Imperator torosus

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Imperator torosus
Boletus torosus 1.JPG
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Boletales
Family: Boletaceae
Genus: Imperator
Species:
I. torosus
Binomial name
Imperator torosus
(Fr.) Assyov, Bellanger, Bertéa, Courtec., Koller, Loizides, G.Marques, J.A.Muñoz, N.Oppicelli, D.Puddu, F.Rich. & P.-A.Moreau (2015)
Synonyms [1] [2]
  • Boletus torosusFr. (1835)
  • Dictyopus torosus(Fr.) Quél. (1886)
  • Boletus purpureus var. xanthocyaneus Romagn. (1948)
  • Boletus xanthocyaneus(Ramain) Romagn. (1976)
Imperator torosus
Information icon.svg
Pores icon.pngPores on hymenium
Convex cap icon.svgFlat cap icon.svg Cap is convex or flat
Bare stipe icon.svg Stipe is bare
Transparent spore print icon.svg
 Spore print is olive-brown
Mycorrhizal fungus.svgEcology is mycorrhizal
Mycomorphbox Poison.pngEdibility is poisonous

Imperator torosus, commonly known as the brawny bolete, is a species of bolete fungus in the family Boletaceae. It is native to southern Europe east to the Caucasus and Israel. It is generally associated with deciduous trees such as hornbeam, oak and beech in warm, dry locales. Although generally rare in Europe, it appears to be relatively common in Hungary. Appearing in summer and autumn on chalky soils, the stocky fruit bodies have an ochre cap up to 20 cm (8 in) across, yellow pores on the cap underside, and a wine-red to brown or blackish stipe up to 6–15 cm (2.4–5.9 in) long by 3–6 cm (1.2–2.4 in) wide. The pale yellow flesh changes to different colours when broken or bruised depending on age; younger mushrooms become reddish, and older ones additionally take on bluish tones.

Contents

Elias Magnus Fries and Christopher Theodor Hök first described this species as Boletus torosus in 1835, a name by which it came to be known for many years. Modern molecular phylogenetics shows that it is only distantly related to Boletus edulis —the type species of Boletus —and it was duly placed in the new genus Imperator in 2015. Eating raw mushrooms of this species leads to gastroenteritis. Illness has also occurred after eating cooked specimens, though this does not always happen.

Taxonomy

Swiss mycologist Louis Secretan described the brawny bolete as Boletus pachypus in his 1833 work Mycographie Suisse. [3] Many of his names have been rejected for nomenclatural purposes because Secretan had a narrow species concept, dividing many taxa into multiple species that were not supported by other authorities, and his works did not use binomial nomenclature consistently. [4] [5] Swedish mycologists Elias Magnus Fries and Christopher Theodor Hök described Boletus torosus in 1835 based on Secretan's B. pachypus—distinct from the B. pachypus described by Fries himself. [6] [lower-alpha 1] Fries reported in his 1838 book Epicrisis Systematis Mycologici seu Synopsis Hymenomycetum that he had not actually observed the species, [7] and he did not designate a type specimen or illustration. [lower-alpha 2] [8] The specific epithet torosus, which derives from Latin, means "muscular". [9] In the United Kingdom, it is known commonly as the "brawny bolete". [10] The German name Ochsen-Röhrling means "oxen bolete" [11] and the French bolet vigoureux is "strong bolete". [12]

French naturalist Lucien Quélet transferred the species to the now-obsolete genus Dictyopus in 1886, which resulted in the synonym Dictyopus torosus. [13] Boletus xanthocyaneus, first described by Henri Romagnesi in 1948 as Boletus purpureus var. xanthocyaneus and classified as a species in 1976, [14] was considered by Italian mycologist Carlo Luciano Alessio to be synonymous with B. torosus. [1] Others, however, regard this fungus a distinct species. [15] [16] [17] In 2013, Italian mycologists Valerio Bertolini and Giampaolo Simonini observed that the brevity of the original species description meant that some subsequent papers used B. torosus for specimens that aligned more closely with descriptions of Boletus rhodopurpureus or B. luteocupreus (both now placed in the genus Imperator). They did note that Swiss and French authors had adhered to a more detailed description by Quélet, and that this was the only description faithful to the original description with a grey cap and yellow pores that slowly turn red. They concluded the original description must have been based on specimens growing near the French-Swiss border and hence selected a neotype specimen from this region. [8]

