Lactarius torminosus - Wikipedia https://en.wikipedia.

org/wiki/Lactarius_torminosus

Lactarius torminosus

Lactarius torminosus, commonly known as the woolly

milkcap or the bearded milkcap, is a large agaric fungus. A Lactarius torminosus
common and widely distributed species, it is found in North
Africa, northern Asia, Europe, and North America. It was first
described scientifically by Jacob Christian Schäffer in 1774 as an
Agaricus, and later transferred to the genus Lactarius in 1821
by Samuel Frederick Gray. A variety, L. torminosus var.
nordmanensis, is known from the United States, Canada, and
Switzerland. L. torminosus officially became the type species of
Lactarius in 2011 after molecular studies prompted the
taxonomic reshuffling of species between several Russulaceae
genera.
Scientific classification
A mycorrhizal species, L. torminosus associates with various
trees, most commonly birch, and its fruit bodies (mushrooms) Kingdom: Fungi
grow on the ground singly or in groups in mixed forests. The Division: Basidiomycota
caps of L. torminosus mushrooms are convex with a central
depression, and attain a diameter of up to 10 cm (3.9 in). A Class: Agaricomycetes
blend of pink and ochre hues, the cap sometimes has concentric Order: Russulales
zones of alternating lighter and darker shades. The edge of the
Family: Russulaceae
cap is rolled inward, and shaggy when young. On the underside
of the cap are narrow flesh-colored gills that are crowded closely Genus: Lactarius
together. The cylindrical stem is a pale flesh color with a
Species: L. torminosus
delicately downy surface and brittle flesh; it is up to 8 cm (3.1 in)
long and 0.6–2 cm (0.2–0.8 in) thick. When cut or injured, the Binomial name
fruit bodies ooze a bitter white latex that does not change color
Lactarius torminosus
upon exposure to air. The variety nordmanensis, in contrast, has
latex that changes from white to yellow. Lactarius torminosus (Schaeff.) Gray (1821)
can be distinguished from similar species like L. pubescens or Synonyms[1]
L. villosus by differences in morphology and coloration, or by
microscopic characteristics like spore shape and size.
Agaricus torminosus Schaeff.
Although it is valued for its peppery flavor and eaten after (1774)
suitable preparation in Russia and Finland, the species is highly Lactarius necator (Bull.) Pers.
irritating to the digestive system when eaten raw. The toxins, (1800)
also responsible for the strongly bitter or acrid taste, are Galorrheus torminosus (Schaeff.)
destroyed by cooking. Studies have identified several chemicals P.Kumm. (1871)
present in the mushrooms, including ergosterol and derivatives Lactifluus torminosus (Schaeff.)
thereof, and the pungent-tasting velleral. Kuntze (1891)
Lactarius torminosus var.
sublateritius Kühner & Romagn.
(1954)
Contents

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Taxonomy and phylogeny Lactarius torminosus

Description Mycological characteristics
Microscopic characteristics
gills on hymenium
Variety nordmanensis
Similar species cap is convex or
Edibility and toxicity depressed

Chemistry hymenium is decurrent

Ecology and distribution stipe is bare

See also
spore print is cream to
References
yellow
Cited texts
ecology is mycorrhizal

edibility: edible but not

Taxonomy and phylogeny recommended

German naturalist Jacob Christian Schäffer was the first to describe the species, placing it in Agaricus
in 1774.[2] Seven years later in 1781, Jean Bulliard described a species he called Agaricus necator and
illustrated it in the first volume of his Herbier de la France;[3] this name and the synonym Lactarius
necator, resulting from Christian Hendrik Persoon's 1800 transfer[4] to Lactarius, are both
considered to refer to L. torminosus.[5] Otto Kuntze, for his part chose to put it in Lactifluus,[6] while
Paul Kummer thought Galorrheus[7] was the appropriate placement; until the recent resurrection of
Lactifluus, both genera had been long considered to be unnecessary segregates of Lactarius.[8]
According to Index Fungorum,[1] another synonym is Samuel Frederick Gray's[9] Lactarius necans.
Gray also gave the species its modern name when he transferred it to Lactarius in his 1821 Natural
Arrangement of British Plants.[1][9]

