellus indecisus, commonly known as the indecisive bolete, is a species of bolete fungus in the family Boletaceae native to North America. It was described in 1888 by Charles Horton Peck.

<i>Harrya chromapes</i> Species of fungus

Harrya chromapes, commonly known as the yellowfoot bolete or the chrome-footed bolete, is a species of bolete fungus in the family Boletaceae. The bolete is found in eastern North America, Costa Rica, and eastern Asia, where it grows on the ground, in a mycorrhizal association with deciduous and coniferous trees. Fruit bodies have smooth, rose-pink caps that are initially convex before flattening out. The pores on the cap undersurface are white, aging to a pale pink as the spores mature. The thick stipe has fine pink or reddish dots (scabers), and is white to pinkish but with a bright yellow base. The mushrooms are edible but are popular with insects, and so they are often infested with maggots.

<i>Sutorius eximius</i> Species of fungus

Sutorius eximius, commonly known as the lilac-brown bolete, is a species of fungus in the family Boletaceae. This bolete produces fruit bodies that are dark purple to chocolate brown in color with a smooth cap, a finely scaly stipe, and a reddish-brown spore print. The tiny pores on the cap underside are chocolate to violet brown. It is widely distributed, having been recorded on North America, South America, and Asia, where it grows in a mycorrhizal relationship with both coniferous and deciduous trees.

<i>Imleria badia</i> Edible species of fungus in the family Boletaceae found in Europe and North America

Imleria badia, commonly known as the bay bolete, is an edible, pored mushroom found in Eurasia and North America, where it grows in coniferous or mixed woods on the ground or on decaying tree stumps, sometimes in prolific numbers. Both the common and scientific names refer to the bay- or chestnut-coloured cap, which is almost spherical in young specimens before broadening and flattening out to a diameter up to 15 cm (6 in). On the cap underside are small yellowish pores that turn dull blue-grey when bruised. The smooth, cylindrical stipe, measuring 4–9 cm long by 1–2 cm thick, is coloured like the cap, but paler. Some varieties have been described from eastern North America, differing from the main type in both macroscopic and microscopic morphology.

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