Within the large genus Boletus, the brawny bolete was classified in the section Luridi, [18] which included species producing medium to large fruit bodies with thick, swollen stipes, and minute pores. [19] In 1996, Czech mycologist Jiří Hlaváček further subdivided the section Luridi, defining and naming the subsection Torosi—for B. torosus—to contain boletes that strongly bruised blue-black with handling. [20]

In a molecular analysis of Boletaceae phylogeny, the brawny bolete was most closely related to Boletus luteocupreus; these two species formed a clade that was sister to B. luridus . [21] Genetic analysis published in 2013 showed that B. torosus and many (but not all) red-pored boletes were part of a dupainii clade (named for B. dupainii ), well-removed from the core group of Boletus edulis (the type species of genus Boletus) and relatives within the Boletineae. This indicated that the brawny bolete and its relatives needed to be placed in a new genus. [22] It was made the type species of the new genus Imperator , [lower-alpha 3] becoming Imperator torosus, in 2015. [23]

Description

The base of the thick stipe is wine-red coloured. Boletus torosus 2.JPG
The base of the thick stipe is wine-red coloured.

Quite heavy compared with other mushrooms of similar size, the stocky fruit body contains relatively high amounts of chitin. Holding the brawny bolete has been described as "having a stone in your hand". [24] The cap is 6–20 cm (2.4–7.9 in) in diameter. Hemispherical when young, it extends and flattens into a convex and finally a flattish shape, sometimes forming a slight depression in the centre. Initially curled downward and inward, the cap margin gradually turns outward, eventually flattening in maturity, and sometimes protruding beyond the pored undersurface. The surface texture starts slightly velvety, but becomes smoother and hairless with age; in moist weather, the cap is slightly sticky. Changing over time, the colour of the cap is initially vivid yellow or golden yellow, then ochre, then darkening in age to brown. However, if the cap cuticle is bruised, or even touched, there are sudden changes in the colour that depend on the age of the mushroom: young, yellow caps become cherry-red, while ochre-brown cap tissue bruises to reddish-purple. These colour changes soon give way to bluish or greyish tones, which, when combined, give the cap a variegated appearance. [1]

The pores on the cap underside are round and small, [25] the sulfur-yellow colour of the pore surface becoming less intense with age. The squat stipe measures 6–15 cm (2.4–5.9 in) long by 3–6 cm (1.2–2.4 in) thick. Egg-shaped when young, it lengthens somewhat as the mushroom grows, but is still bulbous in maturity; a mature stipe is typically a little shorter than the diameter of the cap. It is initially roughly the same colour as the cap, but as it matures, develops a wine-red colouration near the base and a dirty brown to bluish-black colouration elsewhere. The surface has a mesh-like pattern (reticulation) that has a colour development similar to that of the cap: initially yellow, then purplish, and finally dark brown. The flesh is thick and hard. It is yellow, except for the stipe base where it is deep red or dark brownish in older mushrooms. The pores, stipe and flesh turn greenish-blue with bruising or cutting. [1] The smell is unpleasant, but the taste is mild. [25]

The spore print is olive-brown. The smooth oval spores measure 12–15 by 5–6  μm, and sometimes have an internal oil drop. Basidia (spore-bearing cells) are club-shaped, four-spored, and measure 22–34 by 8–9 μm. The fusiform (fuse-shaped) cystidia are colourless with oil droplets, and have dimensions of 40–50 by 7–8 μm. [1] Cap cuticle tissue is in the form of a trichoderm, where the outermost hyphae emerge roughly parallel, like hairs, perpendicular to the cap surface. These cylindrical, interwoven hyphae contain septa. [25]

