Caloboletus Calopus
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''Caloboletus calopus'', commonly known as the bitter bolete, bitter beech bolete or scarlet-stemmed bolete, is a species of
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the bolete family. Christiaan Persoon first described ''Boletus calopus'' in 1801. Modern
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
showed that it was only distantly related to the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of ''
Boletus ''Boletus'' is a genus of mushroom-producing fungi, comprising over 100 species. The genus ''Boletus'' was originally broadly defined and described by Carl Linnaeus in 1753, essentially containing all fungi with hymenial pores instead of gills. ...
'' and required placement in a new genus; '' Caloboletus'' was erected in 2014, with ''C. calopus'' designated as the type species. The stout
fruit bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
are attractively coloured, with a beige to olive
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. The origin of the word "cap" comes from the Old French word "chapeau" which means "head co ...
up to 15 cm (6 in) across, yellow pores, and a reddish stipe up to long and wide. The pale yellow
flesh Flesh is any aggregation of soft tissues of an organism. Various multicellular organisms have soft tissues that may be called "flesh". In mammals, including humans, ''flesh'' encompasses muscles, fats and other loose connective tissues, ...
stains blue when broken or bruised. Its red stipe distinguishes it from ''
Boletus edulis ''Boletus edulis'' (English: cep, penny bun, porcino) is a basidiomycete fungus, and the type species of the genus ''Boletus''. It is prized as an edible mushroom. The fungus produces Basidiospore, spore-bearing basidiocarp, fruit bodies ...
''. Appearing in
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
ous and deciduous woodland in summer and autumn, the species is found in Eurasia and North America. Although not typically considered
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from " eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
due to an intensely bitter taste that does not disappear with cooking, there are reports of it being consumed in eastern Europe.


Taxonomy

''Caloboletus calopus'' was originally published under the name ''Boletus olivaceus'' by
Jacob Christian Schäffer Jacob Christian Schäffer, alternatively Jakob, (31 May 1718 – 5 January 1790) was a German Dean (Christianity), dean, professor of theology, botanist, mycology, mycologist, entomology, entomologist, ornithology, ornithologist and inventor. He ...
in 1774, but this name is unavailable for use as it was later sanctioned for another species.
Johann Friedrich Gmelin Johann Friedrich Gmelin (8 August 1748 – 1 November 1804) was a German natural history, naturalist, chemist, botanist, entomologist, herpetologist, and malacologist. Education Johann Friedrich Gmelin was born as the eldest son of Philipp F ...
's 1792
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
''Boletus lapidum'' is also illegitimate.
Christiaan Hendrik Persoon Christiaan Hendrik Persoon (31 December 1761 – 16 November 1836) was a Cape Colony mycologist who is recognized as one of the founders of mycology, mycological Taxonomy (biology), taxonomy. Early life Persoon was born in Cape Colony at ...
described the mushroom in 1801; its
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
is derived from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
''καλος/kalos'' ("pretty") and ''πους/pous'' ("foot"), referring to its brightly coloured stipe. The German name, ''Schönfußröhrling'' or "pretty-foot bolete", is a literal translation. Alternate
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...
s are scarlet-stemmed bolete and bitter beech bolete. Other synonyms include binomials resulting from generic transfers to ''Dictyopus'' by
Lucien Quélet Lucien Quélet (; 14 July 1832 – 25 August 1899) was a French natural history, naturalist and mycologist. Quélet discovered several species of fungi and was the founder of the Société mycologique de France, a society devoted to mycological ...
in 1886, and ''Tubiporus'' by
René Maire René Charles Joseph Ernest Maire (; 29 May 1878, Lons-le-Saunier – 24 November 1949) was a French botanist and mycologist. His major work was the ''Flore de l'Afrique du Nord'' in 16 volumes published posthumously in 1953. He collected plants ...
in 1937. ''Boletus frustosus'', originally published as a distinct species by
Wally Snell Walter Henry "Doc" Snell (May 19, 1889 – July 23, 1980) was an American baseball player and coach, college athletics administrator, and mycologist. He played briefly as a catcher in Major League Baseball with the Boston Red Sox during the seas ...
and Esther Dick in 1941, was later described as a
variety Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
of ''B. calopus'' by Orson K. Miller and
Roy Watling Roy Watling (born 1938) is a Scottish mycologist who has made significant contributions to the study of fungi both in the identification of new species and correct taxonomic placement, as well as in fungal ecology. Biography Watling was the Hea ...
in 1968. Estadès and Lannoy described the variety ''ruforubraporus'' and the
form Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens. Form may also refer to: *Form (document), a document (printed or electronic) with spaces in which to write or enter dat ...
''ereticulatus'' from Europe in 2001. In his 1986 infrageneric
classification Classification is the activity of assigning objects to some pre-existing classes or categories. This is distinct from the task of establishing the classes themselves (for example through cluster analysis). Examples include diagnostic tests, identif ...
of the genus ''Boletus'',
Rolf Singer Rolf Singer (June 23, 1906 – January 18, 1994) was a German mycologist and Taxonomy (biology), taxonomist of gilled mushrooms (agarics). He wrote the book "The Agaricales in Modern Taxonomy (biology), Taxonomy". He fled to various countries d ...
placed ''C. calopus'' as the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the
section Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
''Calopodes'', which includes species characterised by having a whitish to yellowish flesh, bitter taste, and a blue staining reaction in the tube walls. Other species in section ''Calopodes'' include '' C. radicans'', '' C. inedulis'', '' B. peckii'', and '' B. pallidus''.
Genetic analysis Genetic analysis is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts ...
published in 2013 showed that ''C. calopus'' and many (but not all) red-pored boletes were part of a ''dupainii''
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
(named for ''Boletus'' (now ''Rubroboletus'') ''dupainii''), well-removed from the core group of the type species '' B. edulis'' and relatives within the
Boletineae The Boletineae are a suborder of the fungal order Boletales. Families in the Boletineae include the Boletaceae and the Paxillaceae. Taxa *Paxillaceae **'' Alpova'' **'' Austrogaster'' **'' Gyrodon'' **'' Hydnomerulius'' **'' Meiorganum'' **''Me ...
. This indicated it needed placement in a new genus. This took place in 2014, ''B. calopus'' was transferred to (and designated the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of) the new genus '' Caloboletus'' by Italian mycologist Alfredo Vizzini.


