Leccinum holopus
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''Leccinum holopus'', commonly known as the white birch bolete, white bog bolete, or ghost bolete, is a species of
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 ...
fungus in the family
Boletaceae The Boletaceae are a family of mushroom-forming fungi, primarily characterised by small pores on the spore-bearing hymenial surface (at the underside of the mushroom), instead of gills as are found in most agarics. Nearly as widely distributed ...
found in northern Asia, Europe, and northeastern North America. It associates with
birch 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 ...
trees and is typically found in boggy or swampy areas, often growing among
sphagnum moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store w ...
. Fruitbodies (
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 t ...
s) of ''L. holopus'' have convex
caps Caps are flat headgear. Caps or CAPS may also refer to: Science and technology Computing * CESG Assisted Products Service, provided by the U.K. Government Communications Headquarters * Composite Application Platform Suite, by Java Caps, a Java ...
measuring up to in diameter. Often pure white—especially in young fruitbodies—the caps sometimes become flushed with
buff Buff or BUFF may refer to: People * Buff (surname), a list of people * Buff (nickname), a list of people * Johnny Buff, ring name of American world champion boxer John Lisky (1888–1955) * Buff Bagwell, a ring name of American professional ...
or brownish tints. The whitish surface of the stipe is covered with small, stiff, projecting scales (scabers) that become tan or darker in age. Some
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
of ''Leccinum holopus'' have been described that vary in cap color or staining reaction, but DNA evidence suggests that most are the same
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 nam ...
. Although the fruitbodies are
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
, opinions vary as to their culinary desirability.


Taxonomy

Initially named as a species of ''
Boletus ''Boletus'' is a genus of mushroom-producing fungi, comprising over 100 species. The genus ''Boletus'' was originally broadly defined and described by Carl Linnaeus in 1753, essentially containing all fungi with hymenial pores instead of ...
'' by German mycologist Friedrich Rostkovius in 1844, the fungus was later transferred to ''
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 ...
'' by
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 fung ...
in 1960.
Synonyms 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 all ...
resulting from transfer to different genera include: ''Krombholzia holopoda'' and ''K. holopus'' (both published by
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, even ...
in 1951); ''Krombholziella holopus'' (Josef Šutara, 1989); ''Trachypus holopus'' (
Paul Konrad Paul Konrad (1 April 1877 in Le Locle – 19 December 1948 in Neuchâtel) was a Swiss geometrician and amateur mycologist. From 1902 he was an employee of the Compagnie des Tramways de Neuchâtel, of which, he served as a director from 1938 t ...
and
André Maublanc André Pierre Jules Maublanc (24 July 1880, in Nantes – 30 April 1958, in Paris) was a French mycologist and plant pathologist. Beginning in 1902, he worked as a préparateur at the Station de Pathologie végétale in Paris. In 1912, he traveled ...
, 1952), and ''Trachypus scaber'' f. ''holopus'' (
Henri Romagnesi Henri Charles Louis Romagnesi (7 February 1912 – 18 January 1999) was a French mycologist who was notable for a thorough review and monograph of the agaric genus '' Entoloma'' (or ''Rhodophyllus'' as it was known in the early 20th century), as ...
, 1939). Other synonyms, according to
Index Fungorum ''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and the Institute of M ...
, include ''Leccinum olivaceosum'', described from France in 1994, and ''Leccinum aerugineum'' (1991). ''Leccinum holopus'' is classified in
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 ...
''Scabra'' of genus ''Leccinum'', a grouping that includes
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
species associating exclusively with birch. 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, bot ...
''holopus'' is
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 ...
for "with perfect stalk".
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 contrast ...
s given to the fungus include white birch bolete, white bog bolete, and ghost bolete. Several subtaxa of ''Leccinum holopus'' have been described. In
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 ...
''aerugineum'', described by Josef Šutara in 2009, the flesh discolors green after injury. The
variety Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
''americanum'', described by
Alexander H. Smith Alexander Hanchett Smith (December 12, 1904 – December 12, 1986) was an American mycologist known for his extensive contributions to the taxonomy and phylogeny of the higher fungi, especially the agarics. Early life Smith, born in Crandon, Wi ...
and
Harry Delbert Thiers Harry Delbert Thiers (January 22, 1919 in Fort McKavett, Texas – August 8, 2000 in Ohio) was an American mycologist who studied and named many fungi native to North America, particularly California. Thiers taught mycology at San Francisco S ...
in 1971 from collections made in
Michigan Michigan () is a state in the Great Lakes region of the upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the 10th-largest state by population, the 11th-largest by area, and the ...
, injured flesh stains reddish. Lannoy & Estadès described ''Leccinum nucatum'' in 1993, a taxon that was later (2007) published as variety ''nucatum'' of ''L. holopus''; no
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...
evidence was found supporting the existence of this as a distinct taxon, and it is therefore placed into synonymy with ''L. holopus''. ''Leccinum holopus'' var. ''majus'', described by
Rolf Singer Rolf Singer (June 23, 1906 – January 18, 1994) was a Germany, German-born mycologist and one of the most important Taxonomy (biology), taxonomists of gilled mushrooms (agarics) in the 20th century. After receiving his Ph.D. at the University ...
in 1966 (originally published by Singer as ''Krombholzia scabra'' f. ''majus''), is another historical variety without independent
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
significance.


