Boletaceae
Encyclopedia
Boletaceae are a family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 of mushrooms, primarily characterized by developing their spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s in small pores on the underside of the mushroom, instead of gills, as are found in agaric
Agaric
An agaric is a type of fungal fruiting body characterized by the presence of a pileus that is clearly differentiated from the stipe , with lamellae on the underside of the pileus. "Agaric" can also refer to a basidiomycete species characterized by an agaric-type fruiting body...

s. Nearly as widely distributed as agarics, they include the Cep or King Bolete (Boletus edulis)
Boletus edulis
Boletus edulis, commonly known as penny bun, porcino or cep, 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 occur naturally in the Southern Hemisphere, although it has been...

, much sought after by mushroom hunters
Mushroom hunting
Mushroom hunting, mushrooming, mushroom picking, and similar terms describe the activity of gathering mushrooms in the wild, typically for eating...

. As a whole, the typical members of the family are commonly known as boletes.

Boletes are a relatively safe group of mushrooms for human consumption, as none are known to be deadly to adults, and they are the most sought after fungi for mushroom hunting
Mushroom hunting
Mushroom hunting, mushrooming, mushroom picking, and similar terms describe the activity of gathering mushrooms in the wild, typically for eating...

. They are especially suitable for novice mushroom hunters, since there is little danger of confusing them with deadly mushrooms, like various Amanita
Amanita
The genus Amanita contains about 600 species of agarics including some of the most toxic known mushrooms found worldwide. This genus is responsible for approximately 95% of the fatalities resulting from mushroom poisoning, with the death cap accounting for about 50% on its own...

agarics, which are the most poisonous mushrooms in the world. They are easily distinguished from agaric
Agaric
An agaric is a type of fungal fruiting body characterized by the presence of a pileus that is clearly differentiated from the stipe , with lamellae on the underside of the pileus. "Agaric" can also refer to a basidiomycete species characterized by an agaric-type fruiting body...

s, and easily recognized for colour, pores and thick stems
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...

 and caps
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...

.

Description

Most species in Boletaceae produce large fleshy mushrooms with a central stipe. The spore print
Spore print
thumb|300px|right|Making a spore print of the mushroom Volvariella volvacea shown in composite: mushroom cap laid on white and dark paper; cap removed after 24 hours showing pinkish-tan spore print...

 colours are commonly olivaceous (yellowish-green), yellowish, brownish, or vinaceous (red-wine coloured). In many species, flesh
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....

 that is bruised or cut will turn blue, a result of the oxidation of pulvinic acid derivatives, like variegatic, xerocomic, and atrotomentinic acid. The mushrooms usually have tubular hymenophore
Hymenophore
A hymenophore refers to the hymenium-bearing structure of a fungal fruiting body. Hymenophores can be smooth surfaces, lamellae, folds, tubes, or teeth....

s, although some species (like those in the genus Phylloporus
Phylloporus
Phylloporus is a genus of fungi in the Boletaceae family . The genus has a cosmopolitan distribution, and contains about 50 species, mostly in tropical areas.-Species:*Phylloporus albocarnosus Heinem....

) are lamellate.

Taxonomy

Boletaceae were first described by the French botanist François Fulgis Chevallier in 1826 as a family distinct from Agaricaceae
Agaricaceae
The Agaricaceae are a family of basidiomycete fungi and includes the genus Agaricus, as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. The genus contains 85 genera and 1340 species.-Genera:...

. Five genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...

 were included in Chevallier's circumscription: Boletus
Boletus
Boletus is a genus of mushroom, comprising over 100 species. The genus Boletus was originally broadly defined and described by Elias Magnus Fries in 1821, essentially containing all fungi with pores...

, Cladosporus (now synonymous with Laetiporus), Physisporus (now Perenniporia
Perenniporia
Perenniporia is a cosmopolitan genus of bracket-forming or encrusting polypores containing about 60 currently recognized species. They are dimitic or trimitic with smooth, thick-walled basidiospores and cause a white rot in affected wood.-Species:...

), Polyporus
Polyporus
Polyporus is a genus of fungi in the Polyporaceae family. It is a genus used for the production of single cell proteins, sources of mixed protein extracted from pure or mixed cultures of algae, yeasts, fungi or bacteria used as a substitute for protein-rich foods, in human and animal feeds.The...

, and Fistulina
Fistulina
Fistulina is a genus of fungi in the Fistulinaceae family. Species in the genus cause a brown rot of both dead and living hardwood trees....

.