Description

Up to or rarely 20 cm (8 in) in diameter, the
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. The origin of the word "cap" comes from the Old French word "chapeau" which means "head co ...
is beige to olive and initially almost globular before opening out to a hemispherical and then convex shape. The surface of the cap is smooth or has minute hairs, and sometimes develops cracks with age. The
cap cuticle The pileipellis is the uppermost layer of hyphae in the pileus of a fungal fruit body. It covers the trama, the fleshy tissue of the fruit body. The pileipellis is more or less synonymous with the cuticle, but the cuticle generally describes th ...
hangs over the cap margin. The pore surface is initially pale yellow before deepening to an olive-yellow in maturity, and quickly turns blue when it is injured. The pores, numbering one or two per millimetre, are circular when young but become more angular as the mushroom ages. The tubes are up to deep. The attractively coloured stipe is typically yellow above to pink-red below, with a straw-coloured network (reticulation) near the top or over the upper half; occasionally the entire stipe is reddish. It measures long by thick, and is either fairly equal in width throughout, or thicker towards the base. Sometimes, the reddish stipe colour of mature mushrooms or harvested specimens that are a few days old disappears completely, and is replaced with ochre-brown tones. The pale yellow
flesh Flesh is any aggregation of soft tissues of an organism. Various multicellular organisms have soft tissues that may be called "flesh". In mammals, including humans, ''flesh'' encompasses muscles, fats and other loose connective tissues, ...
stains blue when broken, the discolouration spreading out from the damaged area. Its smell can be strong, and has been likened to ink. The
spore print 300px, Making a spore print of the mushroom ''Volvariella volvacea'' shown in composite: (photo lower half) mushroom cap laid on white and dark paper; (photo upper half) cap removed after 24 hours showing warm orange ("tussock") color spore print. ...
is olive to olive-brown. The
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s are smooth and elliptical, measuring 13–19 by 5–6 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
. The
basidia A basidium (: basidia) is a microscopic spore-producing structure found on the hymenophore of reproductive bodies of basidiomycete fungi. The presence of basidia is one of the main characteristic features of the group. These bodies are also ...
(spore-bearing cells) are club-shaped, four-spored, and measure 30–38 by 9–12 μm. The
cystidia A cystidium (: cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that are o ...
are club-shaped to spindle-shaped,
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
, and measure 25–40 by 10–15 μm. Variety ''frustosus'' is morphologically similar to the main type, but its cap becomes areolate (marked out into small areas by cracks and crevices) in maturity. Its spores are slightly smaller too, measuring 11–15 by 4–5.5 μm. In the European form ''ereticulatus'', the reticulations on the upper stipe are replaced with fine reddish granules, while the variety ''ruforubraporus'' has pinkish-red pores.


