Hydnellum peckii

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Hydnellum peckii
Hydnellum peckii2.jpg
Bleeding tooth fungus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Thelephorales
Family: Bankeraceae
Genus: Hydnellum
Species:
H. peckii
Binomial name
Hydnellum peckii
Banker (1912)
Synonyms [1]
Hydnellum peckii
Information icon.svg
Teeth icon.png Teeth on hymenium
Depressed cap icon.svgConvex cap icon.svg Cap is depressed or convex
NA cap icon.svg Hymenium attachment is not applicable
NA cap icon.svgLacks a stipe
Transparent spore print icon.svg
 Spore print is brown
Mycorrhizal fungus.svgEcology is mycorrhizal
Mycomorphbox Inedible.pngEdibility is inedible

Hydnellum peckii is a fungus in the genus Hydnellum of the family Bankeraceae. It is a hydnoid species, producing spores on the surface of vertical spines or tooth-like projections that hang from the undersurface of the fruit bodies. It is found in North America, Europe, and was recently discovered in Iran (2008) and Korea (2010). Hydnellum peckii is a mycorrhizal species, and forms mutually beneficial relationships with a variety of coniferous trees, growing on the ground singly, scattered, or in fused masses.

Contents

The fruit bodies typically have a funnel-shaped cap with a white edge, although the shape can be highly variable. Young, moist fruit bodies can "bleed" bright red guttation droplets that contain a pigment known to have anticoagulant properties similar to heparin. The unusual appearance of the young fruit bodies has earned the species several descriptive common names, including strawberries and cream, the bleeding Hydnellum, the bleeding tooth fungus, the red-juice tooth, and the Devil's tooth. Although Hydnellum peckii fruit bodies are readily identifiable when young, they become brown and nondescript when they age.

Taxonomy, phylogeny, and naming

The species was first described scientifically by American mycologist Howard James Banker in 1913. [2] Italian Pier Andrea Saccardo placed the species in the genus Hydnum in 1925, [3] while Walter Henry Snell and Esther Amelia Dick placed it in Calodon in 1956; [4] Hydnum peckii (Banker) Sacc. and Calodon peckii Snell & E.A. Dick are synonyms of Hydnellum peckii. [1]

The fungus is classified in the stirps (species thought to be descendants of a common ancestor) Diabolum of the genus Hydnellum, a grouping of similar species with the following shared characteristics: flesh that is marked with concentric lines that form alternating pale and darker zones (zonate); an extremely peppery taste; a sweetish odor; spores that are ellipsoid, and not amyloid (that is, not absorbing iodine when stained with Melzer's reagent), acyanophilous (not staining with the reagent Cotton Blue), and covered with tubercules; the presence of clamp connections in the hyphae. [5]

The specific eponym honors mycologist Charles Horton Peck. [6] The fungus is known in the vernacular by several names, including "strawberries and cream", the "bleeding Hydnellum", [7] the "red-juice tooth", "Peck's hydnum", [8] the "bleeding tooth fungus", [9] and the "devil's tooth". [10]

Description

As in all mushroom-producing fungi, the fruit bodies (sporocarps) are the reproductive structures that are produced from fungal mycelium when the appropriate environmental conditions of temperature, humidity and nutrient availability are met. Hydnellum peckii is a stipitate hydnoid fungus, meaning that it has a cap atop a stipe (stem), and a form resembling a Hydnum —characterized by a teeth-like hymenium, rather than gills or pores on the underside of the cap. Fruit bodies growing closely together often appear to fuse together (this is called "confluence"). They can reach a height of up to 10.5 cm (4+1⁄8 in). [5] Fresh fruit bodies exude a striking, thick red fluid when they are moist, [2] present even in young specimens, which are lumplike in appearance. [11]

The "teeth" covering the cap's underside are specialized structures that produce spores. Hydnellum peckii 62842.jpg
The "teeth" covering the cap's underside are specialized structures that produce spores.

