Lactarius torminosus

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Lactarius torminosus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Russulales
Family: Russulaceae
Genus: Lactarius
Species:
L. torminosus
Binomial name
Lactarius torminosus
(Schaeff.) Gray (1821)
Synonyms[1]
List
  • Agaricus torminosus Schaeff. (1774)
  • Galorrheus torminosus (Schaeff.) P.Kumm. (1871)
  • Lactifluus torminosus (Schaeff.) Kuntze (1891)
  • Agaricus intermedius sensu Krombholz (1887)
  • Lactarius intermedius sensu Krombholz (1887)
  • Agaricus cilicioides Fr. (1821)
  • Lactarius cilicioides (Fr.) Fr. (1838)
  • Galorrheus cilicioides (Fr.) P. Kumm. (1871)
  • Lactarius torminosus var. cilicioides (Fr.) Quél. (1886)
  • Lactifluus cilicioides (Fr.) Kuntze (1891)
  • Lactarius torminosus subsp. cilicioides (Fr.) Konrad & Maubl. (1935)
  • Lactarius intermedius Krombh. ex Berk. & Broome (1881)
  • Lactarius cilicioides var. albus Peck (1885)
  • Lactarius cilicioides subsp. intermedius Krombh. ex Sacc. (1887)
  • Lactifluus intermedius Krombh. ex Kuntze (1891)
  • Lactarius citriolens var. intermedius (Krombh. ex Kuntze) Krieglst. (1999)
  • Lactarius torminosus f. albida Killerm. (1933)
  • Lactarius torminosus var. gracillimus J.E. Lange (1938)
  • Lactarius torminosus var. gracillimus J.E. Lange (1940)
  • Lactarius torminosus var. typicus Kühner & Romagn. (1953)
  • Lactarius torminosus var. sublateritius Kühner & Romagn. (1954)
  • Lactarius nordmanensis A.H. Sm. (1960)
  • Lactarius torminosus var. nordmanensis (A.H. Sm.) Hesler & A.H. Sm. (1979)
Lactarius torminosus
View the Mycomorphbox template that generates the following list
Gills on hymenium
Cap is convex or depressed
Hymenium is decurrent
Stipe is bare
Spore print is cream to yellow
Ecology is
mycorrhizal
Edibility is not recommended

Lactarius torminosus, commonly known as the woolly milkcap or the bearded milkcap, is a large species of

molecular studies prompted the taxonomic reshuffling of species between several Russulaceae
genera.

A

mixed forests. The caps of L. torminosus mushrooms are convex with a central depression, and attain a diameter of up to 10 cm (3.9 in). A blend of pink and ochre hues, the cap sometimes has concentric zones of alternating lighter and darker shades. The edge of the cap is rolled inward, and shaggy when young. On the underside of the cap are narrow flesh-colored gills that are crowded closely together. The cylindrical stem is a pale flesh color with a 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 fruit bodies ooze a bitter white latex that does not change color upon exposure to air. The variety nordmanensis, in contrast, has latex that changes from white to yellow. Lactarius torminosus can be distinguished from similar species like L. pubescens or L. villosus by differences in morphology and coloration, or by microscopic characteristics like spore
shape and size.

Although it is valued for its peppery flavor and eaten after suitable preparation in Russia and Finland, the species is highly irritating to the digestive system when eaten raw. The toxins, also responsible for the strongly bitter or acrid taste, are destroyed by cooking. Studies have identified several chemicals present in the mushrooms, including ergosterol and derivatives thereof, and the pungent-tasting velleral.

Taxonomy and phylogeny

German naturalist

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

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

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

Young fruit body showing the tomentose cap margin and forked gills

The

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 more or less smooth.[22] The surface starts off somewhat sticky with clear concentric rings of darker shade (a zonate pattern); these rings, 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]

Although the cap surface is typically zonate—marked by concentric colored bands—in young specimens ...
... this feature is less prominent in mature mushrooms.

