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''Caloboletus calopus'', commonly known as the bitter beech bolete or scarlet-stemmed bolete, is a
fungus A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
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 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 c ...
are attractively coloured, with a beige to olive cap 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, but ...
stains blue when broken or bruised. 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 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 species that contains the biological type specimen( ...
of '' Boletus'' and required placement in a new
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
; ''
Caloboletus ''Caloboletus'' is a fungal genus in the family Boletaceae. It was circumscribed by Italian mycologist Alfredo Vizzini with ''Caloboletus calopus'' as the type species. The erection of ''Caloboletus'' follows recent molecular studies that outlin ...
'' was erected in 2014, with ''C. calopus'' designated as the type species. Although ''Caloboletus calopus'' is not typically considered edible due to an intensely bitter taste that does not disappear with cooking, there are reports of it being consumed in eastern Europe. Its red stipe distinguishes it from edible species, such as ''
Boletus edulis ''Boletus edulis'' (English: cep, penny bun, porcino or porcini) is a basidiomycete fungus, and the type species of the genus ''Boletus''. Widely distributed in the Northern Hemisphere across Europe, Asia, and North America, it does not occu ...
''.


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, professor, botanist, mycologist, entomologist, ornithologist and inventor. Biography From 1736 to 1738 he studied Theology at the Universit ...
in 1774, but this name is unavailable for use as it was later sanctioned for another species. Johann Friedrich Gmelin's 1792
synonym A synonym is a word, morpheme, or phrase that means exactly 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 al ...
''Boletus lapidum'' is also illegitimate.
Christiaan Hendrik Persoon Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy. Early life Persoon was born in South Africa at the Cape of Good Hope, the third child of an immi ...
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: Greece 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 ...
''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 contra ...
s are scarlet-stemmed bolete and bitter beech bolete. Other synonyms include binomials resulting from generic transfers to ''Dictyopus'' by
Lucien Quélet Lucien Quélet in 1869 Lucien Quélet (14 July 1832 – 25 August 1899) was a French naturalist and mycologist. Quélet discovered several species of fungi and was the founder of the Société mycologique de France, a society devoted to myco ...
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 fr ...
in 1937. ''Boletus frustosus'', originally published as a distinct species by
Wally Snell Walter Henry "Doc" Snell (May 19, 1889 – July 23, 1980) was a pinch-hitter/catcher in Major League Baseball who played briefly for the Boston Red Sox during the season. Following this brief baseball career he became a successful mycologist ...
and Esther Dick in 1941, was later described as a variety of ''B. calopus'' by
Orson K. Miller Orson Knapp Miller Jr. (December 19, 1930 – June 9, 2006) was an American mycologist. He published numerous papers in mycology and was responsible for the naming of many taxa, as well as being one of the authors erecting the genus '' Chroogo ...
and
Roy Watling Roy Watling , PhD., DSc, FRSE, F.I.Biol., C.Biol., FLS (born 1938) is a Scottish mycologist who has made significant contributions to the study of fungi both in identification of new species and correct taxonomic placement, as well as in fungal ...
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 also refers 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 a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of the genus ''Boletus'',
Rolf Singer Rolf Singer (June 23, 1906 – January 18, 1994) was a German-born mycologist and one of the most important taxonomists of gilled mushrooms (agarics) in the 20th century. After receiving his Ph.D. at the University of Vienna in 1931 he worked ...
placed ''C. calopus'' as the
type species In 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 species that contains the biological type specimen( ...
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 sign ...
''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 published in 2013 showed that ''C. calopus'' and many (but not all) red-pored boletes were part of a ''dupainii'' clade (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'' **'' Mel ...
. 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 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 species that contains the biological type specimen( ...
of) the new genus ''
Caloboletus ''Caloboletus'' is a fungal genus in the family Boletaceae. It was circumscribed by Italian mycologist Alfredo Vizzini with ''Caloboletus calopus'' as the type species. The erection of ''Caloboletus'' follows recent molecular studies that outlin ...
'' by Italian mycologist Alfredo Vizzini.


