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''Lactifluus volemus'', formerly known as ''Lactarius volemus'', is a species of
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
in the family
Russulaceae The Russulaceae are a diverse family of fungi in the order Russulales, with roughly 1,900 known species and a worldwide distribution. They comprise the brittlegills and the milk-caps, well-known mushroom-forming fungi that include some edible ...
. It is widely distributed in the
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 ...
, in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions of Europe, North America and Asia as well as some subtropical and tropical regions of Central America and Asia. 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 fungus, its
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 ...
grow on the ground at the base of various species of trees from summer to autumn, either individually or in groups. It is valued as an
edible mushroom Edible mushrooms are the fleshy and edible fruit bodies of several species of macrofungi (fungi which bear fruiting structures that are large enough to be seen with the naked eye). They can appear either below ground (hypogeous) or above ground ...
, and is sold in markets in Asia. Several other ''Lactifluus'' mushrooms resemble ''L. volemus'', such as the closely related edible species '' L. corrugis'', but these can be distinguished by differences in distribution, visible
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
, and microscopic characteristics. ''L. volemus'' produces a white
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 ...
and has roughly spherical
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, f ...
s about 7–8 
micrometre 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 Unit ...
s in diameter. The colour of the ''L. volemus'' mushroom varies from
apricot An apricot (, ) is a fruit, or the tree that bears the fruit, of several species in the genus ''Prunus''. Usually, an apricot is from the species '' P. armeniaca'', but the fruits of the other species in ''Prunus'' sect. ''Armeniaca'' are also ...
to tawny, and 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. Caps typically have a visor, or no brim at all. They are popular in casual and informal se ...
may be up to 11 cm ( in) wide. The pale golden yellow
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 ...
on the underside of the cap are closely spaced and sometimes forked. One of the mushroom's most distinctive features is the large amount of
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latexes are found in nature, but synthetic latexes are common as well. In nature, latex is found as a milky fluid found in 10% of all flowering plants (angiosperms ...
("milk") that it exudes when the gills are damaged, leading to the
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 weeping milk cap and voluminous-latex milky. It also has a distinctive fishy smell, which does not affect the taste. The fruit bodies have been chemically analysed and found to contain several
sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...
s related to
ergosterol Ergosterol (ergosta-5,7,22-trien-3β-ol) is a sterol found in cell membranes of fungi and protozoa, serving many of the same functions that cholesterol serves in animal cells. Because many fungi and protozoa cannot survive without ergosterol, the ...
, some of which are unique to this species. The mushroom also contains a
natural rubber Rubber, also called India rubber, latex, Amazonian rubber, ''caucho'', or ''caoutchouc'', as initially produced, consists of polymers of the organic compound isoprene, with minor impurities of other organic compounds. Thailand, Malaysia, and ...
that has been chemically characterized.
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 o ...
analysis suggests that ''Lactifluus volemus'' represents several species or
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 species ...
, rather than a single
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 ...
.