Description

Fruitbodies of ''Leccinum holopus'' have convex to flattened
caps Caps are flat headgear. Caps or CAPS may also refer to: Science and technology Computing * CESG Assisted Products Service, provided by the U.K. Government Communications Headquarters * Composite Application Platform Suite, by Java Caps, a Java ...
measuring in diameter, with a narrow band of sterile tissue surrounding the margin. The caps are initially whitish, but can develop gray,
buff Buff or BUFF may refer to: People * Buff (surname), a list of people * Buff (nickname), a list of people * Johnny Buff, ring name of American world champion boxer John Lisky (1888–1955) * Buff Bagwell, a ring name of American professional ...
, tan, or pinkish tints during maturity; the color may also darken and become greenish with age. The cap surface is initially covered with very fine hairs, but later becomes more or less smooth, often with a sticky texture in age or in moist conditions. The
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 muscle Skeletal muscles (commonly referred to as mu ...
is white and lacks any distinct odor or taste; it can have either little or no bruising color reaction with injury, or may become light pink in variety ''americanum''. On the cap underside is a porous surface comprising pores numbering 2 to 3 per millimeter, each of which is the end of a tube that extends to deep. The color of the pore surface ranges from whitish to grayish to dingy brown, and has little color reaction to injury, although it may discolor yellowish or brownish. There is a depression where the pores meet the stipe. The stipe measures long by wide. Its whitish surface is covered with scabers that darken in age to tan or darker. The stipe base often stains bluish. ''Leccinum holopus'' produces a 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 ...
.
Spores 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, f ...
are somewhat fusoid (spindle-shaped) and measure 14–20 by 5–6.5  µm. 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-c ...
(spore-bearing cells) are four-spored and measure 28.5–36.5 by 11.5–12.5 µm.
Cystidia A cystidium (plural cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that ar ...
on the pores are flask-shaped (lageniform) to fusiform, and 39.0–45.5 by 7.5–9.0 µm, while those of the stipe (caulocystidia) are fusiform, club-shaped, or cylindrical, measuring 39.0–54.5 x 9.1–13.5 µm. There are no
clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rec ...
s present in the
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e of ''L. holopus''. The
cap cuticle The pileipellis is the uppermost layer of hyphae in the pileus of a fungal fruit body. It covers the trama, the fleshy tissue of the fruit body. The pileipellis is more or less synonymous with the cuticle, but the cuticle generally describes ...
is arranged in the form of a cutis—with hyphae that run parallel to the cap surface. Several chemical tests can be used to help verify an identification of ''L. holopus''. A drop of
ammonium hydroxide Ammonia solution, also known as ammonia water, ammonium hydroxide, ammoniacal liquor, ammonia liquor, aqua ammonia, aqueous ammonia, or (inaccurately) ammonia, is a solution of ammonia in water. It can be denoted by the symbols NH3(aq). Although ...
solution turns the cap cuticle a pinkish color, but has no reaction with the flesh. A drop of dilute
potassium hydroxide Potassium hydroxide is an inorganic compound with the formula K OH, and is commonly called caustic potash. Along with sodium hydroxide (NaOH), KOH is a prototypical strong base. It has many industrial and niche applications, most of which exp ...
(KOH) has no reaction on the cap surface, and either no reaction or a brownish reaction with the flesh. Application of
iron(II) sulfate Iron(II) sulfate (British English: iron(II) sulphate) or ferrous sulfate denotes a range of salts with the formula Fe SO4·''x''H2O. These compounds exist most commonly as the heptahydrate (''x'' = 7) but several values for x are know ...
solution does not have a reaction on the cap surface, and either no reaction to slightly olive coloration on the flesh.


