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Boletus pinophilus
Boletus pinophilus
Boletus pinophilus
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Boletus pinophilus
Scientific classification Edit this classification
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Boletales
Family: Boletaceae
Genus: Boletus
Species:
B. pinophilus
Binomial name
Boletus pinophilus
Pilát & Dermek (1973)[1]
Synonyms
  • Boletus aestivalis var. pinicola
    (Vittad.) Sacc[2]
  • Boletus edulis var. pinicola
    Vitt.[3]
  • Boletus edulis f. pinicola
    (Vittad.) Vassilkov[4]
  • Boletus pinicola
    (Vitt.) Venturi[3]
  • Boletus vinosulus
    (Kluzák, Papoušek & Šutara)[5]
Boletus pinophilus
Mycological characteristics
Pores on hymenium
Cap is convex
Hymenium is adnexed
Stipe is bare
Spore print is olive-brown
Ecology is mycorrhizal
Edibility is choice

Boletus pinophilus, commonly known as the pine bolete or pinewood king bolete, is a basidiomycete fungus of the genus Boletus. Described by Italian naturalist Carlo Vittadini in 1835, B. pinophilus was for many years considered a subspecies or form 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.

The fungus produces spore-bearing fruit bodies (i.e. mushrooms) 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.

The fungus is found throughout Europe and western Asia. It grows predominantly in coniferous forests on sandy soil, forming ectomycorrhizal associations in symbiosis with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. Host trees include various species of pine, the European silver fir and European spruce, as well as deciduous trees such as chestnut trees, oak and beech. B. pinophilus is edible, and may be preserved and cooked.

Taxonomy

[edit]

Italian naturalist Carlo Vittadini was the first to recognise the pine bolete as a distinct taxon, describing it as B. edulis var. pinicola in 1835.[6] It was raised to species status (as B. pinicola) by Antonio Venturi in 1863. Pier Andrea Saccardo treated it as a variety of Boletus aestivalis in 1910.[7] It gained its current name in 1973, described by Czech mycologists Albert Pilát and Aurel Dermek. A new binomial name had to be coined as B. pinicola as authored by Venturi was invalid due that name having been previously applied to fungus now known as Fomitopsis pinicola.[8] The specific epithet is a mix of Latin pinus "pine",[9] and Ancient Greek philus "loving".[10] B. pinophilus is classified in Boletus section 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.[11]

In 2008, a taxonomic revision of western North American populations of this species was published, formally establishing them as a distinct species, B. rex-veris.[12] 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.[5]

Phylogenetic analysis has shown B. pinophilus as a member of a clade, or closely related group, with the North American species B. subcaerulescens, B. subalpinus, B. regineus, B. fibrillosus, and B. rex-veris.[13] 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.[14] 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.[14]

The British Mycological Society approved the name "pine bolete" for B. pinophilus.[15] Other common names include red king bolete,[5] the pinewood king bolete,[16] and cèpe des pins ("pine tree cep").[17]

Description

[edit]

The fruiting body has a convex-shaped cap, 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.[18] It is thicker than other porcini-like boletes and is gelatinous.[11] These characteristics distinguish it visually from relatives such as Boletus edulis, 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 can vary greatly. The cap diameter can be as much as 40 centimetres (16 inches), the stem height 25 cm (10 in) and stem diameter 16 cm (6+1⁄4 in).[18] Measuring 4 to 10 cm (1+1⁄2 to 4 in) tall by 3 to 8 cm (1+1⁄4 to 3+1⁄4 in) wide, the bulbous stipe is often large, swollen and imposing, bearing a network pattern, much coarser in this species than other porcinis.[11] 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 spores are cylindric-ellipsoid, smooth, with oil drops and dimensions 15.5–20 by 4.5–5.5 μm. They produce an olive-brown spore print.[18]

Distribution and habitat

[edit]

Boletus pinophilus is the rarest of the porcini-like mushrooms in Europe, though is found across temperate regions.[5] It is found in Britain,[18] where it is more common in Scotland, and in France, where it is more common in the south,[19] Leningrad Oblast and Vologda Oblast in Russia,[20] and Ukraine.[21] The bolete is considered vulnerable in the Czech Republic.[22] It extends into Asia Minor and southwestern Asia,[5] specifically as far as the Perm Krai in central Russia,[23] and Irkutsk Oblast in Siberia.[20] It is also found in exotic pine plantations (such as Pinus sylvestris) in eastern North America, Mexico, Chile, South Africa and New Zealand.[5] Native populations from China and North America have been mostly confirmed as other distinct species.[5]

