Auriscalpium vulgare

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Auriscalpium vulgare

Scientific classification
Kingdom: Fungi
Phylum: Basidiomycota
Class: Agaricomycetes
Order: Russulales
Family: Auriscalpiaceae
Genus: Auriscalpium
Species: A. vulgare
Binomial name
Auriscalpium vulgare
Gray (1821)
[show]Synonyms[1]
Hydnum auriscalpium L. (1753)
Scutiger auriscalpium (L.) Paulet (1812)
Pleurodon auriscalpium (L.) P.Karst. (1881)
Leptodon auriscalpium (L.) Quél. (1886)
Hydnum atrotomentosum Schwalb (1891)
Auriscalpium auriscalpium (L.) Kuntze
(1898)
Auriscalpium auriscalpium (L.) Banker
(1906)
Hydnum fechtneri Velen. (1922)
 Pleurodon fechtneri (Velen.) Cejp (1928)
Auriscalpium fechtneri (Velen.) Nikol.
(1964)

Auriscalpium vulgare

Mycological characteristics
teeth on hymenium

cap is offset

stipe is bare

spore print is white

ecology is saprotrophic

edibility: inedible

Auriscalpium vulgare, commonly known as the pinecone mushroom, the cone tooth, or the ear-
pick fungus, is a species of mushroom in the family Auriscalpiaceae of the Russulales order. It was
first described in 1753 by Carl Linnaeus, who included it as a member of the spine fungi genus
Hydnum, but in 1821, British mycologist Samuel Frederick Gray recognized its uniqueness and
made it the type species of the genus Auriscalpium that he created to contain it. The fungus is
widely distributed in Europe, Central America, North America, and temperate Asia. Although
common, its small size and nondescript colors lead it to be easily overlooked in the pine woods
where it grows. A. vulgare is not generally considered edible because of its tough texture, but some
older literature says it used to be consumed in France and Italy.
The fruit bodies typically grow on conifer litter or conifer cones, which may be partially or
completely buried in soil. The small, spoon-shaped mushroom has a dark brown cap covered with
fine brown fibrils, and reaches diameters of up to 2 cm (0.8 in). On the underside of the cap are a
crowded array of tiny spines up to 3 mm (0.12 in) long; they are initially whitish to purplish-pink
before turning brown in age. The stem, up to 55 mm (2.2 in) long and 2 mm (0.08 in) thick, is dark
brown and hairy, and attached to one side of the cap. The mushroom produces a white spore print,
and spores that are roughly spherical in shape, measuring 4.6–5.5 by 4–5 micrometers. Scientists
have investigated in detail the process of cell division and the ultrastructure of its hyphae using
electron microscopy. The mycelium of A. vulgare can be grown in pure culture in petri dishes, and
can be induced to produce fruit bodies under suitable conditions.
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History,
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[edit] History, taxonomy and phylogeny

Margaret Bentinck is credited with the first British collection in the 1760s.
The species was first described in the scientific literature by Carl Linnaeus under the name Hydnum
auriscalpium in his 1753 Species Plantarum. Linnaeus placed three other "spine" fungi in this
genus: H. imbricatum, H. repandum, and H. tomentosum.[2] The name was later sanctioned by Elias
Fries in his Systema Mycologicum.[3] The first recorded British collection was from Welbeck
Abbey, in the 1760s by the Duchess of Portland, Margaret Cavendish Bentinck. It was identified for
her by an English botanist, Reverend John Lightfoot.[4]
In 1821, Samuel Frederick Gray recognized that the fungus was distinct from the other spine fungi
described by Linnaeus, and he made it the type species of the genus Auriscalpium, in the process
changing its name to A. vulgare.[5] Both Otto Kuntze in 1898[6] and Howard James Banker[7] in
1906 tried to restore the original species name, resulting in the combination Auriscalpium
auriscalpium; however, tautonyms are illegitimate according to the rules for botanical nomenclature
(ICBN rule 23.4).[8] Additional synonyms include Hydnum fechtneri, named by Josef Velenovský
in 1922,[9] and later combinations based on this name: Pleurodon fechtneri published by Karel
Cejp in 1928, and Auriscalpium fechtneri by Taisiya Lvovna Nikolayeva in 1964.[1]