Similar species

The Mediterranean species Boletus poikilochromus somewhat resembles I. torosus, but can usually be distinguished by a smaller fruit body, a cylindrical stipe, and the lighter colours of younger mushrooms. Also, B. poikilochromus does not feature the colour change of the stipe base in mature fruit bodies that is seen in I. torosus. [26] Imperator luteocupreus and Caloboletus radicans are also similar but the pores of the former are red, and the flesh of the latter has a bitter taste. [25]

Several chemical tests can be used to distinguish I. torosus from other similar boletes, such as I. rhodopurpureus. I. torosus displays the following characteristic colour changes with tests performed on cut flesh less than 12 hours old: ammonium hydroxide (NH4OH)–yellow centre bordered by a blue circular outer ring; potassium hydroxide (KOH)–deep buff (instantly); ferrous sulfate (FeS04)–no change; Melzer's reagent–dark blue; phenol–blue green (very slow); formalin–mid blue (slow). [16]

Toxicity

Like many boletes, Imperator torosus causes gastrointestinal symptoms such as abdominal pain, diarrhoea, and vomiting when eaten raw. These reactions can also happen to some people when the mushroom is eaten cooked, though others have consumed it with no ill effects. [24] In a 1994 study, researchers Ulrich Kiwitt and Hartmut Laatsch looked for the antabuse-like compound coprine in Suillellus luridus and similar species that had been suspected of inducing Antabuse-like reactions with alcohol. [27] Coprine ingestion results in heat and flushing in the face, tingling in arms and legs, nausea and vomiting, and increased heart rate within five to ten minutes of consuming alcohol. [28] They found none in the suspect species, but did find indications for it in Imperator torosus. They concluded that the most likely explanation for historical poisoning incidents was a misidentification of I. torosus with Suillellus luridus, though they could not rule out the latter species containing a hitherto unidentified compound causing alcohol-related reactions. [27] No clinical cases of alcohol-related sensitivity have been recorded for I. torosus. [24]

Distribution and habitat

Imperator torosus is native to southern Europe east to the Caucasus [25] and Israel. [18] A mycorrhizal species, the fungus is generally associated with deciduous trees such as hornbeam (Carpinus), oak (Quercus) and beech (Fagus) in warm, dry locales, and is generally rare in Europe. Russian mycologist Anna Kiyashko has proposed it be listed on the IUCN Red List due to ongoing threats to its habitat. [29] In the United Kingdom, it is listed as a "priority species"—a threatened species requiring conservation action under the UK Biodiversity Action Plan. [30] Similarly, it is on the list of protected species of macrofungi in Montenegro, [31] and Italy, where it has a highly restricted range and is considered threatened by changed forest fire intervals and habitat degradation. [32] It appears to be relatively common in Hungary. [33] It is found under Quercus coccifera trees in the Bar'am forest of Upper Galilee. [18] Fruitings typically occur from July to September in calcareous (chalky) soil, with fruit bodies appearing either singly or in small groups. [1]

Notes

  1. ↑ Fries' Boletus pachypus is a synonym of Caloboletus radicans .
  2. ↑ Type specimens were not necessary for a valid species description at this time.
  3. ↑ Boletus luteocupreus and B. rhodopurpureus were also placed in this new genus. [23]

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>Rubroboletus satanas</i> Species of fungus

Rubroboletus satanas, commonly known as Satan's bolete or the Devil's bolete, is a basidiomycete fungus of the bolete family (Boletaceae) and one of its most infamous members. It was known as Boletus satanas before its transfer to the new genus Rubroboletus in 2014, based on molecular phylogenetic data. Found in broad-leaved and mixed woodland in the warmer regions of Europe, it is classified as a poisonous mushroom, known to cause violent gastroenteritis. However, reports of poisoning are rare, due to the striking coloration and unpleasant odor of the fruiting bodies, which discourage experimentation.