Ecology

[edit]

B. pinophilus forms ectomycorrhizal relationships in symbiosis with pine (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) and European spruce (Picea abies).[5] It can therefore be located wherever those trees grow, particularly with Scots pine in Britain,[24] preferring the poor, acidic, and sandy soils associated with coniferous forests. 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.[11] However, it is not confined to coniferous trees and may also be found fruiting in deciduous forests, such as under chestnut trees (Castanea sativa),[19] oak (Quercus), beech (Fagus sylvatica), and possibly birch (Betula species), and hornbeam (Carpinus).[5]

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.[25] A 2007 field study on four species of boletes revealed little correlation between the abundance of fruit bodies and presence of its mycelia below ground, even when soil samples 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.[26]

Uses

[edit]
Undersides of collected specimens

Boletus pinophilus is edible and may be used fresh, preserved, dried and cooked in a manner similar to that of other edible boletes.[27][28] It is highly regarded and can be quite expensive in central Mexico, and is often sold dried there.[29] The flesh is white, soft in mature specimens and does not change colour upon bruising. The taste and smell is pleasant.[27] People of La Malinche have likened the flavour to pork and pork crackling.[29] It is easily misidentified as the porcini B. edulis, due to the similar habitat and appearance.[30] It is a highly regarded food item, especially across the southern European regions of Portugal, the Basque Country and Navarre in Spain, France, Italy, Bulgaria and Serbia.[31] 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.[32] It is a commonly eaten mushroom in Turkey, especially in the Marmara and Western Black Sea regions, and is exported to Europe.[33] It is sold commercially in Finland.[34]

Fresh mushrooms are up to 90% water, and rich in carbohydrates.[31] 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 and 1,7,7-trimethyl-heptan-2-one, 2-propenoic acid and 1,3-octadiene are the main volatile compounds in B. pinophilus.[33] B. pinophilus is known to be a bioaccumulator of the heavy metals mercury, cadmium and selenium.[16][35] 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.[36]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Boletus pinophilus

View on Grokipedia
from Grokipedia
Boletus pinophilus, commonly known as the pine bolete or pinewood king bolete, is a species of ectomycorrhizal in the family , recognized for its robust, fruiting bodies that feature a vinaceous to reddish-brown cap up to 20 cm in diameter and a stout, reticulate stem. This forms symbiotic associations primarily with pine trees, such as , in coniferous or mixed forests on acidic, sandy soils, contributing to nutrient cycling in these ecosystems. Native to , it is widespread from Britain to the and eastern regions, fruiting in summer and autumn, and is valued for its culinary qualities similar to the closely related . First described in 1973 by Czech mycologists Augustin Pilát and Vladimír Dermek, B. pinophilus was previously classified under synonyms like Boletus pinicola or as a variety of B. edulis, reflecting ongoing taxonomic refinements within the Boletus section Edules. Morphologically, the cap is hemispherical when young, becoming convex to flat with age, dry to slightly viscid, and colored in shades of reddish-brown without changing upon bruising; the tubes are adnate, starting white and turning yellow, with pores that remain unchanging or faintly bluing. The stem, often clavate and up to 12 cm tall, is pale brown with a prominent white to pale brown , especially near the apex, while the firm, white flesh has a mild, pleasant odor and taste. Spores are elliptical, measuring 13–21 × 3.5–6.5 μm, producing an olive-brown . Ecologically, B. pinophilus plays a key role in forest health by enhancing tree growth through mycorrhizal partnerships, preferentially occurring in mature plantations and old-growth woodlands at elevations up to 1,800 meters in cool, humid climates. It is uncommon in parts of Britain but more abundant in , including , , and , where it favors well-drained, nutrient-poor sites under Scots pine or occasionally and . Though not reported in , its European range supports commercial , and it is considered a when young and firm, with no significant toxic look-alikes when properly identified.