Taxonomy and naming

The first mention of ''Lactifluus volemus'' in the scientific literature was in
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
's 1753 ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
'', under the name ''Agaricus lactifluus''. In 1821, Swedish mycologist
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. Career Fries was born at Femsjö (Hylte Municipality), Småland, the son of the pastor there. He attended school in Växjö. He acquired ...
called it ''Agaricus volemus'' in his ''
Systema Mycologicum Systema Mycologicum is a systematic classification of fungi drawn up in 1821 by the Swedish mycologist and botanist Elias Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. Career F ...
''. In this work he proposed a grouping of related species (called a ''tribus'', or tribe) within the genus ''Agaricus'', which he named ''Galorrheus''. Fries later recognised ''Lactarius'' as a distinct genus in his 1838 ''Epicrisis Systematis Mycologici'', citing ''Galorrheus'' as a
synonym 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 ...
. Although Linnaeus had published the species before Fries, Fries's name is sanctioned and thus has nomenclatural priority. In 1871
Paul Kummer Paul Kummer (22 August 1834 – 6 December 1912) was a minister, teacher, and scientist in Zerbst, Germany, known chiefly for his contribution to mycological nomenclature. Earlier classification of agarics by pioneering fungal taxonomist Elias Ma ...
raised most of Fries's tribes to generic rank, and so renamed the species ''Galorrheus volemus''. 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), ...
''L. volemus'' var. ''subrugosus'' was identified by
Charles Horton Peck Charles Horton Peck (March 30, 1833 – July 11, 1917) was an American mycologist of the 19th and early 20th centuries. He was the New York State Botanist from 1867 to 1915, a period in which he described over 2,700 species of North American fun ...
in 1879, but is now classified as a separate species, '' L. corrugis''. In 1891,
Otto Kuntze Carl Ernst Otto Kuntze (23 June 1843 – 27 January 1907) was a German botanist. Biography Otto Kuntze was born in Leipzig. An apothecary in his early career, he published an essay entitled ''Pocket Fauna of Leipzig''. Between 1863 and 1866 he ...
moved the species into ''Lactifluus'', which was afterwards long considered a synonym of ''Lactarius'' but confirmed as a separate genus through
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
in 2008 and subsequent taxonomical rearrangements within the family Russulaceae. Another historical synonym is ''Lactarius lactifluus'', used by
Lucien Quélet Lucien Quélet in 1869 Lucien Quélet (14 July 1832 – 25 August 1899) was a French naturalist and mycologist. Quélet discovered several species of fungi and was the founder of the Société mycologique de France, a society devoted to mycolo ...
in 1886, a renaming based on Linnaeus's ''Agaricus lactifluus''. ''Lactarius wangii'', reported by Hua-An Wen and Jian-Zhe Ying to be a new species from China in 2005, was synonymised two years later with ''L. volemus''. 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 ...
"''volemus''" is derived from the
Latin Latin (, or , ) is a classical language belonging to the 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 the power of the ...
''vola'', meaning "the hollow of the hand", suggestive of Fries's reference to the large amount of latex "flowing enough to fill the hand".
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 for ''L. volemus'' include the weeping milk cap, the tawny milkcap, the orange-brown milky, the voluminous-latex milky, the lactarius orange, the fishy milkcap, and the apricot milk cap. In the
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the Bur ...
n mountains of the United States, the mushroom is called a "leatherback" or a "bradley". The latter name may originate from its German name ''Brätling''.Bessette ''et al''. (2009), p. 5.


Phylogeny

''Lactifluus volemus'' was the
type species In 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 species that contains the biological type specimen ...
of the
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 ...
''Dulces'' in
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
''Lactarius'', and is currently classified in ''Lactifluus'' section ''Lactifluus''. The group around this ''L. volemus'' includes species with a dry cap, abundant latex, and a white or pale cream
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 ...
. Because the closely related ''L. corrugis'' has overlapping morphological characters, including similar colouration in the cap and stem, it has been difficult to reliably distinguish between the two species. The difficulty in discerning the two is exacerbated by the fact that both species have several colour forms: Japanese specimens of ''L. volemus'' may have a red cap, a yellow cap with a long stem, or a velvet-like surface texture; the caps of ''L. corrugis'' may be either red, commonly
rust Rust is an iron oxide, a usually reddish-brown oxide formed by the reaction of iron and oxygen in the catalytic presence of water or air moisture. Rust consists of hydrous iron(III) oxides (Fe2O3·nH2O) and iron(III) oxide-hydroxide (FeO(OH ...
-coloured. In 2005, Japanese researchers clarified the relationships between these two species and others in section ''Dulces'' using
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, and by comparing differences in
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, fr ...
composition,
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
, and taste. The colour variants group
phylogenetically 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 o ...
into different
subclades In genetics, a subclade is a subgroup of a haplogroup. Naming convention Although human mitochondrial DNA (mtDNA) and Y chromosome DNA (Y-DNA) haplogroups and subclades are named in a similar manner, their names belong to completely separate syst ...
, suggesting that they might better be considered as "different species, subspecies, or varieties". A 2010 molecular study of ''L. volemus'' of northern Thailand found that 79 tested specimens could be divided into 18 distinct phylogenetic species; six of these were described as new species: '' Lactifluus acicularis'', '' L. crocatus'', '' L. distantifolius'', '' L. longipilus'', '' L. pinguis'' and '' L. vitellinus''.