Similar species

The overall colouration of ''Caloboletus calopus'', with its pale cap, yellow pores and red stipe, is not shared with any other bolete. Large pale specimens resemble '' Suillellus luridus'', and the cap of '' Rubroboletus satanas'' is a similar colour but this species has red pores. Fruit bodies in poor condition could be confused with '' Xerocomellus chrysenteron'' but the stipes of this species are not reticulated. Edible species such as '' B. edulis'' lack a red stipe. It closely resembles the similarly inedible ''C. radicans'', which lacks the redness on the stipe. Like ''C. calopus'', the western North American species '' C. rubripes'' also has a bitter taste, similarly coloured cap, and yellowish pores that bruise blue, but it lacks reticulation on its reddish stipe. Found in northwestern North America, '' B. coniferarum'' lacks reddish or pinkish colouration in its yellow reticulate stipe, and has a darker, olive-grey to deep brown cap. Two eastern North American species, ''C. inedulis'' and '' C. roseipes'', also have an appearance similar to ''C. calopus''. ''C. inedulis'' produces smaller fruit bodies with a white to greyish-white cap, while ''C. roseipes'' associates solely with hemlock. '' C. firmus'', found in the eastern United States, eastern Canada, and Costa Rica, has a pallid cap colour, reddish stipe, and bitter taste, but unlike ''C. calopus'', has red pores and lacks stipe reticulation. '' C. panniformis'', a Japanese species described as new to science in 2013, bears a resemblance to ''C. calopus'', but can be distinguished by its rough cap surface, or microscopically by the
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
-
staining Staining is a technique used to enhance contrast in samples, generally at the Microscope, microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissue (biology), tissues), in cytology (microscopic ...
cells in the flesh of the cap, and morphologically distinct
cystidia A cystidium (: cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that are o ...
on the stipe.


Distribution and habitat

An
ectomycorrhiza An ectomycorrhiza (from Greek ἐκτός ', "outside", μύκης ', "fungus", and ῥίζα ', "root"; ectomycorrhizas or ectomycorrhizae, abbreviated EcM) is a form of symbiotic relationship that occurs between a fungal symbiont, or mycobio ...
l species, ''Caloboletus calopus'' grows in
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
ous and
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
woodland, often at higher altitudes, especially under
beech Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
and
oak An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
. Fruit bodies occur singly or in large groups. The species grows on chalky ground from July to December, in Northern Europe, and North America's
Pacific Northwest The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...
and
Michigan Michigan ( ) is a peninsular U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest, Upper Midwestern United States. It shares water and land boundaries with Minnesota to the northwest, Wisconsin to the west, ...
. In North America, its range extends south to Mexico. Variety ''frustosus'' is known from
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
and the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
of
Idaho Idaho ( ) is a landlocked U.S. state, state in the Pacific Northwest and Mountain states, Mountain West subregions of the Western United States. It borders Montana and Wyoming to the east, Nevada and Utah to the south, and Washington (state), ...
. In 1968, after comparing European and North American collections, Miller and Watling suggested that the typical form of ''C. calopus'' does not occur in the United States. Similar comparisons by other authors have led them to the opposite conclusion, and the species has since been included in several North American
field guide A field guide is a book designed to help the reader identify wildlife (flora or fauna or funga) or other objects of natural occurrence (e.g. rocks and minerals). It is generally designed to be brought into the " field" or local area where suc ...
s. The bolete has been recorded from the
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
region in
Turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
, from under '' Populus ciliata'' and ''
Abies pindrow ''Abies pindrow'', the pindrow fir, West Himalayan fir, or silver fir, is a fir native to the western Himalaya and adjacent mountains, from northeast Afghanistan east through northern Pakistan and India to central Nepal. Description It is a ...
'' in
Rawalpindi Rawalpindi is the List of cities in Punjab, Pakistan by population, third-largest city in the Administrative units of Pakistan, Pakistani province of Punjab, Pakistan, Punjab. It is a commercial and industrial hub, being the list of cities in P ...
and
Nathia Gali Nathia Gali or Nathiagali () is a hill station and mountain resort town located in the Abbottabad District of Khyber Pakhtunkhwa, Pakistan. It is located at the centre of the Galyat range, where several hill stations are situated. Nathia ...
in Pakistan,
Yunnan Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
Province in China, Korea, and Taiwan.