Taxonomy

Nomenclature and History

Boletus pinophilus was first described as a variety of the porcini mushroom, var. pinicola, by the Italian naturalist Carlo Vittadini in his 1835 work Descrizione dei funghi mangerecci più comuni d'Italia. This initial recognition highlighted its distinct characteristics associated with pine habitats, though it was treated as a subordinate under the broadly defined B. edulis at the time. In 1863, Italian mycologist Antonio Venturi elevated this taxon to species level, renaming it Boletus pinicola in his publication I Miceti dell'Agro Bresciano. This change reflected growing appreciation for its morphological and ecological differences from B. edulis, particularly its preference for forests, but the name Boletus pinicola proved problematic as it was a later of the bracket fungus Boletus pinicola Swartz (now ) described in 1810. To resolve the nomenclatural conflict, Czech mycologist Augustin Pilát and Slovak mycologist Vladimír Dermek proposed the current accepted name, , in 1973 within the journal Česká Mykologie. This replacement epithet, derived from the Latin pinus (pine) and philus (loving), underscores the fungus's strong mycorrhizal association with Pinus species. These key publications by Vittadini, Venturi, Pilát, and Dermek mark the primary taxonomic developments, establishing B. pinophilus as a distinct species within the .

Synonyms and Classification

Boletus pinophilus has several historical synonyms, including Boletus edulis var. pinicola Vittad. (1835), which was the original description based on Italian specimens associated with pines, and Boletus aestivalis var. pinicola (Vittad.) Sacc. (1910), reflecting early taxonomic confusion within the Boletus edulis complex. Other synonyms include Boletus pinicola (Vittad.) Rea (1922), though this name is a later homonym and not accepted. The species was formally elevated to specific rank as Boletus pinophilus Pilát & Dermek in 1973, distinguishing it from the summer-fruiting B. aestivalis. In modern taxonomy, pinophilus is placed in the Boletus Fr. (1821), section Boletus, family Chevall. (1826), order E.-J. Gilbert (1931). This placement reflects its poroid hymenophore and reticulate stipe, characteristic of the core porcini group. North American populations previously identified as B. pinophilus have undergone reclassification based on molecular data. Western collections from and the , fruiting in spring under pines, were described as the distinct species Boletus rex-veris Arora & Simonini in 2008, supported by morphological and unpublished ITS sequence differences. Eastern populations under hard pines, such as Pinus spp. in the southeastern U.S., were recognized as Boletus pseudopinophilus A.R. Bessette, A.E. Bessette, J. Craine & J.L. Frank in 2019, following ITS rDNA analysis that placed it in a supported sister to European B. pinophilus. Phylogenetic studies using multi-locus (ITS, LSU, RPB1, ATP6) have confirmed B. pinophilus as a monophyletic species separate from the B. edulis complex, with divergence estimated at 42–54 million years ago, highlighting cryptic diversity within the porcini .

Description

Macroscopic Features

The fruiting body of Boletus pinophilus features a robust that measures 5–25 cm in diameter, initially convex to hemispherical but flattening with age, often developing wrinkles or cracks on the surface; the cap is smooth to slightly felty or greasy, turning viscid in wet conditions, and colored vinaceous brown to maroon or chocolate-brown. The stipe is stout and clavate to bulbous, reaching 5–15 cm in height and 3–8 cm thick at the base, with a pale yellow to light brown or orange-red hue marked by coarse reticulation especially on the upper portion; it often tapers upward and features white basal . The pore surface under the cap starts white to pale yellow, becoming olive-brown with maturity and arranged in 2–4 pores per millimeter; bruising reaction is absent to slight bluing or blue-green in some specimens. The flesh is white, firm, and unchanging upon exposure or cutting, though occasionally tinged vinaceous below the cap cuticle or with pinkish hues near the stipe base. The is mild and pleasant, mushroom-like, while the taste is nutty and agreeable. The is olive-brown.

