Boletus pinophilus
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''Boletus pinophilus'', commonly known as the pine bolete or pinewood king bolete, is a
basidiomycete Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Bas ...
fungus of the genus ''
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 ...
'' found throughout Europe and western Asia. Described by Italian naturalist
Carlo Vittadini Carlo Vittadini (11 June 1800, in Bertonico – 20 November 1865, in Milan) was an Italian doctor and mycologist. Life He studied in Milan and at the University of Pavia, where he attended the classes given by Giuseppe L. Moretti (1782–1 ...
in 1835, ''B. pinophilus'' was for many years considered a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
or
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 data * ...
of the porcini mushroom '' B. edulis'' before genetic studies confirmed its distinct status. In 2008, ''B. pinophilus'' in western North America were reclassified as a new species, '' B. rex-veris''. ''B. pinophilus'' is edible, and may be preserved and cooked. The fungus grows predominantly in
coniferous forest Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All exta ...
s on sandy soils, forming
ectomycorrhizal 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 my ...
associations in
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. Host trees include various species of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
, the
European silver fir ''Abies alba'', the European silver fir or silver fir, is a fir native to the mountains of Europe, from the Pyrenees north to Normandy, east to the Alps and the Carpathians, Slovenia, Croatia, Bosnia and Herzegovina, Montenegro, Serbia, and s ...
and
European spruce ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, 9–17 cm long. It is very close ...
, as well as deciduous trees such as
chestnut tree The chestnuts are the deciduous trees and shrubs in the genus ''Castanea'', in the beech family Fagaceae. They are native to temperate regions of the Northern Hemisphere. The name also refers to the edible nuts they produce. The unrelated ...
s, oak and
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 ...
. The fungus produces
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, ...
-bearing fruit bodies (known as "
mushroom A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans. The standard for the name "mushroom" is ...
s") above ground under pine trees in summer and autumn. It has a red-brown to maroon-coloured cap and a large and bulbous stipe, covered with coarse orange-red reticulation. As with other boletes, the size of the fruiting body is variable.


Description

The fruiting body has a convex-shaped
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. Caps typically have a visor, or no brim at all. They are popular in casual and informal se ...
, at first small in relation to its stipe, expanding in volume as it matures. The skin of the cap is dry, matte and can be coloured from maroon to chocolate brown with a reddish tint. It is thicker than other porcini-like boletes and is gelatinous. These characteristics distinguish it visually from relatives 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 ...
'', '' B. reticulatus'' and '' B. aereus''. The young, immature cap may have a pale pink colour and a white, powdery flush. As with all boletes, the size of the
fruiting body 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 cy ...
can vary greatly. The cap diameter can be as much as , the stem height and stem diameter . Measuring tall by wide, the bulbous stipe is often large, swollen and imposing, bearing a network pattern, much coarser in this species than other porcinis. The overall colour may have an orange-red tinge which is more obvious in the lowest parts, although this is also common in other species. Like other boletes, ''B. pinophilus'' has small pores on the underside of its cap rather than gills. These are coloured white at first, becoming yellow with age and olive-brown at full maturity. The
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 cylindric-ellipsoid, smooth, with oil drops and dimensions 15.5–20 by 4.5–5.5 
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 Uni ...
. They produce an olive-brown
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 ...
.