Genera

Rolf Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

, in the 4th edition (1986) of his Agaricales in Modern Taxonomy, included 26 genera and 415 species in the Boletaceae. Molecular phylogenetic studies of the 2000s have revised our concept of the family; in a highly-cited 2006 publication, Manfred Binder and David Hibbett included 38 genera. Even after recent changes in classification that have moved many members out of the Boletaceae, it remains a large family with many genera. According to the Dictionary of the Fungi (10th edition, 2008), 35 genera are recognized in Boletaceae, which collectively contain 787 species.
Genus Authority  Year # of species Distribution
Afroboletus
Afroboletus
Afroboletus is a genus of fungi in the family Boletaceae. Afroboletus luteolus is known to be found in Togo.-Species:* Afroboletus azureotinctus* Afroboletus costatisporus* Afroboletus elegans* Afroboletus lepidellus...

Pegler & T.W.K.Young 1981 7 tropical Africa
Aureoboletus
Aureoboletus
Aureoboletus is a genus of bolete fungi in the family Boletaceae. It was circumscribed by Czech mycologist Zdeněk Pouzar in 1957. A taxonomic monograph was recently published in 2010.-Species:*A. auriporus*A. citriniporus*A. flavimarginatus...

Pouzar 1957 5 widespread
Austroboletus
Austroboletus
Austroboletus is a genus of fungi in the family Boletaceae. Formerly a subgenus of Boletus, Austroboletus was raised to genus level in 1979. Although they resemble the boletes macroscopically, Austroboletus is differentiated microscopically with spores that are pitted, rather than...

Wolfe 1980 America; Australia
Boletellus
Boletellus
Boletellus is a genus of fungi in the family Boletaceae. The genus has a widespread distribution, especially in subtropical regions, and contains about 50 species. The genus was first described by American mycologist William Alphonso Murrill in 1909...

Murrill 1909 widespread
Boletochaete
Boletochaete
Boletochaete is a genus of fungi in the family Boletaceae. The genus contains three species found in Africa and southeast Asia. American mycologist Rolf Singer circumscribed the genus in 1944....

Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

1944 3 Africa; Southeast Asia
Boletus
Boletus
Boletus is a genus of mushroom, comprising over 100 species. The genus Boletus was originally broadly defined and described by Elias Magnus Fries in 1821, essentially containing all fungi with pores...

Fr. 1821 widespread
Bothia
Bothia
Bothia is a genus of fungi in the Boletaceae family. A monotypic genus, it contains the single species Bothia castanella, found in North America...

Halling, T.J.Baroni, & Binder 2007 1 North America
Buchwaldoboletus
Buchwaldoboletus
Buchwaldoboletus is a genus of bolete fungi in the family Boletaceae. It was circumscribed by mycologist Albert Pilát in 1969....

Pilát 1962 3 Europe; Australia
Chalciporus
Chalciporus
Chalciporus is a genus of fungi in the family Boletaceae . There are approximately 25 species in the genus.-Species:*Chalciporus africanus*Chalciporus aurantiacus*Chalciporus cervinococcineus*Chalciporus chontae...

Bataille 1908 25 widespread
Chamonixia
Chamonixia
Chamonixia is a genus of truffle-like fungi in the Boletaceae family. The genus is widely distributed, especially in temperate regions, and contains eight species. Chamonixia was circumscribed by French mycologist Léon Louis Rolland in 1899....

Rolland 1899 8 widespread
Fistulinella
Fistulinella
Fistulinella is a genus of bolete fungi in the family Boletaceae. The genus has a pantropical distribution, and contains 15 species. Fistulinella was circumscribed by German mycologist Paul Christoph Hennings in 1901.-Species:...

Henn. 1901 15 pantropical
Pantropical
In biogeography, a pantropical distribution one which covers tropical regions of all of the major continents, i.e. in Africa, in Asia and in the Americas. Examples include the plant genera Acacia and Bacopa....

Gastroboletus
Gastroboletus
Gastroboletus is a genus of fungi in the family Boletaceae. Species in the genus have misshapen caps, poorly developed or absent stipes, and are often buried or partially buried. Gastroboletus has tubes arranged irregularly, rather than vertically as in typical boletes...

Lohwag 1962 13 widespread
Gastroleccinum
Gastroleccinum
Gastroleccinum is a genus of fungi in the Boletaceae family. It is a monotypic genus, represented by the single species Gastroleccinum scabrosum.-External links:*...

Thiers 1989 1 North America
Heimioporus
Heimioporus
Heimioporus is a genus of fungi in the Boletaceae family. The genus is widely distributed in tropical and subtropical regions, and contains about 15 species.-Species:*Heimioporus alveolatus*Heimioporus anguiformis*Heimioporus betula...