Microscopic Features

The microscopic features of Boletus pinophilus are characteristic of the section Boletus within the family, aiding in its identification through detailed examination of reproductive and structural elements. The basidiospores are to cylindric-ellipsoid in shape, measuring 13–21 × 3.5–6.5 μm, with a smooth surface and thick walls; they contain oil drops and appear yellowish brown under the . These spores are typically inamyloid in Melzer's reagent, consistent with other species in the genus. The basidia, which bear the spores, are club-shaped (clavate) and four-spored, with dimensions of 25–35 × 7–10 μm; they are produced abundantly on the hymenial surface of . Hymenial cystidia are absent or present only sparsely. The pileipellis, or cuticle, consists of a trichodermial layer of interwoven, septate hyphae that are slightly to moderately gelatinized, with short cylindrical end-cells that may appear incrusted or granulated; these elements measure up to 25–90 × 14–35 μm and contribute to the 's greasy texture when fresh. Caulocystidia on the stipe surface are cylindrical to slightly lageniform, 58–66 × 10–14 μm, but do not significantly alter the overall microscopic profile.

Distribution and Habitat

Geographic Range

Boletus pinophilus is native to and western , where it inhabits temperate to boreal pine forests. It is widespread across the continent, with records from numerous countries including , , , , , , , , Italy, the Netherlands, , the Russian Federation, , , , , , and the . The species is particularly abundant in and , reflecting its preference for coniferous-dominated landscapes. In , B. pinophilus is common in and , often fruiting prolifically in Scots pine () stands. Similarly, it occurs frequently in , where it contributes to the mycorrhizal community in forested areas. Its distribution extends southward into the Mediterranean region, though abundance decreases in warmer climates. In western Asia, the fungus is reported in areas such as Asia Minor and southwestern regions, associated with native pine species. The species is rarer in certain peripheral regions, such as the , where it is uncommon and primarily found in mature pine plantations in and . In the , B. pinophilus is classified as vulnerable due to limited distribution and habitat pressures, while it is assessed as Least Concern globally by the IUCN.

Environmental Preferences

Boletus pinophilus prefers sandy, acidic soils with a pH typically ranging from 3.5 to 6.5, which are well-drained and often nutrient-poor, facilitating its growth in environments with limited fertility such as lichen-heathland forests. These conditions are commonly associated with coniferous landscapes where organic matter accumulation is low, promoting the fungus's ectomycorrhizal development without excessive competition from other soil microbes. The species thrives in temperate to boreal climates featuring cool summers, mild winters, and moderate , with fruiting bodies emerging mainly during summer and autumn following periods of rainfall and subsequent dry spells. Such climatic patterns support mycelial expansion in forested areas, aligning with the thermal tolerances observed in related species. It forms mycorrhizal associations primarily with and other pine species (Pinus spp.), but also occurs with (Abies spp.), spruces (Picea spp.), and occasionally deciduous trees such as oaks (Quercus spp.) or birches (Betula spp.). These partnerships are most productive in open woodlands, plantations, and mixed coniferous stands up to altitudes of approximately 1,500–1,800 m, where light penetration and tree spacing enhance sporocarp formation.

Ecology

Mycorrhizal Associations

Boletus pinophilus is an ectomycorrhizal that forms mutualistic associations with the roots of (Pinus) species, where it envelops fine root tips to create a structure that facilitates nutrient exchange between the symbionts. These associations enhance the host tree's uptake of immobile nutrients, particularly and , which are often limiting in forest soils. In return, the fungus receives carbohydrates from the tree's photosynthetic activity, providing the source for its growth and maintenance. The fungus exhibits a strong host specificity for Pinus species, such as and , forming robust ectomycorrhizae that support tree vigor in nutrient-poor environments. However, it is also compatible with other conifers including Picea and Abies, though such associations are less frequent and sometimes designated as varieties like B. pinophilus var. fuscoruber. Compatibility extends occasionally to deciduous trees like , broadening its potential ecological interactions in mixed forests. In forest , B. pinophilus plays a vital role by promoting growth on acidic, sandy, or impoverished through improved acquisition and relations. Its extensive extraradical mycelial networks further contribute to soil aggregation and structure, enhancing overall ecosystem resilience and facilitating cycling among plants.