<i>Caloboletus calopus</i> Species of fungus in the family Boletaceae found in Asia, Northern Europe and North America

Caloboletus calopus, commonly known as the bitter bolete, bitter beech bolete or scarlet-stemmed bolete, is a fungus of the bolete family, found in Asia, Northern Europe and North America. Appearing in coniferous and deciduous woodland in summer and autumn, the stout fruit bodies are attractively coloured, with a beige to olive cap up to 15 cm (6 in) across, yellow pores, and a reddish stipe up to 15 cm (6 in) long and 5 cm (2 in) wide. The pale yellow flesh stains blue when broken or bruised.

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

Tylopilus felleus, commonly known as the bitter bolete or the bitter tylopilus, is a fungus of the bolete family. Its distribution includes east Asia, Europe and eastern North America, extending south into Mexico and Central America. A mycorrhizal species, it grows in deciduous and coniferous woodland, often fruiting under beech and oak. Its fruit bodies have convex to flat caps that are some shade of brown, buff or tan and typically measure up to 15 cm (6 in) in diameter. The pore surface is initially white before turning pinkish with age. Like most boletes it lacks a ring and it may be distinguished from Boletus edulis and other similar species by its unusual pink pores and the prominent dark-brown net-like pattern on its stalk.

<i>Neoboletus luridiformis</i> Species of fungus

Neoboletus luridiformis, also previously known as Boletus luridiformis and (invalidly) as Boletus erythropus, is a fungus of the bolete family, all of which produce mushrooms with tubes and pores beneath their caps. It is found in Northern Europe and North America, and is commonly known as the scarletina bolete, for its red pores, which are yellow when young. Other common names include the red foot bolete, dotted stemmed bolete, or dotted stem bolete.

<i>Suillellus luridus</i> Species of edible fungus of the bolete family, found in Asia, Europe, and eastern North America

Suillellus luridus, commonly known as the lurid bolete, is a fungus of the family Boletaceae, found in calcareous broadleaved woodlands in Europe. Fruit bodies appear in summer and autumn and may be locally abundant. It is a firm bolete with an olive-brown cap up to 20 cm (8 in) in diameter, with small orange or red pores on the underside. The stout ochre stem reaches 8–14 cm (3–6 in) high and 1–3 cm (0.4–1.2 in) wide, and is patterned with a red network. Like several other red-pored boletes, it stains blue when bruised or cut.

<i>Rubroboletus legaliae</i> Species of fungus

Rubroboletus legaliae, previously known as Boletus splendidus, B. satanoides, and B. legaliae is a basidiomycete fungus of the family Boletaceae. It is poisonous, with predominantly gastrointestinal symptoms, and is related to Rubroboletus satanas.

<i>Gyrodon lividus</i> Species of fungus

Gyrodon lividus, commonly known as the alder bolete, is a pored mushroom bearing close affinity to the genus Paxillus. Although found predominantly in Europe, where it grows in a mycorrhizal association with alder, it has also recorded from China, Japan and California. Fruit bodies are distinguished from other boletes by decurrent bright yellow pores that turn blue-grey on bruising. G. lividus mushrooms are edible.

<i>Xerocomus subtomentosus</i> Species of fungus

Xerocomus subtomentosus, commonly known as suede bolete, brown and yellow bolete , boring brown bolete or yellow-cracked bolete, is a species of bolete fungus in the family Boletaceae. The fungus was initially described by Carl Linnaeus in 1753 and known for many years as Boletus subtomentosus. It is edible, though not as highly regarded as other bolete mushrooms.

<i>Exsudoporus frostii</i> Species of fungus in the family Boletaceae found in North America

Exsudoporus 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.

<i>Rubroboletus rhodoxanthus</i> Species of fungus

Rubroboletus rhodoxanthus is a species of bolete in the family Boletaceae, native to Europe. Previously known as Boletus rhodoxanthus, it was transferred in 2014 to the newly erected genus Rubroboletus, based on DNA data.