Description


Macromorphology

The
fruit body In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the Ovary (plants), ovary after flowering plant, flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their ...
of ''Lactifluus volemus'' has a fleshy and firm
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 ...
with a velvety or smooth surface and a shape that changes with maturity: it starts off convex, with edges curved inwards, then later grows flat with a depression in the middle. With a typical diameter of 5 to 11 cm (2– in), its colour ranges from apricot to tawny. The cap colouration, however, is somewhat variable, as has been noted in Asian, European, and North American specimens. The
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
, whose height varies between 4 and 12 cm ( and 5 in), and which is typically between thick, has a slightly lighter colouration than the cap. It is firm, with a velvety or smooth surface that sometimes has depressions running longitudinally up and down its length. 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
adnate Adnate may refer to: * Adnation, in botany, the fusion of two or more whorls of a flower * Adnate, in mycology, a classification of lamellae (gills) * Conjoined twins Conjoined twins – sometimes popularly referred to as Siamese twins – are ...
to slightly
decurrent ''Decurrent'' (sometimes decurring) is a term used in botany and mycology to describe plant or fungal parts that extend downward. In botany, the term is most often applied to leaf blades that partly wrap or have wings around the stem or petio ...
, brittle, narrow, quite closely spaced and sometimes forked. Normally a pale golden yellow colour, the gills turn brown when bruised. Interspersed between the gills are lamellulae, short gills that do not extend to the stem. 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 whitish and firm. The mushroom smells somewhat fishy; one source suggests the odour is "like a dead
shad The Alosinae, or the shads,Alosinae
, which anglers will tell you is probably the most malodorous freshwater fish". The odour is concentrated when the fruit bodies are dried. One of the mushroom's most distinctive features is the abundant
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latexes are found in nature, but synthetic latexes are common as well. In nature, latex is found as a milky fluid found in 10% of all flowering plants (angiosperms ...
, so plentiful that a small nick on the gills will cause it to "weep" the milky substance. The latex tends to impart brown stains on whatever it contacts.


Micromorphology

The
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 ...
is whitish. 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, f ...
s are roughly spherical, translucent (
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
), and typically measure 7.5–10.0 by 7.5–9.0 µm. The spore surface is
reticulate Reticulation is a net-like pattern, arrangement, or structure. Reticulation or Reticulated may refer to: * Reticulation (single-access key), a structure of an identification tree, where there are several possible routes to a correct identificatio ...
—covered with ridges that form a complete network. The ridges are up to 0.8 µm high and have conspicuous projections up to 1.2 µm high. The spore-bearing cells 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 others some ...
, 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 ...
, are club-shaped, hyaline, four-spored, and have dimensions of 40–62 by 7.2–10.4 µm. Interspersed among the basidia are sterile cells called
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 ...
. The pleurocystidia (cystidia on the side of a gill) are roughly spindle- to club-shaped, and measure 48–145 by 5–13 µm. The cheilocystidia (cystidia on the edge of a gill) may be spindle-, club-, or awl-shaped (
subulate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
), or intermediate in between these forms, and measure 27–60 by 5–7 µm. Additionally, there are cystidia present on both the surface of the cap and the stem. If a drop of ferric sulphate (used as a chemical test in mushroom identification) is applied to the mushroom flesh, it will immediately stain dark bluish-green.Bessette ''et al''. (2009), pp. 264–66.