Cladogram showing the phylogeny of A. vulgare and other Auriscalpiaceae species based on rDNA
sequences.[10]
Despite vast differences in appearance and morphology, A. vulgare is related to such varied taxa as
the gilled fungi of Lentinus, the poroid genus Albatrellus, the coral-like Clavicorona, and fellow
spine fungi Hericium.[11] The relationship of all of these taxa—all members of the Auriscalpiaceae
family of the order Russula—has been demonstrated through molecular phylogenetics.[10][12]
Auriscalpium vulgare is commonly known as the "pinecone mushroom",[13] the "cone tooth",[14]
"pine cone tooth", or the "ear-pick fungus".[15] Gray called it the "common earpick-stool";[5] it
was also referred to as the "fir-cone Hydnum", when it was still a member of that genus.[16] The
specific epithet vulgare means "common".[15] The generic name Auriscalpium is Latin for "ear
pick" and refers to a small, scoop-shaped instrument used to remove foreign matter from the ear.
[17]

[edit] Description

The cap surface is bristly when young.

The spore-bearing surface features "teeth" up to 3 mm long.

The fruit body of A. vulgare is fibrous when fresh, and becomes stiff when dry. The cap is round or
kidney-shaped, flat on one side and rounded on the other, and between 0.5 and 2 cm (0.2 and 0.8
in)in diameter—although it has been known to reach up to 4 cm (1.6 in).[18] The surface is covered
with bristles, but becomes smooth with maturity; it is initially dark chestnut-brown but can become
almost black as it ages. The cap margin, usually lighter in color than the center, becomes rolled
inward in maturity and often wavy.[19] The color of the cap margin is buff to light brown, roughly
the same color as the spines.[20] The spines on the underside of the cap are up to 3 mm (0.12 in)
long, cylindrical down to the base with sharp tips. They are white to light brown when young, later
covered with a white spore mass and then ashy gray.[14][19]
Auriscalpium vulgare usually has a single stem, but occasionally several stems arise from a thick
common base. The stem is 14–55 mm (0.55–2.2 in) long, by 1–2 mm (0.04–0.08 in) thick. It is
attached to the side of the cap, and cylindrical or slightly flattened with a bulbous base. The surface
is covered with hairy fibers (especially near the base), and is a dark chestnut-brown color. There is
no distinct taste or odor. The flesh is up to 0.5 mm thick, and composed of two distinct layers. The
upper compact cortical layer is thin, black-brown, and covered with hairs. The lower layer is thick,
soft, white to light brown, and made of thin, thread-like filaments arranged in a roughly parallel
fashion.[14] Similarly, the stem is layered, with a thin dark cortical layer covered by hairs, which
encircles the inner ochraceous-colored flesh.[19] A drop of potassium hydroxide applied to the
surface of the mushroom will cause it to instantly stain black.[21] The mushroom is generally
considered inedible because of its toughness and small size;[22] however, an 1887 textbook noted
that it was "commonly eaten in France and Italy".[23]

[edit] Microscopic characteristics "... the mushroom

In deposit, the spores are white. Viewed with a light microscope, the look(s) like a little
spores appear hyaline (translucent), covered with minute wart-like bumps, periscope sent up from
and spherical or nearly so, with dimensions of 4.6–5.5 by 4–5 µm. The a pine-cone
spores are amyloid and cyanophilous, meaning they will absorb stain from submarine."
Melzer's reagent and methyl blue, respectively.[14] The basidia (spore- Michael Kuo[21]
bearing cells of the hymenium) are four-spored, have basal clamps, and
measure 15–24 by 3–4 µm. The sterigmata (slender extensions that attach the spores to the basidia)
are swollen at the base and roughly 3 µm long. The hyphal system is dimitic, meaning it made of
both generative hyphae (relatively undifferentiated hyphae that can develop reproductive structures)
and skeletal hyphae (long and thick-walled hyphae that provide structural support). The thin-walled
generative hyphae are 1.7–4 µm wide, hyaline, and have clamp connections; the thick-walled
skeletal hyphae are 1.5–4.8 µm wide. The cortex is made of branch-free generative hyphae that are
brown, thick-walled, parallel, clumped together, frequently clamped, and 2–4 µm wide. The
mushroom flesh is made of interwoven generative and skeletal hyphae. The gloeoplerous hyphae
(defined as containing resinous or granular contents) are 3–5 µm in diameter, with oily contents,
protruding into the hymenium as club-like or sharp-pointed gloeocystidia.[19]
The hyphae of basidiomycetous fungi are partitioned by cross-walls called septa, and these septa
have pores that permit the passage of cytoplasm or protoplasm between adjacent hyphal
compartments. The ultrastructure of the septal pore apparatus as well as nuclear division in
A. vulgar have been studied using electron microscopy, and have revealed certain characteristics
that are unique among the Russulales. The septa in hyphae of the fertile spore-bearing tissue have
bell-shaped perforated septal pore "caps". These caps can extend along the septum, along with a
zone surrounding the septal pore apparatus that is free of organelles. The process of metaphase I of
meiosis is similar to metaphase of mitosis. Globular spindle pole bodies with electron-opaque
inclusions are set within polar fenestrae (gaps in the nuclear membrane) of the nuclear envelope.
The nuclear envelope was shown to be mostly intact with occasional gaps. Fragments of
endoplasmic reticulum are present near the spindle pole bodies, but do not form a polar cap (a
chromophilic body beneath the anterior spore wall contained in the polar sac). According to the
authors, these microscopic characteristics are useful as additional data in phylogenetic analyses.[24]