Description

Up to 15 cm (6 in) or rarely 20 cm (8 in) in diameter, the cap 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 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, but ...
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 pinkish-tan spore print. A 3.5-centimeter ...
is olive to olive-brown.
Spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s are smooth and elliptical, measuring 13–19 by 5–6 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
. The
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
(spore-bearing cells) are club-shaped, four-spored, and measure 30–38 by 9–12 µm. The cystidia are club-shaped to spindle-shaped,
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
, 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 ''Suillellus luridus'' (formerly ''Boletus 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 a ...
'', 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 ''Xerocomellus chrysenteron'', formerly known as ''Boletus chrysenteron'' or ''Xerocomus chrysenteron'', is a small, edible, wild mushroom in the family Boletaceae. These mushrooms have tubes and pores instead of gills beneath their caps. It ...
'' 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 7–13 nm in diameter, a beta sheet (β-sheet) secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the hu ...
-
staining Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in ...
cells in the flesh of the cap, and morphologically distinct cystidia on the stipe.


Distribution and habitat

An
ectomycorrhiza An ectomycorrhiza (from Greek ἐκτός ', "outside", μύκης ', "fungus", and ῥίζα ', "root"; pl. ectomycorrhizas or ectomycorrhizae, abbreviated EcM) is a form of symbiotic relationship that occurs between a fungal symbiont, or mycob ...
l species, ''Caloboletus calopus'' grows in coniferous 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 leaves, usually in the autumn; to the shedding of petals, a ...
woodland, often at higher altitudes, especially under
beech Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engl ...
and oak. 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 (sometimes Cascadia, or simply abbreviated as 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 ...
and
Michigan Michigan () is a U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest, upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the List of U.S. states and ...
. In North America, its range extends south to Mexico. Variety ''frustosus'' is known from
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
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 straight-line distance from the northernmost part of western Canada, to New Mexico in ...
of
Idaho Idaho ( ) is a state in the Pacific Northwest region of the Western United States. To the north, it shares a small portion of the Canada–United States border with the province of British Columbia. It borders the states of Montana and W ...
. 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 other objects of natural occurrence (e.g. rocks and minerals). It is generally designed to be brought into the " field" or local area where such objects e ...
s. The bolete has been recorded from the
Black Sea The Black Sea is a marginal mediterranean sea of the Atlantic Ocean 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 bounded by Bulgaria, Georgia, ...
region in
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...
, from under '' Populus ciliata'' and '' Abies pindrow'' in
Rawalpindi Rawalpindi ( or ; Urdu, ) is a city in the Punjab province of Pakistan. It is the fourth largest city in Pakistan after Karachi, Lahore and Faisalabad, and third largest in Punjab after Lahore and Faisalabad. Rawalpindi is next to Pakistan's ...
and
Nathia Gali Nathia Gali or Nathiagali ( ur, ) 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. N ...
in Pakistan,
Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
Province in China, Korea, and Taiwan.


Biochemistry

Although it is an attractive-looking bolete, ''Caloboletus calopus'' 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. The bitter taste is largely due to the compounds calopin and a δ-
lactone Lactones are cyclic carboxylic esters, containing a 1-oxacycloalkan-2-one structure (), or analogues having unsaturation or heteroatoms replacing one or more carbon atoms of the ring. Lactones are formed by intramolecular esterification of the co ...
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 3-Methylcatechol is a chemical compound, a methylbenzenediol. Metabolism The enzyme 1,2-dihydroxy-6-methylcyclohexa-3,5-dienecarboxylate dehydrogenase uses 1,2-dihydroxy-6-methylcyclohexa-3,5-dienecarboxylate and NAD+ to produce 3-methylcate ...
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 sy ...
s. A
total synthesis Total synthesis is the complete chemical synthesis of a complex molecule, often a natural product, from simple, commercially-available precursors. It usually refers to a process not involving the aid of biological processes, which distinguishes i ...
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 Inhibitor or inhibition may refer to: In biology * Enzyme inhibitor, a substance that binds to an enzyme and decreases the enzyme's activity * Reuptake inhibitor, a substance that increases neurotransmission by blocking the reuptake of a neurotr ...
cytochrome P450 Cytochromes P450 (CYPs) are a superfamily of enzymes containing heme as a cofactor that functions as monooxygenases. In mammals, these proteins oxidize steroids, fatty acids, and xenobiotics, and are important for the clearance of various compo ...
—major
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
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. Other compounds found in the fruit bodies include calopin B, and the sesquiterpenoid compounds cyclopinol and boletunones A and B. The latter two highly oxygenated compounds have significant free-radical scavenging activity ''
in vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology and ...
''. The compounds 3-octanone (47.0% of total volatile compounds), 3-octanol (27.0%), 1-octen-3-ol (15.0%), and
limonene Limonene is a colorless liquid aliphatic hydrocarbon classified as a cyclic monoterpene, and is the major component in the oil of citrus fruit peels. The -isomer, occurring more commonly in nature as the fragrance of oranges, is a flavoring ...
(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 ...


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