Varieties

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), ...
''Lactifluus volemus'' var. ''flavus'' was 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
Lexemuel Ray Hesler Lexemuel Ray Hesler (20 February 1888 – 20 November 1977) was an American mycologist. He was the son of Clinton F. Hesler and Laura Iris (née Youngblood). He obtained his B.A. degree with Wabash College in 1911 and his Ph.D. at the University ...
in their 1979
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
of North American ''Lactarius'' species. This rare variety, found in the southeastern United States (ranging from
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
to
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
and extending west to
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
), has a cap that stays yellow throughout its development. It also has slightly smaller spores than the regular variety: 6.5–9.0 by 6–8 µm. It is also a good edible. Some authors have considered the rarely collected ''L. volemus'' var. ''oedematopus'', found in central and southern Europe, to be a distinct variety distinguished from the common variety by a darker reddish-brown cap and a swollen stem. This assessment is not universally accepted, possibly because it falls within the range of morphological variation shown by the main variety. ''L. volemus'' var. ''asiaticus'' was named in 2004 based on Vietnamese specimens; associating with Khasi pine (''Pinus khasya''), it has small, dull brown, velvety fruit bodies. In general, little taxonomical significance has been ascribed to the several varieties of ''L. volemus'' that have been proposed.


Similar species

''Lactifluus volemus'' is closely related to ''L. corrugis'', and generally similar in appearance. ''L. corrugis'' usually has more surface wrinkles, darker gills, weaker or absent scent, and less orange colouration; however, intermediate colour forms can be found. The two can be distinguished more definitively by microscopic characteristics: ''L. corrugis'' has larger spores—typically 10.4–12.8 by 9.6–11.8 µm—with a coarser surface reticulum, and larger pleurocystidia. The species '' Lactifluus austrovolemus'' is closely related, but has more crowded gills, while '' L. lamprocystidiatus'' can only be reliably distinguished from ''L. volemus'' by microscopic characteristics: the reticulations on its spores are taller and more acute, and the meshes formed by the intersections of the reticulations are smaller. Both ''L. austrovolemus'' and ''L. lamprocystidiatus'' are known only from
Papua New Guinea Papua New Guinea (abbreviated PNG; , ; tpi, Papua Niugini; ho, Papua Niu Gini), officially the Independent State of Papua New Guinea ( tpi, Independen Stet bilong Papua Niugini; ho, Independen Stet bilong Papua Niu Gini), is a country i ...
. '' Lactifluus hygrophoroides'' also resembles ''L. volemus'', but differs in having widely spaced gills, and spores that lack surface reticulations. Some species of the genus ''Lactarius'' are also similar: The tropical African '' Lactarius chromospermus'' has a superficial resemblance to ''L. volemus'', but the former species, in addition to its African distribution, can be identified by its cinnamon-brown spore print—unique in the
Russulaceae The Russulaceae are a diverse family of fungi in the order Russulales, with roughly 1,900 known species and a worldwide distribution. They comprise the brittlegills and the milk-caps, well-known mushroom-forming fungi that include some edible ...
. '' Lactarius subvelutinus'' is also similar to ''L. volemus'', but lacks the fishy odour, has a dull yellow-orange to bright golden orange cap, narrow gills, and a white latex that does not change colour.


Edibility and other uses

Despite the unappealing fishy scent that develops after the mushroom is picked, ''Lactifluus volemus'' 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 ...
and recommended for culinary usage, though, typical of milk caps, it has a slightly granular texture that some may find unappetizing. The odor disappears during cooking. The latex only has a mild taste. The species is considered good for novice mushroom hunters to eat, and is best prepared by slow cooking to prevent it from becoming too hard; specimens that have been rehydrated after having been dried will require longer cooking times to eliminate the grainy texture. The mushroom has also been suggested for use in
casserole A casserole ( French: diminutive of , from Provençal 'pan') is a normally large deep pan or bowl a casserole is anything in a casserole pan. Hot or cold History Baked dishes have existed for thousands of years. Early casserole recipes ...
s and thick sauces.
Pan frying Pan frying or pan-frying is a form of frying food characterized by the use of minimal cooking oil or fat (compared to shallow frying or deep frying), typically using just enough to lubricate the pan. In the case of a greasy food such as bacon, n ...
is not a recommended cooking technique, due to the large amounts of latex it exudes. ''L. volemus'' is one of several species of milk caps that are sold in rural markets in
Yunnan Province Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the C ...
, China, and it is among the most popular wild edible mushroom species collected for consumption and sale in Nepal. In their 2009 book on milk caps of North America, Bessette and colleagues consider the mushroom "the best-known and most popular edible milk mushroom" in the eastern United States. A Turkish study of the nutritional composition of the fruit bodies concluded that ''L. volemus'' is a good source of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
and
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s. Two elderly people developed a transient pancreatitis after consuming ''L. volemus'' in central
Anatolia 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 ...
in Turkey. Both had eaten the mushroom, which they knew as ''Tirmit'', many times before. The condition resolved spontaneously.