E.Horak 2004 widespread
Heliogaster
Heliogaster
Heliogaster is a sequestrate genus of fungi in the Boletaceae family. This is a monotypic genus, containing the single species Heliogaster columellifer, found in Japan. The genus was first described in 2010, to accommodate the species formerly named Octaviania columellifera by Kobayasi in 1937....

(Kobayasi) Orihara & Iwase 2010 1 Japan
Leccinellum
Leccinellum
Leccinellum is a genus of fungi in the family Boletaceae. The genus is widespread in northern temperate areas, and contains an estimated 10 species.-Species:*Leccinellum albellum*Leccinellum corsicum*Leccinellum crocipodium...

Bresinsky & Manfr. Binder 2003 10 widespread
Leccinum
Leccinum
Leccinum is a genus of fungi in the family Boletaceae. It was the name given first to a series of fungi within the genus Boletus, then erected as a new genus last century. Their main distinguishing feature is the small, rigid projections that give a rough texture to their stalks...

Gray
Samuel Frederick Gray
Samuel Frederick Gray was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray.-Background:...

1821 widespread
Mycoamaranthus
Mycoamaranthus
Mycoamaranthus is a genus of fungi in the family Boletaceae. The genus contains three species, found in Australasia, Africa, and Southeast Asia....

Castellano, Trappe & Malajczuk 1992 3 Australasia; Africa, Southeast Asia
Octaviania
Octaviania
Octaviania is a genus of fungi in the family Boletaceae. The widespread genus is estimated to contain 15 species.-Species:*Octaviania asterosperma*Octaviania cyanescens*Octaviania hinsbyi*Octaviania ivoryana...

Vittad. 1831 15 widespread
Paxillogaster
Paxillogaster
Paxillogaster is a genus of fungi in the Boletaceae family. This is a monotypic genus, containing the single species Paxillogaster luteus.-External links:*...

E.Horak 1966 1 South America
Phylloboletellus
Phylloboletellus
Phylloboletellus is a genus of fungi in the Boletaceae family.-External links:*...

Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

1952 1 Central and South America
Phyllobolites
Phyllobolites
Phyllobolites is a genus of fungi in the family Boletaceae. The genus is monotypic, containing the single species Phyllobolites miniatus, found in tropical South America....

Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

1942 1 South America
Phylloporus
Phylloporus
Phylloporus is a genus of fungi in the Boletaceae family . The genus has a cosmopolitan distribution, and contains about 50 species, mostly in tropical areas.-Species:*Phylloporus albocarnosus Heinem....

Quel. 1888 cosmopolitan
Pseudoboletus
Pseudoboletus
Pseudoboletus is a genus of fungi in the family Boletaceae. The genus contains two species found in north temperate areas that grow in a parasitic association with species of Scleroderma and Pisolithus....

Šutara 1991 2 north temperate regions
Pulveroboletus
Pulveroboletus
Pulveroboletus is a genus of fungi in the Boletaceae family. The genus has a cosmopolitan distribution and contains 25 species.-Taxonomy:The genus was first described by American mycologist William Alphonso Murrill in 1909...

Murrill 1909 25 cosmopolitan
Retiboletus
Retiboletus
Retiboletus is a genus of fungi in the Boletaceae family. The genus, first described in 2002, contains six species distributed in north temperate regions....

Manfr. Binder & Bresinksy 2002 5 north temperate regions
Rhodactina
Rhodactina
Rhodactina is a genus of fungi in the family Boletaceae. The genus is monotypic, containing the single species Rhodactina himalayensis, found in India....

Pegler & T.W.K.Young 1989 1 India
Royoungia
Royoungia
Royoungia is a genus of fungi in the Boletaceae family. This is a monotypic genus, represented by the single species Royoungia boletoides.-External links:*...

Castellano, Trappe & Malajczuk 1992 1 Australia
Setogyroporus
Setogyroporus
Setogyroporus is a genus of fungi in the Boletaceae family. This is a monotypic genus, containing the single species Setogyroporus verus.-External links:*...

Heinem. & Rammeloo 1999 1 tropical Africa
Singeromyces
Singeromyces
Singeromyces is a genus of fungi in the Boletaceae family. This is currently a monotypic genus, represented by the single species Singeromyces ferrugineus.-External links:*...

M.M.Moser 1966 1 Argentina
Sinoboletus
Sinoboletus
Sinoboletus is a genus of fungi in the Boletaceae family.-External links:*...