The cap's surface is convex to flattened, more or less uneven and sometimes slightly depressed in the center. It is usually densely covered with "hairs" that give it a texture similar to felt or velvet; these hairs are sloughed off in age, leaving the caps of mature specimens smooth. [5] Its shape varies from somewhat round to irregular, 4 to 10 cm (1+5⁄8 to 3+7⁄8 in), or even as much as 20 cm (7+7⁄8 in) wide as a result of confluence. The cap is initially whitish, but later turns slightly brownish, with irregular dark-brown to nearly black blotches where it is bruised. In maturity, the surface is fibrous and tough, scaly and jagged, grayish brown in the upper part of the cap, and somewhat woody. The flesh is a pale pinkish brown. [12] The spore print is brown. [13]

The spines are slender, cylindrical and tapering (terete), less than 5 mm (1⁄4 in) long, and become shorter closer to the cap edge. They are crowded together, with typically between three and five teeth per square millimeter. [2] Pinkish white initially, they age to a grayish brown. [6] The stem is thick, very short, and often deformed. It becomes bulbous where it penetrates the ground, and may root into the soil for several centimeters. Although it may reach up to 5 cm (2 in) in total length, and is 1 to 3 cm (3⁄8 to 1+1⁄8 in) wide, only about 0.1 to 1 cm (1⁄16 to 3⁄8 in) appear above ground. The upper part is covered with the same teeth found on the underside of the cap, whereas the lower part is hairy and often encases debris from the forest floor. [14] The odor of the fruit body has been described as "mild to disagreeable", [6] or, as Banker suggested in his original description, similar to hickory nuts. [2]

Microscopic features

Hydnellum peckii, aka Bleeding Tooth fungus, Gaspereau Mountain (Wolfville) Nova Scotia, Canada, 16 August 2023 BleedingToothFungus.jpg
Hydnellum peckii, aka Bleeding Tooth fungus, Gaspereau Mountain (Wolfville) Nova Scotia, Canada, 16 August 2023

In deposit, the spores appear brown. Viewing them with a light microscope reveals finer details of their structure: they are roughly spherical but end abruptly in a small point, their surfaces are covered with small, wart-like nodules, and their size is between 5.0–5.3 by 4.0–4.7  μm. The spores are inamyloid, meaning they do not absorb iodine when stained with Melzer's reagent. [5]

Hydnellum peckii's cells (the hyphae) also present various characters useful for its characterization. The hyphae that form the cap are hyaline (translucent), smooth, thin-walled, and 3–4 μm thick. They collapse when dry, but may be readily revived with a weak (2%) solution of potassium hydroxide. Those in the cap form an intricate tangle with a tendency to run longitudinally. They are divided into cellular compartments (septa) and have clamp connections—short branches connecting one cell to the previous cell to allow passage of the products of nuclear division. [2] The basidia, the spore-bearing cells in the hymenium, are club-shaped, four-spored, and measure 35–40 by 4.7–6 μm. [5]

Similar species

Hydnellum diabolus (the species epithet is given the neuter diabolum in some publications) [5] [15] has a very similar appearance, so much so that some consider it and H. peckii to be synonymous; H. diabolus is said to have a sweetish pungent odor that is lacking in H. peckii. [16] The differences between the two species are amplified in mature specimens: H. diabolus has an irregularly thickened stem, while the stem of H. peckii is thickened by a "definite spongy layer". Additionally, old specimens of H. peckii have a smooth cap, while H. diabolus is tomentose. [5] The related species H. pineticola also exudes pink droplets of liquid when young and moist. Commonly found growing under conifers in northeastern North America, H. pineticola tastes "unpleasant", but not acrid. [7] Fruit bodies tend to grow singly, rather than in fused clusters, and, unlike H. peckii, they do not have bulbous stems. [17]