Taxonomy

Italian naturalist
Carlo Vittadini Carlo Vittadini (11 June 1800, in Bertonico – 20 November 1865, in Milan) was an Italian doctor and mycologist. Life He studied in Milan and at the University of Pavia, where he attended the classes given by Giuseppe L. Moretti (1782–1 ...
was the first to recognise the pine bolete as a distinct
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
, describing it as ''B. edulis'' var. ''pinicola'' in 1835. It was raised to species status (as ''B. pinicola'') by Antonio Venturi in 1863.
Pier Andrea Saccardo Pier Andrea Saccardo (23 April 1845 in Treviso, Treviso – 12 February 1920 in Padua) was an Italian botanist and mycologist. Life Saccardo studied at the Lyceum in Venice, and then at the Technical Institute of the University of Padua wher ...
treated it as a variety of ''Boletus aestivalis'' in 1910. It gained its current name in 1973, described by Czech mycologists
Albert Pilát Albert Pilát (November 2, 1903 – May 29, 1974) was a Czech botanist and mycologist. He studied at the Faculty of Science at Charles University, under the guidance of Professor Josef Velenovský. In 1930, he joined the National Museum, eventuall ...
and Aurel Dermek. A new
binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bo ...
had to be coined as ''B. pinicola'' as authored by Venturi was
invalid Invalid may refer to: * Patient, a sick person * one who is confined to home or bed because of illness, disability or injury (sometimes considered a politically incorrect term) * .invalid, a top-level Internet domain not intended for real use As ...
due that name having been previously applied to fungus now known as ''
Fomitopsis pinicola ''Fomitopsis pinicola'', is a stem decay fungus common on softwood and hardwood trees. Its conk (fruit body) is known as the red-belted conk. The species is common throughout temperate Europe and Asia. It is a decay fungus that serves as a s ...
''. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bo ...
is a mix of
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''pinus'' "pine", and
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
''philus'' "loving". ''B. pinophilus'' is classified in ''Boletus''
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 ...
''Boletus''; genetic analysis of European members in this group confirmed it is close to but genetically distinct from ''B. edulis'' and proposed maintaining its status as a separate species. In 2008, a taxonomic revision of
western North America Western North America is the western edge of the North American continent that borders the Pacific Ocean. It consists of Alaska at the farthest north, down through the western Canadian province of British Columbia, the western U.S. states of W ...
n populations of this species was published, formally establishing them as a distinct species, '' B. rex-veris''. Populations from eastern North America under pine have been reclassified as a new species '' B. pseudopinophilus''. Conversely, ''B. vinosulus''—described from the Czech Republic in 1992—has been since synonymised to this species.
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis has shown ''B. pinophilus'' as a member of a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
, or closely related group, with the North American species '' B. subcaerulescens'', '' B. subalpinus'', '' B. regineus'', '' B. fibrillosus'', and '' B. rex-veris''. Despite the diverse appearances, these taxa are close genetically, leading Feng and colleagues to speculate on combining the first four taxa above as a single species. These four diverged from the lineage that gave rise to ''B. fibrillosus'' and ''B. rex-veris'' around 5 million years ago; the common ancestor of all these diverged from the ancestor of ''B. edulis'' around 10 million years ago. The
British Mycological Society The British Mycological Society is a learned society established in 1896 to promote the study of fungi. Formation The British Mycological Society (BMS) was formed by the combined efforts of two local societies: the Woolhope Naturalists' Field ...
approved the name "pine bolete" for ''B. pinophilus''. Other common names include red king bolete, the pinewood king bolete, and ''cèpe des pins'' ("pine tree cep").


Distribution and habitat

''Boletus pinophilus'' is the rarest of the porcini-like mushrooms in Europe, though is found across temperate regions. It is found in Britain, where it is more common in Scotland, and in France, where it is more common in the south,
Leningrad Oblast Leningrad Oblast ( rus, Ленинградская область, Leningradskaya oblast’, lʲɪnʲɪnˈgratskəjə ˈobləsʲtʲ, , ) is a federal subject of Russia (an oblast). It was established on 1 August 1927, although it was not until 1 ...
and Vologda Oblast in Russia, and Ukraine. The bolete is considered vulnerable in the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
. It extends into
Asia Minor Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
and southwestern Asia, specifically as far as the
Perm Krai Perm Krai (russian: Пе́рмский край, r=Permsky kray, p=ˈpʲɛrmskʲɪj ˈkraj, ''Permsky krai'', , ''Perem lador'') is a federal subject of Russia (a krai) that came into existence on December 1, 2005 as a result of the 2004 re ...
in central Russia, and
Irkutsk Oblast Irkutsk Oblast (russian: Ирку́тская о́бласть, Irkutskaya oblast; bua, Эрхүү можо, Erkhüü mojo) is a federal subject of Russia (an oblast), located in southeastern Siberia in the basins of the Angara, Lena, and N ...
in Siberia. It is also found in exotic
pine plantation A tree plantation, forest plantation, plantation forest, timber plantation or tree farm is a forest planted for high volume production of wood, usually by planting one type of tree as a monoculture forest. The term ''tree farm'' also is used to ...
s (such as ''
Pinus sylvestris ''Pinus sylvestris'', the Scots pine (UK), Scotch pine (US) or Baltic pine, is a species of tree in the pine family Pinaceae that is native to Eurasia. It can readily be identified by its combination of fairly short, blue-green leaves and or ...
'') in eastern North America, Mexico, Chile, South Africa and New Zealand. Native populations from China and North America have been mostly confirmed as other distinct species.