The specific epithet torminosus means "tormenting" or "causing colic", in reference to the
gastrointestinal distress associated with consuming the raw mushroom.[10] Early English vernacular
names were Gray's "bellyach milk-stool" (1821),[9] and James Edward Smith's "bearded pepper agaric"
(1824).[11] More recent common names include "shaggy milkcap",[12] "powderpuff milkcap",[13] "pink-
fringed milkcap",[14] "bearded milkcap",[15] and the British Mycological Society-recommended "woolly
milkcap".[16]

According to Hesler and Smith's 1979 classification of the genus Lactarius, L. torminosus belongs to
subgenus Piperites, section Piperites (in which it is the type species), subsection Piperites. Species in
this subsection are characterized by having latex that does not turn yellow after exposure to air,
and/or that does not stain the cut surface of the mushroom surface yellow.[17] A 2004 phylogenetic
analysis of European Lactarius species concluded that L. torminosus falls into a group that includes
L. torminosulus, and that these two species are closely related to a group that includes L. tesquorum,
L. scoticus, and L. pubescens.[18]

A multi-gene molecular analysis published in 2008 demonstrated that species then distributed in the
genera Lactarius and Russula actually consisted of four distinct lineages.[19] The subsequent
reorganization of Russulaceae species—a taxonomic change needed to make Russula and Lactarius
monophyletic—required that a new type species be defined for Lactarius, since the previous type,

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L. piperatus, belonged to the clade that will be transferred to genus Lactifluus. A proposal to conserve
Lactarius with L. torminosus as the type was accepted by the Nomenclatural Committee for Fungi[20]
and passed at the 2011 International Botanical Congress.[21] The change minimizes "taxonomic
disruption", allowing most of the common and well-known Lactarius species to retain their names.[8]

Description
The cap is initially convex, but as it matures the center forms a
depression and the outer edges rise until it assumes the shape of a
shallow funnel; its final width is typically between 2 and 12 cm (0.8
and 4.7 in).[22] The cap margin is strongly curled inward; when
young, it is tomentose (covered with a thick matting of hairs),
forming a veil-like structure that partly covers up the gills.[15] This
tomentum diminishes with age. The cap surface is at first similarly
tomentose, but eventually, the hairs wear off, leaving the surface
Young fruit body showing the more or less smooth.[22] The surface starts off somewhat sticky with
tomentose cap margin and clear concentric rings of darker shade (a zonate pattern); these rings,
forked gills especially the outer ones, usually fade in maturity.[13] The cap color
is pinkish-orange to pale dull pink, becoming orange to whitish
toward the margin as the pink gradually fades. The white to flesh-
colored flesh is firm and brittle, but becomes flaccid in age. The latex that is produced when the
mushroom tissue is cut or injured is white to cream, and does not change color with prolonged
exposure to air, nor does it stain the gills. It has an acrid taste, with a slight to pungent odor.[22]

The gills are subdecurrent (running only a small way down the length of the stem), close to crowded
together, narrow, and sometimes forked near the stem. Their color is whitish, becoming pink-tinged,
turning pale tan with age. The adult stem is 1.5–8 cm (0.6–3.1 in) long, 0.6–2 cm (0.2–0.8 in) thick,
fragile, more or less equal in width throughout, and cylindrical or narrowed at the base. Its surface is
dry, and either smooth to pruinose (covered with a very fine whitish powder on the surface). The color
ranges from pale light pink to yellowish-tinged or slightly pinkish-orange to orange-white, sometimes
spotted. The interior of the stem is firm, beige white, and filled with a soft pith, but it eventually
becomes hollow.[22] Occasionally, white mycelium is visible at the base of the stem where it meets the
ground.[23]

The fruit body formation of L. torminosus is pileostipitocarpic. In this type of development, the
hymenium forms early on the underside of the cap and upper stem of the mushroom primordium. As
the cap enlarges, the margin, made of flaring filamentous hyphae that grow outward and downward,
tends to curve inward, eventually forming a flap of tissue roughly parallel to the stem surface. As
further development takes place, these hyphae make contact with and adhere to the hymenial surface
of the stem, covering basidia and macrocystidia (very long cystidia) already present. The junction
between the two tissues produces a cavity that provides some temporary protection to the basidia,
although they are already fertile when the cap margin starts to grow.[24]

Microscopic characteristics

The spore print of L. torminosus is cream to pale yellow, and the spores 8–10.2 by 5.8–6.6 μm,
roughly spherical to broadly elliptical in side view, and hyaline (translucent). Only the ornamentation
on their surface is amyloid; it is partially reticulate (network-like) with interrupted ridges roughly

