Hygrophorus Eburneus
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''Hygrophorus eburneus'', commonly known as the ivory waxy cap or the cowboy's handkerchief, is a species of
edible mushroom Edible mushrooms are the fleshy fruit bodies of numerous species of macrofungi (fungi that bear fruiting structures large enough to be seen with the naked eye). Edibility may be defined by criteria including the absence of poisonous effect ...
in the waxgill family of
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
. ''Hygrophorus eburneus'' is the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the genus ''Hygrophorus''. The
fruit bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
are medium-sized, pure white, and when wet are covered in a layer of slime thick enough to make the mushroom difficult to pick up. The
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
are broadly attached to the
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
or running down it; as the family name suggests, they feel waxy when rubbed between the fingers. It is widespread in Europe and North America, and has also been collected in northern Africa. Like all ''Hygrophorus'' species, the fungus is
mycorrhiza A mycorrhiza (; , 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's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...
l—having a
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
association whereby the underground fungal
mycelia Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...
penetrate and exchange nutrients with tree roots. They are common in a variety of forest types, where they grow on the ground in thickets or grassy areas. A number of biologically active chemicals have been purified from the fruit bodies of the fungus, including
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s with
bactericidal A bactericide or bacteriocide, sometimes abbreviated Bcidal, is a substance which kills bacteria. Bactericides are disinfectants, antiseptics, or antibiotics. However, material surfaces can also have bactericidal properties based solely on their p ...
and
fungicidal Fungicides are pesticides used to kill parasitic fungi or their spores. Fungi can cause serious damage in agriculture, resulting in losses of yield and quality. Fungicides are used both in agriculture and to fight fungal infections in animals, ...
activity.


Taxonomy

The species was first named as ''Agaricus eburneus'' by French botanist Jean Bulliard in 1783.
Elias Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. He is sometimes called the "Linnaeus of Mycology". In his works he described and assigned botanical names to hundreds of fungus and lichen sp ...
divided the large genus ''Agaricus'' into a number of "tribes" (
taxonomically In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon), and these groups are given ...
equivalent to modern
sections 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 ...
) in his ''Systema Mycologicum I'', and classified ''A. eburneus'' in the tribe ''Limacium''. When In 1836, Fries first defined the genus ''
Hygrophorus ''Hygrophorus'' is a genus of agarics (gilled mushrooms) in the family Hygrophoraceae. Called "woodwaxes" in the UK or "waxy caps" (together with '' Hygrocybe'' species) in North America, basidiocarps (fruit bodies) are typically fleshy, often ...
'' in his ''Epicrisis Systematis Mycologici'', ''H. eburneus'' was included. The fungus has also been named ''Limacium eburneum'' by
Paul Kummer Paul Kummer (22 August 1834 – 6 December 1912) was a Minister (Christianity), minister, teacher, and scientist in Zerbst, Germany, known chiefly for his contribution to mycological botanical nomenclature, nomenclature. Earlier Taxonomy (biology), ...
in 1871, when he raised the tribes of Fries to the rank of genus, and ''Gymnopus eburneus'' by
Samuel Frederick Gray Samuel Frederick Gray (10 December 1766 – 12 April 1828) was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray. Background He was the son of Samuel Gray, a London Se ...
in 1821. ''H. eburneus'' is the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the genus ''Hygrophorus'', and is classified in the section ''Hygrophorus'', subsection ''Hygrophorus''. This includes species with non-
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
, smooth
spores In biology, a spore is a unit of sexual (in fungi) 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 plant ...
, and divergent
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 o ...
e in the tissue of the
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
. Other species in this subsection include '' H. eburneiformis'', '' H. coccus'', '' H. ponderatus'', '' H. chrysaspis'', and '' H. glutinosus''. The mushroom is commonly known as the "ivory waxy cap", the "white waxy cap", or the "cowboy's handkerchief". The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''eburneus'' is a
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
adjective meaning "of ivory".