Ecology

''B. pinophilus'' forms
ectomycorrhizal 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 my ...
relationships in
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
with
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
(''Pinus'') species such as stone pine ('' P. cembra''), black pine ('' P. nigra''), Corsican pine (''P. nigra'' subsp. ''laricio''), cluster pine ('' P. pinaster''), Monterey pine ('' P. radiata''), Scots pine ('' P. sylvestris'') and '' P. uncinata'', as well as European silver fir (''
Abies alba ''Abies alba'', the European silver fir or silver fir, is a fir native to the mountains of Europe, from the Pyrenees north to Normandy, east to the Alps and the Carpathians, Slovenia, Croatia, Bosnia and Herzegovina, Montenegro, Serbia, and sou ...
'') and European spruce (''
Picea abies ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, 9–17 cm long. It is very close ...
''). It can therefore be located wherever those trees grow, particularly with Scots pine in Britain, preferring the poor,
acidic In computer science, ACID ( atomicity, consistency, isolation, durability) is a set of properties of database transactions intended to guarantee data validity despite errors, power failures, and other mishaps. In the context of databases, a ...
, and sandy soils associated with
coniferous forests Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All exta ...
. It appears to favour ''Pinus'', while the form of the mushroom occurring in association with ''Abies'' and ''Picea'' has been labeled ''B. pinophilus'' var. ''fuscoruber''. However, it is not confined to
coniferous Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
trees and may also be found fruiting in
deciduous forest 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 ...
s, such as under chestnut trees (''
Castanea sativa ''Castanea sativa'', the sweet chestnut, Spanish chestnut or just chestnut, is a species of tree in the family Fagaceae, native to Southern Europe and Asia Minor, and widely cultivated throughout the Temperate climate, temperate world. A substa ...
''), oak (''
Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ...
''), beech (''
Fagus sylvatica ''Fagus sylvatica'', the European beech or common beech is a deciduous tree belonging to the beech family Fagaceae. Description ''Fagus sylvatica'' is a large tree, capable of reaching heights of up to tall and trunk diameter, though more ...
''), and possibly birch (''
Betula A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' contains 30 t ...
'' species), and hornbeam (''
Carpinus Hornbeams are hardwood trees in the flowering plant genus ''Carpinus'' in the birch family Betulaceae. The 30–40 species occur across much of the temperate regions of the Northern Hemisphere. Origin of names The common English name ''hor ...
''). Fruiting bodies can occur singly, or in small groups throughout the summer and autumn months, although they are known to appear as early as April in Italy. A 2007 field study on four species of boletes revealed little correlation between the abundance of fruit bodies and presence of its
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
below ground, even when
soil sample Soil test may refer to one or more of a wide variety of soil analysis conducted for one of several possible reasons. Possibly the most widely conducted soil tests are those done to estimate the plant-available concentrations of plant nutrients, i ...
s were taken from directly beneath the mushroom; the study concluded that the triggers leading to formation of mycorrhizae and production of the fruit bodies appear to be more complex than previously thought.