<i>Imperator rhodopurpureus</i> Species of fungus

The oldrose bolete, Imperator rhodopurpureus, is an inedible fungus of the genus Imperator, found under deciduous trees including oak and beech in neutral soils. Initially described as Boletus rhodopurpureus, it was transferred to the new genus Imperator in 2015. The bolete is considered critically endangered in the Czech Republic., and endangered in the United Kingdom, reported most commonly from Berkshire and Hampshire, and typically very rare throughout, although up to a hundred fruiting bodies have been recorded at a few sites.

Zangia is a genus of bolete fungi in the family Boletaceae. The genus, circumscribed in 2011, contains six species found in China. Zangia species grow in forests dominated by Fagaceae mixed with Pinaceae (pines).

<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>Xerocomellus</i> Genus of fungi

Xerocomellus is a genus of fungi in the family Boletaceae. The genus, as it was described in 2008, contained 12 species. However X. rubellus and X. engelii were transferred to the new genus Hortiboletus and X. armeniacus was transferred to the new genus Rheubarbariboletus in 2015. Molecular analysis supports the distinction of Xerocomellus species from Boletus and Xerocomus, within which these species were formerly contained. Xerocomellus in fact is only distantly related to Xerocomus and is most closely related to Tylopilus, Boletus sensu stricto, Porphyrellus, Strobilomyces, and Xanthoconium.

<i>Imperator luteocupreus</i> Species of fungus

Imperator luteocupreus is a species of bolete fungus in the family Boletaceae. It is native to southern Europe, where it is found under chestnut (Castanea) and oak (Quercus). Although it was originally described in genus Boletus, it was placed in the new genus Imperator in 2015, based on phylogenetic inferences.

<i>Butyriboletus</i> Genus of fungi

Butyriboletus is a genus of fungi in the family Boletaceae. The genus was circumscribed in 2014 by mycologists David Arora and Jonathan L. Frank to accommodate "butter bolete" species that were shown by molecular analysis to be phylogenetically distinct from Boletus. Butyriboletus contains 24 ectomycorrhizal species found in Asia, Europe, North America and north Africa.

<i>Hemileccinum impolitum</i> Species of fungus

Hemileccinum impolitum is a basidiomycete fungus of the family Boletaceae, native to Europe. It is commonly referred to as the iodine bolete, because its fruit bodies tend to emit an iodine-like odour when cut, more detectable in the stem base or overripe specimens.

<i>Chalciporus rubinellus</i> Species of fungus

Chalciporus rubinellus, commonly known as the purple-red bolete, is a bolete fungus of the family Boletaceae. It was first described in 1879 as Boletus rubinellus, and later transferred to the genus Chalciporus in 1973.

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

Aureoboletus betula is a species of mushroom producing fungus in the family Boletaceae. It is commonly known as the Shaggy Stalked Bolete.

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  30. ↑ Joint Nature Conservation Committee (15 December 2010). UK priority species pages – Version 2. Boletus torosus Fr. & Hök (PDF) (Report).
  31. ↑ Kasom G, Miličković N (2010). "Protected species of macrofungi in Montenegro" (PDF). Natura Montenegrina, Podgorica. 9 (2): 195–203. Archived from the original (PDF) on 2016-03-04.
  32. ↑ Wagensommer, Robert P.; Angeles Flores, Giancarlo; Arcangeli, Andrea; Bistocchi, Giancarlo; Maneli, Fabio; Materozzi, Giorgio; Perini, Claudia; Venanzoni, Roberto; Angelini, Paola (2021). "Application of IUCN red listing criteria at the regional level: A case study with Boletales across the Apennine province ecoregion and EU-habitats of Central Italy". Plant Biosystems. 156 (3): 1–11. doi:10.1080/11263504.2021.1918782. S2CID   234834743.
  33. ↑ Benedek L, Pál-Fám F (2006). "Rare macrofungi from Central Börzsöny I. Hungarian occurrence data and habitat preference". International Journal of Horticultural Science. 12 (1): 45–52. doi: 10.31421/IJHS/12/1/622 . hdl: 2437/314359 .
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