Molecular analysis based on the sequences of the internal transcribed spacer DNA of several Hydnellum species placed H. peckii as most closely related to H. ferrugineum and H. spongiosipes . [18]

Ecology

Hydnellum peckii is a mycorrhizal fungus, and as such establishes a mutualistic relationship with the roots of certain trees (referred to as "hosts"), in which the fungus exchanges minerals and amino acids extracted from the soil for fixed carbon from the host. The subterranean hyphae of the fungus grow a sheath of tissue around the rootlets of a broad range of tree species, in an intimate association that is especially beneficial to the host (termed ectomycorrhizal), as the fungus produces enzymes that mineralize organic compounds and facilitate the transfer of nutrients to the tree. [19]

The ectomycorrhizal structures of H. peckii are among a few in the Bankeraceae that have been studied in detail. They are characterized by a plectenchymatous mantle—a layer of tissue made of hyphae tightly arranged in a parallel orientation, or palisade, and which rarely branch or overlap each other. These hyphae, along with adhering mineral soil particles, are embedded in a gelatinous matrix. The hyphae of the ectomycorrhizae can become chlamydospores, an adaptation that helps the fungus tolerate unfavorable conditions. Chlamydospores of H. peckii have a peculiar structure—markedly distinct from those of other Bankeraceae—with thick, smooth inner walls and an outer wall that is split radially into warts. The most striking characteristic of the ectomycorrhizae as a whole is the way the black outer layers of older sections are shed, giving a "carbonized appearance". [20] The majority of the underground biomass of the fungus is concentrated near the surface, [21] most likely as "mycelial mats"—dense clusters of ectomycorrhizae and mycelium. [22] The mycelium is also known to extend far beyond the site of the fruit bodies, as far as 337 centimeters (11+1⁄12 ft) away. [22]

Molecular techniques have been developed to help with conservation efforts of stipitate hydnoid fungi, including H. peckii. While the distribution of the fungus has traditionally been determined by counting the fruit bodies, this method has a major drawback in that fruit bodies are not produced consistently every year, and the absence of fruit bodies is not an indication of the absence of its mycelium in the soil. [23] More modern techniques using the polymerase chain reaction to assess the presence of the fungal DNA in the soil have helped alleviate the issues in monitoring the presence and distribution of fungi mycelia. [24]

Habitat and distribution

Fruit bodies can grow solitarily, scattered, or clustered together. Location: Tusby, Finland. Hydnellum peckii - Karvasorakas C IMG 7751.jpg
Fruit bodies can grow solitarily, scattered, or clustered together. Location: Tusby, Finland.

The fruit bodies of Hydnellum peckii are found growing solitary, scattered, or clustered together on the ground under conifers, often among mosses and pine needle litter. H. peckii is a "late-stage" fungus that, in boreal forests dominated by jack pine, typically begins associating with more mature hosts once the canopy has closed. [25] A preference for mountainous or subalpine ecosystems has been noted. [6]

The fungus has a wide distribution in North America, and is particularly common in the Pacific Northwest; [26] its range extends north to Alaska and east to North Carolina. [8] In the Puget Sound area of the U.S. state of Washington, it is found in association with Douglas-fir, fir, and hemlock. [5] Along the Oregon Coast it has been collected under lodgepole pine. [14] In addition to North America, the mushroom is widespread in Europe, and its presence has been documented in Italy, [27] Germany, [28] and Scotland. [29] The species is common in the latter location, but becoming increasingly rare in several European countries, such as Norway, [30] The Netherlands, [31] and the Czech Republic. [32] Increased pollution in central Europe has been suggested as one possible factor in the mushroom's decline there. [29] Reports from Iran in 2008 [33] and Korea in 2010 [34] were the first outside Europe and North America.