Bioactive compounds

Fruit bodies contains a unique
sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...
molecule called volemolide, a
derivative In mathematics, the derivative of a function of a real variable measures the sensitivity to change of the function value (output value) with respect to a change in its argument (input value). Derivatives are a fundamental tool of calculus. F ...
of the common fungal sterol
ergosterol Ergosterol (ergosta-5,7,22-trien-3β-ol) is a sterol found in cell membranes of fungi and protozoa, serving many of the same functions that cholesterol serves in animal cells. Because many fungi and protozoa cannot survive without ergosterol, the ...
that may have application in fungal
chemotaxonomy Webster's Dictionary, Merriam-Webster defines ''chemotaxonomy'' as the method of biology, biological classification based on similarities and dissimilarity in the structure of certain chemical compound, compounds among the organisms being classifi ...
. A 2001 study identified a further nine sterols, three of which were previously unknown to science. According to the authors, these types of highly
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
ated compounds—similar to sterols found in marine
soft coral Alcyonacea, or soft corals, are an order of corals. In addition to the fleshy soft corals, the order Alcyonacea now contains all species previously known as "gorgonian corals", that produce a more or less hard skeleton, though quite different f ...
and
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s—are rare in fungi. The mushroom also contains volemitol (D-glycero-D-mannoheptitol), a seven-carbon
sugar alcohol Sugar alcohols (also called polyhydric alcohols, polyalcohols, alditols or glycitols) are organic compounds, typically derived from sugars, containing one hydroxyl group (–OH) attached to each carbon atom. They are white, water-soluble solids ...
first isolated from the species by the French scientist Émile Bourquelot in 1889. Volemitol occurs as a
free sugar Free may refer to: Concept * Freedom, having the ability to do something, without having to obey anyone/anything * Freethought, a position that beliefs should be formed only on the basis of logic, reason, and empiricism * Emancipate, to procu ...
in many plant and
brown alga Brown algae (singular: alga), comprising the class (biology), class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the ...
l species. Due to their natural
polyisoprene Polyisoprene is strictly speaking a collective name for Polymer, polymers that are produced by polymerization of isoprene. In practice polyisoprene is commonly used to refer to synthetic ''cis''-1,4-polyisoprene, made by the industrial polymerisa ...
content (1.1–7.7% by
dry weight Vehicle weight is a measurement of wheeled motor vehicles; either an actual measured weight of the vehicle under defined conditions or a gross weight rating for its weight carrying capacity. Curb or kerb weight Curb weight (U.S. English) or kerb ...
of fruit bodies), ''L. volemus'' fruit bodies can also be used to produce
rubber Rubber, also called India rubber, latex, Amazonian rubber, ''caucho'', or ''caoutchouc'', as initially produced, consists of polymers of the organic compound isoprene, with minor impurities of other organic compounds. Thailand, Malaysia, and ...
. The chemical structure of rubber from the mushroom consists of a high
molecular mass The molecular mass (''m'') is the mass of a given molecule: it is measured in daltons (Da or u). Different molecules of the same compound may have different molecular masses because they contain different isotopes of an element. The related quanti ...
homologue of
polyprenol Polyprenols are natural long-chain isoprenoid alcohols of the general formula H-(C5H8)n-OH where n is the number of isoprene units. Any prenol with more than 4 isoprene units is a polyprenol. Polyprenols play an important function acting as natura ...
, arranged as a dimethylallyl group, two ''trans''
isoprene Isoprene, or 2-methyl-1,3-butadiene, is a common volatile organic compound with the formula CH2=C(CH3)−CH=CH2. In its pure form it is a colorless volatile liquid. Isoprene is an unsaturated hydrocarbon. It is produced by many plants and animals ...
units, a long sequence of ''cis'' isoprenes (between 260 and 300 units), terminated by a
hydroxyl In chemistry, a hydroxy or hydroxyl group is a functional group with the chemical formula and composed of one oxygen atom covalently bonded to one hydrogen atom. In organic chemistry, alcohols and carboxylic acids contain one or more hydroxy ...
or
fatty acid ester Fatty acid esters (FAEs) are a type of ester that result from the combination of a fatty acid with an alcohol. When the alcohol component is glycerol, the fatty acid esters produced can be monoglycerides, diglycerides, or triglycerides. Dietary f ...
. Biosynthetically, the creation of the polyisoprene begins with the compound ''trans'',''trans''-
farnesyl pyrophosphate Farnesyl pyrophosphate (FPP), also known as farnesyl diphosphate (FDP), is an intermediate in the biosynthesis of terpenes and terpenoids such as sterols and carotenoids. It is also used in the synthesis of CoQ (part of the electron transport cha ...
, and is thought to terminate by
ester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides ar ...
ification of polyisoprenyl pyrophosphate. The enzyme isopentenyl-diphosphate delta isomerase has been identified as required for the initiation of rubber synthesis in ''L. volemus'' and several other milk cap species.