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0.5–0.7 μm high, and a few isolated warts. Spores have a

conspicuous apiculus, demarcating where it was once attached to
the basidia via the sterigma. The basidia are four-spored, hyaline
and club-shaped to cylindrical, measuring 30–47.7 by
7.3–8.2 μm.[22]

Pleurocystidia are present only in the form of macrocystidia

embedded and originating in the hymenium and just below it,
they reach 40.3–80.0 by 5.1–9.5 μm. Macrocystidia are abundant
in the hymenium. Characterized by their spindle-shaped to
ventricose (swollen on one side) form that gradually tapers in
width, they have granular hyaline contents. Gill edge cystidia
(cheilocystidia) are smaller: 30–52 by 4.5–8.0 μm. The cap Although the cap surface is typically
cuticle is made of gelatinized, interwoven hyphae arranged more zonate—marked by concentric
or less parallel to the cap surface (a form known as ixocutis); the colored bands—in young
thin-walled, threadlike hyphae of this layer are 2.5–7.3 μm specimens ...
wide.[22]

Variety nordmanensis

Lactarius nordmanensis was described by Alexander Smith in

1960 to account for a North American species closely resembling
L. torminosus in appearance, but with a slightly larger range of
spore sizes (9–11 by 6.5–8 μm). In contrast to the unchanging
latex of L. torminosus, L. nordmanensis has whitish latex that ... this feature is less prominent in
slowly changes to pale yellow upon exposure to air; the latex also mature mushrooms.
stains mushroom tissues and paper yellow. [25] Hesler and Smith
reduced L. nordmanensis to the status of a variety under
L. torminosus in 1979.[26] Lactarius torminosus var. nordmanensis has been recorded from
California,[27] Idaho, Michigan, and Wisconsin in the United States, Quebec in Canada, and
Switzerland. The variety resembles Lactarius pubescens var. betulae, but differs in its longer
pleurocystidia, larger spores with slightly different spore ornamentation, and strongly burning acrid
taste.[26] The holotype specimen of L. torminosus var. nordmanensis was collected by Smith in 1956
near Nordman, Idaho.[28]

Similar species

The woolly cap margin,

pinkish tones in the cap,
acrid latex, and association
with birch are reliable field
characteristics to help
identify L. torminosus.[15]
However, there are several
hairy Lactarius species with
which it is often confused, Lactarius pubescens Lactarius mairei
and sometimes examination
of microscopic characteristics

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is necessary to distinguish between them. The closely related L. torminosulus is a dwarf version of
L. torminosus, an arctic species associated with the birches Betula nana or B. glandulosa.[29]
Immature fruit bodies of L. scrobiculatus resemble L. torminosus, but they have a white latex that
soon turns yellow upon exposure to air, and their stems have shiny depressed spots. The caps of the
poorly known species L. cilicioides are not zonate, and its spores are smaller. L. pubescens is
physically quite similar, but can be distinguished by its paler color and smaller spores (6.0–7.5 by
5.0–6.5 μm).[22] L. controversus has a cap margin that is not as hairy, whitish to cream-colored gills,
and larger spores measuring 7.5–10 by 6–7.5 μm.[30] L. mairei has a coloration similar to
L. torminosus, but is rarer and typically found associated with oak trees on calcareous soil.[31] Known
only from North Carolina and western Canada,[32] L. subtorminosus was named for its similarity to
L. torminosus. It can be distinguished by its mild-tasting latex and smaller, roughly spherical spores
measuring 5.5–7 by 5.5–6.5 μm.[33][34]