M.Zang 1992 10 China
Spongiforma
Spongiforma
Spongiforma thailandica is a species of fungus in the Boletaceae family, genus Spongiforma. The stemless sponge-like species, first described in 2009, was found in Khao Yai National Park in central Thailand, where it grows in soil in old-growth forests...

Desjardin, Manf. Binder, Roekring & Flegel 2009 2 Thailand
Strobilomyces
Strobilomyces
Strobilomyces is a genus of boletes . The only well-known European species is the type species S. strobilaceus Strobilomyces is a genus of boletes (mushrooms having a spongy mass of pores under the cap). The only well-known European species is the type species S. strobilaceus Strobilomyces is a...

Berk. 1851 cosmopolitan
Tubosaeta
Tubosaeta
Tubosaeta is a genus of fungi in the Boletaceae family. The genus was circumscribed by mycologist Egon Horak in 1967....

E.Horak 1967 5 Africa; Asia
Tylopilus
Tylopilus
Tylopilus is a large genus of around 75 species of mycorrhizal bolete fungi separated from Boletus. Its best known member is the bitter bolete , the only species found in Europe. More species are found in North America, such as the edible species Tylopilus chromapes and T. alboater, and the...

P.Karst 1881 widespread
Veloporphyrellus
Veloporphyrellus
Veloporphyrellus is a genus of fungi in the Boletaceae family. The genus, circumscribed in 1984, contain two species....

L.D.Gómez & Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

1984 1 Central America
Wakefieldia
Wakefieldia
Wakefieldia is a genus of two species of fungi in the Boletaceae family.-External links:* at Index Fungorum...

Corner & Hawker 1952 2 Asia; Europe
Xanthoconium
Xanthoconium
Xanthoconium is a genus of fungi in the Boletaceae family. The genus has a cosmopolitan distribution and contains seven species.The concept of Xanthoconium has not been supported by DNA studies.-Species:*Xanthoconium affine...

Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

1944 7 cosmopolitan
Zangia
Zangia (fungus)
Zangia is a genus of fungi in the Boletaceae family. The genus, newly described in 2011, contains six species found in China....

Yan C.Li & Zhu L.Yang 2011 6 China


Many other genera formerly part of this family have been moved into other, smaller families as work with molecular phylogeny
Molecular phylogeny
Molecular phylogenetics is the analysis of hereditary molecular differences, mainly in DNA sequences, to gain information on an organism's evolutionary relationships. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree...

 shows that they are more distantly related, even if physically similar. Representative of this adjustment is the move of the slimy-capped genus Suillus
Suillus
Suillus is a genus of basidiomycete fungi in the family Suillaceae and order Boletales. Species in the genus are associated with coniferous trees, and are mostly distributed in northern temperate locations, although some species have been introduced to the Southern Hemisphere.-Taxonomy:The genus...

to Suillaceae
Suillaceae
The Suillaceae are a family of fungi in the order Boletales , containing the boletus-like Suillus, the small truffle-like Truncocolumella, as well as the monotypic genus Psiloboletinus. As of 2008, there are 54 species in the family...

.

Distribution

Boletes are found worldwide, on every continent except Antarctica. Well-known and well-described in the temperate latitudes in the northern hemisphere, newer research has shown significant diversity in tropical and southern hemisphere regions as well. E. J. H. Corner
E. J. H. Corner
Edred John Henry Corner FRS was a botanist who occupied the posts of assistant director at the Singapore Botanic Gardens and Professor of Tropical Botany at the University of Cambridge...

 found evidence of at least 60 species on the island of Singapore
Singapore
Singapore , officially the Republic of Singapore, is a Southeast Asian city-state off the southern tip of the Malay Peninsula, north of the equator. An island country made up of 63 islands, it is separated from Malaysia by the Straits of Johor to its north and from Indonesia's Riau Islands by the...

 alone. In 1972 he described 140 species from the Malay Peninsula
Malay Peninsula
The Malay Peninsula or Thai-Malay Peninsula is a peninsula in Southeast Asia. The land mass runs approximately north-south and, at its terminus, is the southern-most point of the Asian mainland...

 and Borneo
Borneo
Borneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....

 and estimated there were an equal number again to be found.

Similar statements about the biodiverse richness of Australian Boletaceae have also been made.

Edibility

Many of the boletes are considered to be true culinary delicacies, especially the king bolete (Boletus edulis
Boletus edulis
Boletus edulis, commonly known as penny bun, porcino or cep, 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 occur naturally in the Southern Hemisphere, although it has been...