Edibility

Although an attractive-looking bolete, the species is not considered edible on account of its very bitter taste, which does not disappear upon cooking. There are reports of it being eaten in far eastern Russia and Ukraine.


Biochemistry

The bitter taste is largely due to the compounds calopin and a δ-
lactone Lactones are cyclic carboxylic esters. They are derived from the corresponding hydroxycarboxylic acids by esterification. They can be saturated or unsaturated. Lactones are formed by lactonization, the intramolecular esterification of the corresp ...
derivative, ''O''-acetylcyclocalopin A. These compounds contains a
structural motif In a chain-like biological molecule, such as a protein or nucleic acid, a structural motif is a common three-dimensional structure that appears in a variety of different, evolutionarily unrelated molecules. A structural motif does not have t ...
known as a 3-methylcatechol unit, which is rare in
natural product A natural product is a natural compound or substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural products can also be prepared by chemical s ...
s. A
total synthesis Total synthesis, a specialized area within organic chemistry, focuses on constructing complex organic compounds, especially those found in nature, using laboratory methods. It often involves synthesizing natural products from basic, commercially ...
of calopin was reported in 2003. The ''frustosus'' variety is reported as causing severe sickness in Europe. The
pulvinic acid Pulvinic acids are natural chemical pigments found in some lichens, derived biosynthetically from the aromatic amino acids phenylalanine and tyrosine, via dimerization and oxidative ring-cleavage of arylpyruvic acids, a process that also produces ...
derivatives atromentic acid, variegatic acid, and xerocomic acid are present in ''B. calopus'' mushrooms. These compounds inhibit
cytochrome P450 Cytochromes P450 (P450s or CYPs) are a Protein superfamily, superfamily of enzymes containing heme as a cofactor (biochemistry), cofactor that mostly, but not exclusively, function as monooxygenases. However, they are not omnipresent; for examp ...
—major
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s involved in
drug metabolism Drug metabolism is the metabolic breakdown of drugs by living organisms, usually through specialized enzymatic systems. More generally, xenobiotic metabolism (from the Greek xenos "stranger" and biotic "related to living beings") is the set o ...
and
bioactivation In chemistry and biology, activation is the process whereby something is prepared or excited for a subsequent reaction. Chemistry In chemistry, "activation" refers to the reversible transition of a molecule into a nearly identical chemical or ...
. Other compounds found in the fruit bodies include calopin B, and the
sesquiterpenoid Sesquiterpenes are a class of terpenes that consist of three isoprene units and often have the molecular formula C15H24. Like monoterpenes, sesquiterpenes may be cyclic or contain rings, including many combinations. Biochemical modifications such ...
compounds cyclopinol and boletunones A and B. The latter two highly oxygenated compounds have significant free-radical scavenging activity '' in vitro''. The compounds (47.0% of total volatile compounds), (27.0%),
1-octen-3-ol 1-Octen-3-ol, octenol for short and also known as mushroom alcohol, is a chemical that attracts biting insects such as mosquitoes. It is contained in human breath and sweat, and it is believed that insect repellent DEET works by blocking the i ...
(15.0%), and
limonene Limonene () is a colorless liquid aliphatic hydrocarbon classified as a cyclic monoterpene, and is the major component in the essential oil of citrus fruit peels. The (+)-isomer, occurring more commonly in nature as the fragrance of oranges, ...
(3.6%) are the predominant volatile components that give the fruit body its odour.


See also

*
List of North American boletes __NOTOC__ This is a list of bolete species found in North America. Bolding of the species name, and an asterisk (*) following indicate the species is the type species of that genus. ''Aureoboletus'' *''Aureoboletus auriporus'' *''Aureoboletus ge ...


References


External links

* * {{DEFAULTSORT:Caloboletus calopus Inedible fungi calopus Fungi of Asia Fungi of Europe Fungi of North America Fungi described in 1801 Taxa named by Christiaan Hendrik Persoon Fungus species