Ecology, distribution, and habitat

Like all milk caps,Bessette ''et al''. (2009), p. 4. ''L. volemus'' forms mycorrhiza, ectomycorrhizae, mutually beneficial symbiosis, symbiotic associations with various tree species. In this association, the fungal hyphae grow around the root of the plant and between its cortex (botany), cortical cells, but do not actually penetrate them. The hyphae extend outward into the soil, increasing the surface area for absorption to help the plant absorb nutrients from the soil. It is found growing at the base of both coniferous and deciduous, broad-leaved trees, although it is more common in deciduous woods. It may also sometimes be found in sphagnum, peat moss beds. The fruit bodies, which appear between summer and autumn, are common. They can be found growing solitarily or in groups, and are more abundant in weather that is warm and humid. Fruit bodies can be inhabited by species of Limoniidae, limoniid flies, such as ''Discobola marginata'' or ''Limonia yakushimensis'', as well as several species of fungi-dwelling mites. The flies are hosts for the mites in a symbiosis, symbiotic association known as ''phoresis'', whereby the mites are mechanically carried by its host. Mites are small and unable to migrate the relatively long distances between mushrooms without assistance; the insect hosts, in comparison, are large and can transfer the mites between their preferred feeding habitats. ''Lactifluus volemus'' is found in warm
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions and as well as some subtropical and tropical regions of the Northern Hemisphere. The fungus is widely distributed throughout Europe, although it is in decline in some countries, and has become rare enough in the Netherlands (and Flanders) to be considered local extinction, locally extinct. In the Americas, the northern limit of its distribution reaches southern Canada east of the Great Plains, and the species extends south to the East Coast of the United States and Mexico, and beyond into Central America (Guatemala). It is also known from Asia, including China (Qinling Mountains, Guizhou Province, and
Yunnan Province Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the C ...
), Japan, India, Korea, Nepal, and Vietnam. Collections have also been made from the Middle East, including Iran and Turkey.


See also

*List of Lactifluus species, List of ''Lactifluus'' species


References


Cited text

*


External links

* *
Photos
at Mushroom Observer {{featured article Edible fungi Fungi described in 1753 Taxa named by Carl Linnaeus Fungi of Asia Fungi of Central America Fungi of Europe Fungi of North America Lactifluus, volemus Taxa named by Elias Magnus Fries