Uses

Fruit bodies of H. peckii have been described as resembling "Danish pastry topped with strawberry jam". [7] Hydnellum species are not known to be poisonous, [35] but they are not particularly edible due to their foul taste. [7] This acrid taste persists even in dried specimens. [26]

The fruit bodies of this and other Hydnellum species are prized by mushroom dyers. [36] The colors may range from beige when no mordant is used, to various shades of blue or green depending on the mordant added. [26]

Chemistry

Structural formula of atromentin Atromentin.svg
Structural formula of atromentin

Screening of an extract of Hydnellum peckii revealed the presence of an effective anticoagulant, named atromentin (2,5-dihydroxy-3,6-bis(4-hydroxyphenyl)-1,4-benzoquinone), and similar in biological activity to the well-known anticoagulant heparin. [37] Atromentin also possesses antibacterial activity, inhibiting the enzyme enoyl-acyl carrier protein reductase (essential for the biosynthesis of fatty acids) in the bacteria Streptococcus pneumoniae . [38]

Hydnellum peckii can bioaccumulate the metal caesium. In one Swedish field study, as much as 9% of the total caesium of the topmost 10 cm (3+7⁄8 in) of soil was found in the fungal mycelium. [21] In general, ectomycorrhizal fungi, which grow most prolifically in the upper organic horizons of the soil or at the interface between the organic and mineral layers, are involved in the retention and cycling of caesium-137 in organic-rich forest soils. [39]

Related Research Articles

<i>Hydnum repandum</i> Species of edible fungus of the family Hydnaceae distributed in Europe

Hydnum repandum, commonly known as the sweet tooth, pig's trotter, wood hedgehog or hedgehog mushroom, is a basidiomycete fungus of the family Hydnaceae. First described by Carl Linnaeus in 1753, it is the type species of the genus Hydnum. The fungus produces fruit bodies (mushrooms) that are characterized by their spore-bearing structures—in the form of spines rather than gills—which hang down from the underside of the cap. The cap is dry, colored yellow to light orange to brown, and often develops an irregular shape, especially when it has grown closely crowded with adjacent fruit bodies. The mushroom tissue is white with a pleasant odor and a spicy or bitter taste. All parts of the mushroom stain orange with age or when bruised.

<i>Hydnellum</i> Genus of fungi in the family Bankeraceae

Hydnellum is a genus of tooth fungi in the family Bankeraceae. Widely distributed in the Northern Hemisphere, the genus contains around 40 species. The fruitbodies of its members grow by slowly enveloping nearby bits of grass and vegetation. There is great variability in the form of Hydnellum fruitbodies, which are greatly influenced by environmental conditions such as rainfall and humidity, drying winds, and temperature. They are too tough and woody to eat comfortably. Several species have become the focus of increasing conservation concern following widespread declines in abundance.

<i>Hydnellum concrescens</i> Species of fungus

Hydnellum concrescens is an inedible fungus, commonly known as the zoned hydnellum or zoned tooth fungus. As with other tooth fungi, the spores are produced on spines on the underside of the cap, rather than gills. It has a funnel-shaped cap, typically between 2 and 7 cm in diameter, which has characteristic concentric zones of color. The cap may also have radial ridges extending from the center to the margins. The spines are pink in young specimens, but turn brown with age.

<i>Hydnellum caeruleum</i> Species of fungus

Hydnellum caeruleum, commonly known as the blue-gray hydnellum, blue-green hydnellum, blue spine, blue tooth, or bluish tooth, is an inedible fungus found in North America, Europe, and temperate areas of Asia.

<span class="mw-page-title-main">Hydnaceae</span> Family of fungi

The Hydnaceae are a family of fungi in the order Cantharellales. Originally the family encompassed all species of fungi that produced basidiocarps having a hymenium consisting of slender, downward-hanging tapering extensions referred to as "spines" or "teeth", whether they were related or not. This artificial but often useful grouping is now more generally called the hydnoid or tooth fungi. In the strict, modern sense, the Hydnaceae are limited to the genus Hydnum and related genera, with basidiocarps having a toothed or poroid hymenium. Species in the family are ectomycorrhizal, forming a mutually beneficial relationship with the roots of trees and other plants. Hydnum repandum is an edible species, commercially collected in some countries and often marketed under the French name pied de mouton.