Description

The
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. The origin of the word "cap" comes from the Old French word "chapeau" which means "head co ...
of ''H. eburneus'' is broad, with a shape ranging from convex to flattened, sometimes with an umbo (a raised area in the center of the cap). In age the cap margin sometimes becomes elevated and the center of the cap depressed. The cap is pure white, and depending on the moisture in the environment, may be
glutinous Domestication syndrome refers to two sets of phenotypic traits that are common to either domesticated plants or domesticated animals. Domesticated animals tend to be smaller and less aggressive than their wild counterparts; they may also have ...
to sticky. The cap surface is smooth, the margin is even and in young specimens, rolled inward and covered with short fibrils. 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 muscles, fats and other loose connective tissues, ...
is soft and white, thick in the center of the cap but thinning toward the margin. The odor and taste are mild. The
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
are somewhat arcuate-decurrent, meaning they are shaped like a bow, curving upward and then running down the stem for a short distance. In terms of spacing, they are subdistant to distant, so that space can be seen between them. The gills are moderately broad, broadest near the stem, narrowed in front, pure white, slightly yellowish or
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 wr ...
with age or when dried. The
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
is long, thick, equal in width throughout its length to somewhat tapered downward or with a greatly attenuated base, and glutinous. Its surface is silky beneath the gluten. The top of the stem is covered with short fibrils, pure white, sometimes becoming grayish or dirty with age. It is initially stuffed with cotton-like
mycelia Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...
, then later becomes hollow. The caps of dried fruit bodies will typically remain white, while the stems will dry darker, especially if they are initially waterlogged.Smith, 1947, pp. 253–55.


Microscopic characteristics

When they are viewed in mass, such as with a
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 warm orange ("tussock") color spore print. ...
, the
spores In biology, a spore is a unit of sexual (in fungi) 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 plant ...
appear white. Observing with a light microscope reveals additional details: spores are
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
, smooth, and measure 6–8 by 3.5–5 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
. They are pale yellow in
Melzer's reagent Melzer's reagent (also known as Melzer's iodine reagent, Melzer's solution or informally as Melzer's) is a chemical reagent used by mycologists to assist with the identification of fungi, and by phytopathologists for fungi that are plant pathogens ...
. The
basidia A basidium (: basidia) is a microscopic spore-producing structure found on the hymenophore of reproductive bodies of basidiomycete fungi. The presence of basidia is one of the main characteristic features of the group. These bodies are also ...
(spore-bearing cells) are 42–52 by 6-8 μm, and four-spored. There are no
pleurocystidia A cystidium (: 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 are oft ...
or
cheilocystidia A cystidium (: 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 are oft ...
. The gill tissue is made of branching
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 o ...
e about 7–12 μm wide. 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 th ...
is made of gelatinous, narrow (3–6 μm) hyphae which are repent (bent over) but typically with some erect free ends.
Clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of basidiomycete fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rece ...
s are present in the hyphae.


Similar species

A lookalike species of ''Hygrophorus eburneus'' is '' H. piceae'', which differs by having a less slimy cap, dry to slightly viscid stem, and frequent association with spruce. '' H. gliocyclus'' is just as slimy, but has a cream-colored cap, thicker stalk, and grows with pine. The "snow white waxy cap" ('' H. borealis'') is also similar in appearance, but has a smaller cap diameter of up to —and is not slimy. '' Hygrophorus cossus'', which typically grows with ''
Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
'' species, differs in its pale pinkish-buff cap and gills, and has a distinct sour odor; also, ''H. cossus'' does not have a potassium hydroxide
reaction Reaction may refer to a process or to a response to an action, event, or exposure. Physics and chemistry *Chemical reaction *Nuclear reaction *Reaction (physics), as defined by Newton's third law * Chain reaction (disambiguation) Biology and ...
on the stem as does ''H. eburneus''. '' Limacella illinita'' has non-waxy gills that are free from attachment to the stalk.Roody, 1997
p. 250
Retrieved 2010-08-25.
'' Cuphophyllus virgineus'' has decurrent, distant gills.


Habitat and distribution

The fruit bodies of ''H. eburneus'' grow on the soil, mostly in
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
ous woods, thickets and grassy areas. The fungus prefers soil that is moist, mesic,
loam Loam (in geology and soil science) is soil composed mostly of sand (particle size > ), silt (particle size > ), and a smaller amount of clay (particle size < ). By weight, its mineral composition is about 40–40–20% concentration of sand–si ...
y and
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines. In zoology ''Calcare ...
. The fungus is widely distributed in North America. It is also found in Europe (Poland and Portugal), Israel and North Africa.