Uses

''Boletus pinophilus'' is edible, and may be used fresh, preserved, dried and cooked in a manner similar to that of other edible
bolete {{refimprove, date=July 2020 A bolete is a type of mushroom, or fungal fruiting body. It can be identified thanks to a unique mushroom cap. The cap is clearly different from the stem. On the underside of the cap there is usually a spongy surf ...
s. It is highly regarded and can be quite expensive in central Mexico, and is often sold dried there. The flesh is white, soft in mature specimens and does not change colour upon bruising. The taste and smell is pleasant. People of
La Malinche Marina or Malintzin ( 1500 – 1529), more popularly known as La Malinche , a Nahua woman from the Mexican Gulf Coast, became known for contributing to the Spanish conquest of the Aztec Empire (1519–1521), by acting as an interpreter, ad ...
have likened the flavour to pork and pork crackling. It is easily misidentified as the porcini ''B. edulis'', due to the similar habitat and appearance. It is a highly regarded food item, especially across the southern European regions of Portugal, the Basque Country and
Navarre Navarre (; es, Navarra ; eu, Nafarroa ), officially the Chartered Community of Navarre ( es, Comunidad Foral de Navarra, links=no ; eu, Nafarroako Foru Komunitatea, links=no ), is a foral autonomous community and province in northern Spain, ...
in Spain, France, Italy,
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedo ...
and
Serbia Serbia (, ; Serbian: , , ), officially the Republic of Serbia ( Serbian: , , ), is a landlocked country in Southeastern and Central Europe, situated at the crossroads of the Pannonian Basin and the Balkans. It shares land borders with Hu ...
. In the vicinity of Borgotaro in the Province of Parma of northern Italy, the four species ''B. edulis'', ''B. aereus'', ''B. reticulatus'', and ''B. pinophilus'' have been recognised for their superior taste and officially termed ''Fungo di Borgotaro''. Here, these mushrooms have been collected and exported commercially for centuries. It is a commonly eaten mushroom in Turkey, especially in the Marmara and Western
Black Sea region The Black Sea Region ( tr, Karadeniz Bölgesi) is a geographical region of Turkey. The largest city in the region is Samsun. Other big cities are Trabzon, Ordu, Tokat, Giresun, Rize, Amasya and Sinop. It is bordered by the Marmara Regio ...
s, and is exported to Europe. It is sold commercially in Finland. Fresh mushrooms are up to 90% water, and rich in carbohydrates. Unsaturated alcohols are a major component of the aroma of porcini mushrooms; 1-Octen-3-ol, 2-octen-1-ol, 3-Octanone, (E)-2-octenal,
oct-1-en-3-one Oct-1-en-3-one (CH2=CHC(=O)(CH2)4CH3), also known as 1-octen-3-one, is the odorant that is responsible for the typical "metallic" smell of metals and blood coming into contact with skin. Oct-1-en-3-one has a strong metallic mushroom-like odor with ...
and 1,7,7-trimethyl-heptan-2-one,
2-propenoic acid Acrylic acid (IUPAC: propenoic acid) is an organic compound with the formula CH2=CHCOOH. It is the simplest unsaturated carboxylic acid, consisting of a vinyl group connected directly to a carboxylic acid terminus. This colorless liquid has a ch ...
and 1,3-octadiene are the main volatile compounds in ''B. pinophilus''. ''B. pinophilus'' is known to be a bioaccumulator of the heavy metals mercury,
cadmium Cadmium is a chemical element with the Symbol (chemistry), symbol Cd and atomic number 48. This soft, silvery-white metal is chemically similar to the two other stable metals in group 12 element, group 12, zinc and mercury (element), mercury. Li ...
and
selenium Selenium is a chemical element with the symbol Se and atomic number 34. It is a nonmetal (more rarely considered a metalloid) with properties that are intermediate between the elements above and below in the periodic table, sulfur and tellurium, ...
. To reduce exposure, authorities recommend avoiding mushrooms from polluted areas such as those near mines, smelters, roadways, incinerators and disposal sites. Furthermore, pores should be removed as they contain the highest concentrations of pollutants.


References


External links

* {{Taxonbar, from=Q1415435 pinophilus Fungi of Europe Edible fungi Fungi described in 1973