<i>Phellodon</i> Genus of tooth fungi in the family Bankeraceae

Phellodon is a genus of tooth fungi in the family Bankeraceae. Species have small- to medium-sized fruitbodies with white spines on the underside from which spores are released. All Phellodon have a short stalk or stipe, and so the genus falls into the group known as stipitate hydnoid fungi. The tough and leathery flesh usually has a pleasant, fragrant odor, and develops a cork-like texture when dry. Neighboring fruitbodies can fuse, sometimes producing large mats of joined caps. Phellodon species produce a white spore print, while the individual spores are roughly spherical to ellipsoid in shape, with spiny surfaces.

<i>Sarcodon</i> Genus of fungi

Sarcodon is a genus of fungi in the family Bankeraceae, which is part of the order Thelephorales, known for its almost universal ectomycorrhizal ecology. The genus owes its name to the presence of teeth-like spines on the hymenophore, from ancient Greek; sarco = flesh and odon = tooth. For this reason they are commonly called "tooth fungi", or "Hydnoid fungi".

<i>Hydnellum aurantiacum</i> Species of fungus

Hydnellum aurantiacum is an inedible fungus, commonly known as the orange spine or orange hydnellum for its reddish orange or rusty red colored fruit bodies. Like other tooth fungi, it bears a layer of spines rather than gills on the underside of the cap. Due to substantial declines in sightings, this species is listed as critically endangered in the United Kingdom.

<span class="mw-page-title-main">Hydnoid fungi</span> Group of fungi

The hydnoid fungi are a group of fungi in the Basidiomycota with basidiocarps producing spores on pendant, tooth-like or spine-like projections. They are colloquially called tooth fungi. Originally such fungi were referred to the genus Hydnum, but it is now known that not all hydnoid species are closely related.

<i>Suillus pungens</i> Species of fungus in the family Suillaceae found in California

Suillus pungens, commonly known as the pungent slippery jack or the pungent suillus, is a species of fungus in the genus Suillus. The fruit bodies of the fungus have slimy convex caps up to 14 cm (5.5 in) wide. The mushroom is characterized by the very distinct color changes that occur in the cap throughout development. Typically, the young cap is whitish, later becoming grayish-olive to reddish-brown or a mottled combination of these colors. The mushroom has a dotted stem (stipe) up to 7 cm (2.8 in) long, and 2 cm (0.8 in) thick. On the underside on the cap is the spore-bearing tissue consisting of minute vertically arranged tubes that appear as a surface of angular, yellowish pores. The presence of milky droplets on the pore surface of young individuals, especially in humid environments, is a characteristic feature of this species. S. pungens can usually be distinguished from other similar Suillus species by differences in distribution, odor and taste. The mushroom is considered edible, but not highly regarded.

<i>Hydnellum ferrugineum</i> Species of fungus

Hydnellum ferrugineum, commonly known as the mealy tooth or the reddish-brown corky spine fungus, is a species of tooth fungus in the family Bankeraceae. A widely distributed species, it is found in north Africa, Asia, Europe, and North America. The fungus fruits on the ground singly or in clusters in conifer forest, usually in poor or sandy soil. Fruit bodies are somewhat top-shaped, measuring 3–10 cm (1–4 in) in diameter. Their velvety surfaces, initially white to pink, sometimes exude drops of red liquid. The lower surface of the fruit body features white to reddish-brown spines up to 6 mm long. Mature fruit bodies become dark reddish brown in color, and are then difficult to distinguish from other similar Hydnellum species. H. ferrugineum forms a mat of mycelia in the humus and upper soil where it grows. The presence of the fungus changes the characteristics of the soil, making it more podzolized.