Fruiting and Life Cycle

The mycelium of Boletus pinophilus persists year-round in the , forming extensive networks that maintain the fungus's symbiotic association with tree roots. This vegetative phase allows the to absorb nutrients and from the host while remaining dormant during unfavorable conditions, such as dry winters or hot summers. Fruiting bodies only develop when specific environmental cues stimulate the transition from mycelial growth to reproductive structures, a process that can take several years to establish in new sites. The primary fruiting season for B. pinophilus occurs from late summer to autumn, typically June through October across much of , when conditions favor sporocarp production. In warmer Mediterranean regions like , fruiting can occasionally begin in spring, around , particularly after early rains in pine-dominated woodlands. These seasonal patterns align with the fungus's ectomycorrhizal lifestyle, where it briefly references its dependence on hosts for sustained mycelial health. Fruiting is triggered by elevated following rainfall, combined with moderate temperatures of 15–20°C and high , which signal the to initiate primordia formation. Studies on related ectomycorrhizal boletes indicate that hormonal signals from host trees, such as auxin-like compounds, may further coordinate this response by influencing fungal for reproductive development. These cues ensure synchronized emergence to maximize release during optimal dispersal windows. Primordia initially form underground as compact hyphal aggregates near the soil surface, expanding rapidly into mature fruiting bodies that push through the litter layer over several days. Once emerged, the pores on the underside release billions of olivaceous-brown spores, primarily dispersed by wind currents over short to moderate distances within forests. Animals, including and small mammals, contribute to longer-range dispersal by consuming and excreting viable spores, facilitating colonization of new host trees.

Uses and Edibility

Culinary Applications

Boletus pinophilus is considered a choice , prized for its culinary versatility and safe consumption when properly cooked. It is particularly popular in southern European countries such as , , , and , where it features prominently in regional gastronomy, as well as in , including and , among foraging enthusiasts. The can be utilized fresh, dried, or frozen, allowing for year-round availability in dishes. Common preparation methods include sautéing in with , grilling, or incorporating into soups, stews, risottos, and creamy sauces; young specimens are preferred for their tenderness, as mature ones may develop a tougher texture. Its flavor profile is characterized by nutty, earthy notes with subtle sweetness and a firm, meaty texture that holds up well during cooking; when dried, it retains and even concentrates these qualities, making it ideal for rehydration in or powdered for . Culturally, B. pinophilus is commercially foraged in pine-dominated forests across , supporting local economies through harvesting and export, particularly from Eastern European countries like , , and to markets in .

Nutritional Value and Risks

Boletus pinophilus is valued for its nutritional profile, offering a low-calorie option with approximately 38 kcal per 100 grams of fresh weight, making it suitable for diets emphasizing moderation. On a dry weight basis, it contains moderate protein levels around 7.42 g per 100 g, alongside high carbohydrates at 83.21 g per 100 g, which contribute to its role as a nutrient-dense . It is also rich in , supporting digestive health, and provides essential vitamins including vitamins and , which are naturally present or enhanced through exposure in specimens via conversion of to vitamin D2. Minerals such as , at levels exceeding 11,000 mg/kg dry weight in the cap, and , averaging 19.9 mg/kg dry weight, further bolster its nutritional benefits, with selenium acting as an cofactor. The harbors bioactive compounds, including antioxidants and , which exhibit potential immune-boosting effects through modulation of and enhancement of cellular defenses. from Boletus , including those akin to B. pinophilus, have demonstrated immunomodulatory properties in studies, promoting production and immune cell activity without direct toxicity. These compounds contribute to the overall health-promoting potential of the , though specific quantification in B. pinophilus remains limited compared to related . Despite its edibility, B. pinophilus poses risks due to its capacity to accumulate heavy metals, particularly mercury and cadmium, from polluted soils. Mean mercury concentrations reach 6.9 mg/kg dry weight in the hymenophore and 4.5 mg/kg in the rest of the fruiting body, with cadmium at 0.579 mg/kg dry weight, exceeding safe thresholds in contaminated areas and potentially leading to toxicological concerns upon regular consumption. Rare allergic reactions, including IgE-mediated responses, have been reported for bolete mushrooms, manifesting as gastrointestinal distress or respiratory issues in sensitive individuals. While no inherent toxins are known in B. pinophilus, overconsumption is advised against to mitigate cumulative metal exposure. Post-2000 studies affirm selenium enrichment as a nutritional asset but highlight mercury bioaccumulation as a persistent hazard, recommending harvest from unpolluted sites to minimize risks.

References

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