Similar species

''
Leccinellum albellum ''Leccinellum albellum'' is a species of bolete fungus in the family Boletaceae. Originally described by Charles Horton Peck as a species of ''Boletus'', and, after 1945, usually considered a species of ''Leccinum'', it was transferred to the ne ...
'' is similar in appearance to ''L. holopus'', but grows in association with
oak 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 ''L ...
and has a more southerly distribution. '' L. scabrum'' is a widely distributed lookalike that can be distinguished from ''L. holopus'' by its larger size and generally darker colors.


Edibility

Although commonly considered
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
, opinions vary on the culinary appeal of ''Leccinum holopus'' fruitbodies. Michael Kuo, writing in ''100 Edible Mushrooms'', considers it a good edible; Peter Roberts and Shelley Evans in ''The Book of Fungi'' say "it is edible, but is said to be tasteless and pappy, so is not recommended." Fruitbodies are optimally harvested when they are young, before the flesh becomes too spongy, and before insect larvae establish themselves. Minimal cleaning is required in the field. The mushroom has a mild, somewhat sweet flavor that is enhanced after brief sauteeing. Drying the mushrooms enhances the flavor, but diminishes the sweetness of fresh mushrooms.


Habitat and distribution

''Leccinum holopus'' is a
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l species. It fruits on the ground (often among ''
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
'' moss), singly to scattered in wet areas like
cedar Cedar may refer to: Trees and plants *''Cedrus'', common English name cedar, an Old-World genus of coniferous trees in the plant family Pinaceae *Cedar (plant), a list of trees and plants known as cedar Places United States * Cedar, Arizona * ...
swamps,
bogs A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main Wetland#Types, types of wetlands. Other names for bogs include mire, mosses, quagmire, ...
, or soggy forests. Like most ''Leccinum'' species, the fungus is highly host-specific and associates with
birch 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 ...
(''Betula''). In North America, the range of ''Leccinum holopus'' extends from eastern Canada to
New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
, extending west to the northern
Rocky Mountain 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 ...
s, roughly coinciding with the distribution of the
paper birch ''Betula papyrifera'' (paper birch, also known as (American) white birch and canoe birch) is a short-lived species of birch native to northern North America. Paper birch is named for the tree's thin white bark, which often peels in paper like ...
(''Betula papyrifera''). In this range, it is common and fruits from August to October. ''L. holopus'' var. ''americanum'' is known only from North America. The fungus is rare in southern Europe, but more common in the ''Sphagnum'' swamps in the north. In Asia, it has been recorded from Taiwan and
Qinghai Qinghai (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest po ...
(China). Fruitbodies of ''L. holopus'' are a source of food for
fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
species such as ''
Pegomya winthemi ''Pegomya winthemi'' is a species of fly in the family Anthomyiidae. Found in North America, it was first described as ''Anthomyia winthemi'' in 1829 by Johann Wilhelm Meigen. The insect measures 4.25–5 mm long. Its pedipalps are infuscated ...
'' (family
Anthomyiidae The Anthomyiidae are a large and diverse family of Muscoidea flies. Most look rather like small houseflies, but are commonly drab grey. The genus ''Anthomyia'', in contrast, is generally conspicuously patterned in black-and-white or black-and- ...
) and '' Megaelia pygmaeoides'' (family
Phoridae The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of thei ...
).


See also

* List of ''Leccinum'' species *
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

{{Taxonbar, from=Q1328979 Edible fungi Fungi described in 1844 Fungi of Asia Fungi of Europe Fungi of North America
holopus ''Holopus'' is a genus of echinoderms belonging to the family Holopodidae. The species of this genus are found in Central America, Pacific Ocean (near Australia). Species: *''Holopus alidis'' *''Holopus mikihe'' *''Holopus rangii'' Referenc ...