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

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 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]

cap cuticle is made of gelatinized, interwoven hyphae arranged more or less parallel to the cap surface (a form known as ixocutis); the thin-walled, threadlike hyphae of this layer are 2.5–7.3 μm 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 slowly changes to pale yellow upon exposure to air; the latex also 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

Lactarius pubescens
Lactarius mairei

The woolly cap margin, pinkish tones in the cap, acrid latex, and association with birch are reliable field characteristics to help identify L. torminosus.

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]

A comparison between Lactarius torminosus (left) and Lactarius pubescens (right)
A comparison between Lactarius torminosus (left) and Lactarius pubescens (right)

Edibility and toxicity

"Its taste is biting, worse than Cayenne pepper ... Unless he possessed a stomach built for the purpose, the man who touched such food as this would have a singularly bad time before him."

Jean-Henri Fabre[35]

The intensely peppery taste of the raw mushroom can blister the tongue if sampled in excess.

circulatory collapse. The gastroenteritis will usually resolve without treatment in a couple of days.[39]

Despite these reports of toxicity, L. torminosus mushrooms are prepared in Finland, Russia, and other northern and eastern European countries by parboiling, soaking in brine for several days, or pickling, after which they are valued for their 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

Structure of the sesquiterpene velleral

The compound thought to be responsible for the toxicity of raw L. torminosus is the pungent-tasting

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

1-octen-3-one,[46] an odorant common in mushrooms.[47]

Ecology and distribution

In Västerbotten, Sweden

Lactarius torminosus is a

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

References

  1. ^ a b c "Species synonymy: Lactarius torminosus (Schaeff.) Gray". Index Fungorum. CAB International. Archived from the original on 2012-10-07. Retrieved 2012-03-14.
  2. ^ Schäffer JC. (1774). Fungorum qui in Bavaria et Palatinatu circa Ratisbonam nascuntur icones [The Fungi that Grow in Bavaria and the Palatine, around Ratisbon] (in Latin and German). Vol. 4. Erlangen, Germany: Apud J.J. Palmium. pp. 7–8.
  3. ^ Bulliard JBF. (1781). Herbier de la France [Guide to the Herbs of France] (in French). Vol. 1. Paris, France: Chez l'auteur, Didot, Debure, Belin. pp. 1–48, plate 28.
  4. ^ Persoon Christian Hendrik (1799). Observationes mycologicae seu Descriptiones tam novorum, quam notabilium fungorum (in Latin). Leipzig, Germany: Gesnerus, Usterius & Wolfius. p. 42.
  5. ^ "Lactarius necator (Bull.) Pers. 1800". MycoBank. International Mycological Association. Retrieved 2012-04-22.
  6. ^ Kuntze O. (1898). Revisio generum plantarum (in Latin). Vol. 3. Leipzig, Germany: Arthur Felix. p. 489.
  7. ^ Kummer P. (1871). Der Führer in die Pilzkunde (in German). Zerbst, Germany: E. Luppe Staude. p. 125.
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  9. ^ a b c Gray SF. (1821). A Natural Arrangement of British Plants. London, UK: Baldwin, Cradock, and Joy. p. 623.
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  11. ^ Smith JE. (1824). The English Flora. Vol. 5. London, UK: Longman, Hurst, Rees, Orme, Brown, and Green. p. 24.
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  16. ^ "List of recommended English Names for Fungi in the UK" (PDF). British Mycological Society. 2003. Archived from the original (PDF) on 2011-07-16.
  17. ^ Hesler and Smith (1979), p. 237.
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  19. ^ Buyck B, Hofstetter V, Eberhardt U, Verbeken A, Kauff F (2008). "Walking the thin line between Lactarius and Russula: the dilemma of Russula sect. Ochricompactae" (PDF). Fungal Diversity. 28: 15–40.
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  28. ^ "Lactarius nordmanensis; Russulaceae (MICH5267)". Herbarium Fungus Collection Database. University of Michigan. Retrieved 2012-04-22.
  29. ^ Knudsen H, Borgen T (1994). "The Lactarius torminosus-group in Greenland". Mycologia Helvetica. 2: 49–56.
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