A. vulgare on a Fir-cone, in different stages of development.

[edit] Growth in culture
Auriscalpium vulgare can be grown in pure culture on agar-containing plates supplemented with
nutrients. The colonies that grow are white to pale cream color, and cover the agar surface within
six weeks from the initial inoculation. The growth is made of hyphae that are bent-over, and do not
form aerial hyphae (hyphae that extend above the surface of the agar). Typically, two indistinct
zones develop at about 6 mm (0.24 in) and 15 millimetres (0.59 in) from the initial inoculum spot,
with each zone 3–4 mm (0.12–0.16 in) wide. The zones appears somewhat lighter in color because
the hyphae are more closely packed and form crystalline substances that deposit into the agar.[25]
The advancing edge of the mycelia is composed of two types of hyphae. The first are submerged
into the agar, measuring 0.8–1.5 µm in diameter. They are thin-walled, hyaline, clamped, long-
celled, and branched at nearly 90°. The second are the surface hyphae: they are straight to
occasionally somewhat spiral (subhelicoid), thin-walled, clamped, branched acutely, usually with a
clamp at the branch base, and measure 1.8–3.0 µm diameter. In addition, there are frequently
"empty" hyphae that are short-celled, and occasionally clamped. Gloeocystidia (thin-walled cystidia
with refractive, frequently granular contents) are rare.[25]
The mature mycelium consists of thin-walled, densely packed, vegetative hyphae that are 1.5–
3.2 µm in diameter. They are short-celled, clamped, often gnarled or subhelicoid, frequently
branched at about 45°, with a clamp at base of the branch. Their contents are somewhat granular to
packed with amorphous granules that appear refractive when viewed under phase contrast
microscopy. Their walls are often encrusted with tiny granules. Gloeocystidia are common; they
measure 50–85 by 6.5–8.5 µm, and are club-shaped (sometimes elongated), thin-walled, and often
have one or two lobes with rounded apices. Their contents are foamy and pale yellow, they appear
yellow-refractive under phase contrast. Initially they are erect but they soon fall under their own
weight to lie on agar surface. Crystalline deposits are abundant as small plate-like or star-like,
randomly scattered crystals.[25]
Fruiting begins about six weeks after the initial inoculation on the agar plate, but only when
portions of fruit bodies (teeth or stem sections) are used as the inoculum to initiate growth; the use
of vegetative mycelium as inoculum precludes subsequent fruiting. Mature fruit bodies grow very
close to the initial site of inoculation—within 3 mm (0.12 in)—and take about 60 days to mature
after they first start to form.[25]

[edit] Habitat and distribution

Growing on a Douglas-fir cone

Auriscalpium vulgare is a saprobic species. The fruit bodies grow solitarily or clustered on fallen
pine cones, especially those that are buried or partially buried.[26] It especially prefers Scots Pine
(Pinus sylvestris), but has also been reported to occur on spruce cones.[15] In the California area, it
grows primarily on Douglas-fir cones.[20] One author noted finding the mushroom on spruce
needles on top of squirrel dens where cone bracts were present in the duff.[27] In a Chinese study
conducted in the Laojun Mountain region, southwestern China, A. vulgare was found to be one of
the most dominant species collected from mixed forest at an altitude of 2,600–3,000 m (8,500–
9,800 ft).[28] A study on the effect of slash and burn practices for jhum cultivation in northeast
India showed that the fungus prefers to fruit on burned cones of the Khasi Pine, and that the number
of fruit bodies on unburned cones increases with increased girth size of the cone.[29]
The fungus is widely distributed in Europe, Central America, North America, temperate Asia,[30]
and Turkey.[31] In North America, its range extends north from Canada[27] south to the Trans-
Mexican Volcanic Belt south of Mexico City.[32] The mushroom is common, appearing in the
summer and autumn,[14] although it is easily overlooked because of its small size and nondescript
coloration.[2