Uses

The mushroom is
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 ...
, although it may not be appealing to many due to its sliminess. In China, a yak milk beverage is made with ''H. eburneus'' and
yak The yak (''Bos grunniens''), also known as the Tartary ox, grunting ox, hairy cattle, or domestic yak, is a species of long-haired domesticated cattle found throughout the Himalayan region, the Tibetan Plateau, Tajikistan, the Pamir Mountains ...
milk, by
lactic acid fermentation Lactic acid fermentation is a metabolic process by which glucose or other hexose, six-carbon sugars (also, disaccharides of six-carbon sugars, e.g. sucrose or lactose) are converted into cellular energy and the metabolite lactic acid, lactate, w ...
with ''
Lactobacillus bulgaricus ''Lactobacillus bulgaricus'' is the main bacterium used for the production of yogurt. It also plays a crucial role in the ripening of some cheeses, as well as in other processes involving naturally fermented products. It is defined as homoferment ...
'', ''
Streptococcus thermophilus ''Streptococcus thermophilus'' formerly known as ''Streptococcus salivarius ''subsp.'' thermophilus'' is a gram-positive bacteria, gram-positive bacterium, and a lactic acid fermentation, fermentative facultative anaerobic organism, facultative ...
'' and ''
Lactobacillus acidophilus ''Lactobacillus acidophilus'' (Neo-Latin 'acid-loving milk-bacillus') is a rod-shaped, Gram-positive bacteria, Gram-positive, homofermentative, Anaerobic organism, anaerobic microbe first isolated from infant feces in the year 1900. The species ...
'' as mixed starter.


Bioactive compounds

Several
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s with
bactericidal A bactericide or bacteriocide, sometimes abbreviated Bcidal, is a substance which kills bacteria. Bactericides are disinfectants, antiseptics, or antibiotics. However, material surfaces can also have bactericidal properties based solely on their p ...
and
fungicidal Fungicides are pesticides used to kill parasitic fungi or their spores. Fungi can cause serious damage in agriculture, resulting in losses of yield and quality. Fungicides are used both in agriculture and to fight fungal infections in animals, ...
activity have been isolated and identified from the fruit bodies of ''H. eburneus''. The bioactive fatty acids are built upon a chemical structure called γ-oxocrotonate. The following gamma-oxocrotonate derivatives have been identified from the mushroom: (2''E'',9''E'')-4-oxooctadeca-2,9,17-trienoic acid, (2''E'',11''Z'')-4-oxooctadeca-2,11,17-trienoic acid, (''E'')-4-oxohexadeca-2,15-dienoic acid, (''E'')-4-oxooctadeca-2,17-dienoic acid, (2''E'',9''E'')-4-oxooctadeca-2,9-dienoic acid, (2''E'',11''Z'')-4-oxooctadeca-2,11-dienoic acid, (''E'')-4-oxohexadec-2-enoic acid, and (''E'')-4-oxooctadec-2-enoic acid. The compound (''E'')-4-oxohexadec-2-enoic acid has been investigated for potential use as a fungicide against the
oomycete The Oomycetes (), or Oomycota, form a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms within the Stramenopiles. They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction o ...
species ''
Phytophthora infestans ''Phytophthora infestans'' is an oomycete or Oomycete, water mold, a fungus-like microorganism that causes the serious potato and tomato disease known as late blight or potato blight. Early blight, caused by ''Alternaria solani'', is also often c ...
'', a causal agent of potato and tomato late blight disease. Additional
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s discovered in ''H. eburneus'' include the
ceramide Ceramides are a family of waxy lipid molecules. A ceramide is composed of sphingosine and a fatty acid joined by an amide bond. Ceramides are found in high concentrations within the cell membrane of Eukaryote, eukaryotic cells, since they are co ...
compound named ''hygrophamide'' ((2''S'',3''R'',4''R'',2''R'')-2-(2'-hydroxy-9''Z''-ene-tetracosanoylamino)-octadecane-1,3,4-triol), and the β-carboline
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
s known as ''harmane'' and ''norharmane''. The report of discovery of the latter two compounds in 2008 represents their first known occurrence in fungal fruit bodies.


See also

* List of ''Hygrophorus'' species


References


Literature cited

* *


External links

* *
Mushroom Observer
Images {{Good article Edible fungi Fungi of Africa Fungi of Europe Fungi of North America eburneus Taxa named by Jean Baptiste François Pierre Bulliard Fungus species