Edibility and toxicity

The intensely peppery taste of the raw mushroom can blister the
tongue if sampled in excess.[14] Some authors have reported the
species as outright poisonous,[10][36] or causing "mild to fatal
gastroenteritis".[37] In a 1930 publication, Hans Steidle reported
that although the mushroom was not toxic to "unicellular and A comparison between Lactarius
cold-blooded organisms" when ingested, the liquid extract and the torminosus (left) and Lactarius
pressed juice of the fruit bodies, when injected under the skin of a pubescens (right)
frog, resulted in disturbed breathing, paralysis, and eventually
death.[38] Symptoms that are typically experienced after
consuming raw mushrooms include nausea, vomiting, and severe
"Its taste is biting, worse
diarrhea starting about one hour after ingestion. This combination can than Cayenne pepper ...
lead to dehydration, muscle spasms, and circulatory collapse. The Unless he possessed a
gastroenteritis will usually resolve without treatment in a couple of stomach built for the
days. [39] purpose, the man who
touched such food as this
would have a singularly
Despite these reports of toxicity, L. torminosus mushrooms are prepared bad time before him."
in Finland, Russia, and other northern and eastern European countries by
parboiling, soaking in brine for several days, or pickling, after which they Jean-Henri Fabre[35]
are valued for its peppery taste. In Norway, it is roasted and added to
coffee.[15][31] Mushrooms are harvested for commercial sale in
Finland.[40] The nutrient composition of Finnish specimens has been
analyzed and found to contain the following components (as a percentage of dry weight): protein,
17.20%; phosphorus, 0.46%; calcium, 0.12%; magnesium, 0.09%; potassium, 2.97%; sodium
0.01%.[41]

Chemistry
The compound thought to be responsible for the toxicity of raw L. torminosus is the pungent-tasting
velleral present at a concentration of 0.16 mg/g mushroom. Velleral is a breakdown product of stearyl-
velutinal. Broken lactifers—specialized hyphal cells that produce the mushroom's latex—leak the
precursor chemicals whose breakdown products act as the defensive agents toxic to humans,
effectively deterring certain vertebrates that might consume the mushroom.[42] The lactarane-type
sesquiterpene lactone 15-hydroxyblennin A is one of several sesquiterpenes produced by the species.

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Other lactaranes are found in various Lactarius species, such as

blennin A in L. deliciosus and L. blennius, and lactarorufin N in
Lactarius rufus.[43] Fungal sesquiterpenes are commonly
produced as toxins to defend against predation, and as a result
some have chemical properties that may have applications in
medicinal chemistry.[44]

Fruit bodies of Lactarius torminosus contain a number of sterols,

of which ergosterol (a component of fungal cell walls) is the most Structure of the sesquiterpene
predominant at 60.5% of all sterols, followed by its derivatives velleral
and ergosta-5/7-dien-3-ol (17.0%), ergost-7-en-3-ol (13.7%) and
ergosta-7-22-dien-3-ol (8.3%).[45] Researchers have identified 28
volatile compounds that contribute to the odor of the mushroom. Many of these are alcohols and
carbonyl compounds with eight carbon atoms; the predominant volatile compound (about 90%) is
1-octen-3-one,[46] an odorant common in mushrooms.[47]

Ecology and distribution

Lactarius torminosus is a mycorrhizal species, and as
such plays an important role in facilitating nutrient
and water uptake by trees. It grows in association with
birch (Betula) and hemlock (Tsuga) in mixed forests.
It is also known to grow in urban settings when birch
trees are nearby.[22] A field study in Scotland
concluded that the species is more likely to be present
in older than in younger birch woodlands.[48] Fruit
In Västerbotten, Sweden bodies grow on the ground, scattered or grouped
together. They are a component of the diet of the red
squirrel,[41] and serve as breeding sites for some
fungus-feeding flies in the Drosophilidae and Mycetophilidae families.[49] Lactarius torminosus
mushrooms may be parasitized by the mold Hypomyces lithuanicus, which produces a cream-ochre to
cinnamon-colored granular or velvety growth of mycelium on the surfaces of the gills and causes them
to be deformed.[50]

The species is found in northern temperate and boreal climates, penetrating sometimes into subarctic
regions. It has been recorded from North Africa, northern Asia,[31] Europe, and is common in North
America, where it sometimes grows with aspen (Populus species).[15] The North American distribution
extends north into the Yukon and Alaska and south to Mexico.[51][52]

See also
List of deadly fungi
List of Lactarius species

References
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222358/http://www.indexfungorum.org/Names/SynSpecies.asp?RecordID=228133). Index

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Cited texts
Bessette AR, Bessette AE, Harris DB (2009). Milk Mushrooms of North America: A Field Guide to
the Genus Lactarius. Syracuse, New York: Syracuse University Press. ISBN 978-0-8156-3229-0.
Hesler LR, Smith AH (1979). North American Species of Lactarius. Ann Arbor, Michigan: The
University of Michigan Press. ISBN 978-0-472-08440-1.

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