); the Scandinavian cuisine praises boletes. In Finnish cuisine, the king bolete is universally considered to be the tastiest culinary mushroom. A large number of boletes are delicious or at least edible. Poisonous or otherwise inedible species do exist, however, such as the unpalatable bitter species Boletus calopus
Boletus calopus
Boletus calopus, commonly known as the bitter beech bolete or scarlet-stemmed bolete, is a fungus of the bolete family, found in Northern Europe and North America. Appearing in coniferous and deciduous woodland in summer and autumn, the fruiting bodies are attractively coloured, with a reddish...

and the aptly named bitter bolete (Tylopilus felleus
Tylopilus felleus
Tylopilus felleus, formerly Boletus felleus, is a fungus of the bolete family, found in Northern Europe and North America. Although it is not poisonous, it is not considered edible, due to its overwhelming bitterness.-Taxonomy:...

) with a taste compared to gall bladder juice, and some orange-capped species of Leccinum
Leccinum
Leccinum is a genus of fungi in the family Boletaceae. It was the name given first to a series of fungi within the genus Boletus, then erected as a new genus last century. Their main distinguishing feature is the small, rigid projections that give a rough texture to their stalks...

. As the bitter bolete resembles somewhat the king bolete, it can produce literally a bitter disappointment to the mushroom hunter! The rule of thumb is that bitter bolete has pink pores and brownish stem, while cep has whitish pale grey pores and stem. They also grow on different places and terrain. The peppery bolete (Chalciporus piperatus
Chalciporus piperatus
Chalciporus piperatus, commonly known as the Peppery bolete, is a small pored mushroom of the Boletaceae family found in mixed woodland in Europe....

) has extremely strong taste, and has been used in place of pepper
Black pepper
Black pepper is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. The fruit, known as a peppercorn when dried, is approximately in diameter, dark red when fully mature, and, like all drupes, contains a single seed...

.

Finnish cuisine uses boletes for various soups, sauces, casseroles and hotpots. They are sometimes also used as pizza filling, not unlike champignons, shiitake
Shiitake
The Shiitake is an edible mushroom native to East Asia, which is cultivated and consumed in many Asian countries, as well as being dried and exported to many countries around the world. It is a feature of many Asian cuisines including Vietnamese, Chinese, Japanese, Korean and Thai...

 or portobellos.

Two of the best common edible boletes however are the bay bolete (Boletus badius
Boletus badius
Boletus badius , and commonly known as the Bay Bolete is an edible, pored mushroom from Europe and North America...

), whose pores bruise blue-green, and the orange birch bolete, which is a Leccinum
Leccinum
Leccinum is a genus of fungi in the family Boletaceae. It was the name given first to a series of fungi within the genus Boletus, then erected as a new genus last century. Their main distinguishing feature is the small, rigid projections that give a rough texture to their stalks...

with an orange cap and which bruises a bluish grey.

Several guidebooks recommend avoiding all red-pored boletes, however both B. erythropus
Boletus erythropus
Boletus luridiformis, formerly known as Boletus erythropus, is a wild mushroom of the bolete family, all of which have tubes and pores, instead of gills beneath their caps. It is found in Northern Europe, and North America, and is sometimes commonly known as the Dotted Stem Bolete...

and B. luridus
Boletus luridus
Boletus luridus, commonly known as the lurid bolete, is a fungus of the bolete family, found in deciduous woodlands in Europe and eastern North America. Fruiting bodies arise in summer and autumn and may be common. It is a solid bolete with an olive-brown cap, orange pores and stout ochre stem...

are edible when well-cooked. However, there has been one recorded instance of death from Boletus pulcherrimus
Boletus pulcherrimus
Boletus pulcherrimus, commonly known as Alice Eastwood's boletus or the red-pored bolete, is a species of mushroom in the Boletaceae family. It is a large bolete from Western North America with distinguishing features that include a finely netted surface on the upper third of the stem, a red to...

in 1994; a couple developed gastrointestinal symptoms after eating this fungus with the husband succumbing. Autopsy revealed infarction
Infarction
In medicine, infarction refers to tissue death that is caused by a local lack of oxygen due to obstruction of the tissue's blood supply. The resulting lesion is referred to as an infarct.-Causes:...

 of the midgut. Boletus satanas has also long considered to be poisonous, though it has not been responsible for any deaths. The symptoms are predominantly gastrointestinal in nature. A glycoprotein
Glycoprotein
Glycoproteins are proteins that contain oligosaccharide chains covalently attached to polypeptide side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glycosylation. In proteins that have segments extending...

, bolesatine, has been isolated. A similar compound bolevenine has been isolated from the poisonous Boletus venenatus of Japan.
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