<i>Suillus collinitus</i> Species of fungus

Suillus collinitus is a pored mushroom of the genus Suillus in the family Suillaceae. It is an edible mushroom found in European pine forests. The mushroom has a reddish to chestnut-brown cap that reaches up to 11 cm (4.3 in) in diameter, and a yellow stem measuring up to 7 cm (2.8 in) tall by 1 to 2 cm thick. On the underside of the cap are small angular pores, initially bright yellow before turning greenish-brown with age. A characteristic feature that helps to distinguish it from similar Suillus species, such as S. granulatus, is the pinkish mycelia at the base of the stem.

<i>Auriscalpium vulgare</i> Inedible European fungi

Auriscalpium vulgare, commonly known as the pinecone mushroom, the cone tooth, or the ear-pick fungus, is a species of fungus in the family Auriscalpiaceae of the order Russulales. It was first described in 1753 by Carl Linnaeus, who included it as a member of the tooth fungi genus Hydnum, but British mycologist Samuel Frederick Gray recognized its uniqueness and in 1821 transferred it to the genus Auriscalpium that he created to contain it.

<i>Hydnellum fuscoindicum</i> Species of fungus

Hydnellum fuscoindicum, commonly known as the violet hedgehog, is a species of tooth fungus in the genus Hydnellum. It is found in the Pacific Northwest in moss around western hemlocks. It produces fruit bodies with a violet-black cap, violet flesh, and violet spines on the cap underside. The odor and taste are very farinaceous. The fungus was first described by Kenneth A. Harrison in 1964 as a species of Hydnum, then transferred to Sarcodon in 1967 by Rudolph Arnold Maas Geesteranus. He placed this species in section Violacei of Sarcodon, along with H. fuligineoviolaceum and H. joeides.

<i>Hydnellum compactum</i> Species of fungus

Hydnellum compactum is a rare tooth fungus in the family Bankeraceae. It is characterised by its astringent taste, yellowish flesh. It is found in Europe, where it grows in deciduous forest, typically under beech. Fruit bodies of the fungus grow singly or in groups.

<i>Hydnellum cyanopodium</i> Species of fungus

Hydnellum cyanopodium, commonly known as the blue foot or bleeding blue tooth, is an inedible fungus in the family Bankeraceae. It occurs in the Pacific Northwest region of North America.

<i>Hydnellum scrobiculatum</i> Species of fungus

Hydnellum scrobiculatum, commonly known as the ridged tooth or rough hydnellum, is a tooth fungus in the family Bankeraceae. Widely distributed in the Northern Hemisphere, it is found in Asia, Europe, and North America.

<i>Phellodon niger</i> Species of fungus

Phellodon niger, commonly known as the black tooth, is a species of tooth fungus in the family Bankeraceae, and the type species of the genus Phellodon. It was originally described by Elias Magnus Fries in 1815 as a species of Hydnum. Petter Karsten included it as one of the original three species when he circumscribed Phellodon in 1881. The fungus is found in Europe and North America, although molecular studies suggest that the North American populations represent a similar but genetically distinct species.

<i>Hydnellum joeides</i> Species of fungus

Hydnellum joeides is a species of tooth fungus in the family Bankeraceae.

<i>Hydnellum gracilipes</i> Species of fungus

Hydnellum gracilipes is a species of tooth fungus in the family Bankeraceae. It was first described scientifically in 1886 by Petter Karsten, who called it Hydnum gracilipes. He transferred it to the genus Hydnellum in 1879. Fruit bodies of the fungus have a pinkish to reddish-brown colour, a delicate texture described as "felty or papery", and flimsy stipes. Its spores are roughly spherical with a diameter of no more than 5 μm. H. gracilipes is found in northern Europe, where it is mycorrhizal with pine. Collections made in Scotland have been found by lifting the dense ground cover of common heather, which the fungus seems to use to as support to compensate for its flimsy stipe.

References

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