Phellodon

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Phellodon
Phellodon niger 496570.jpg
Phellodon niger
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Thelephorales
Family: Bankeraceae
Genus: Phellodon
P.Karst. (1881)
Type species
Phellodon niger
(Fr.) P.Karst. (as 'nigrum') (1881)
Species
about 20

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.

Contents

The genus, with about 20 described species, has a distribution that includes to Asia, Europe, North America, South America, Australia, and New Zealand. About half of the species are found in the southeastern United States, including three species added to the genus in 2013–14. Several Phellodon species were placed on a preliminary Red List of threatened British fungi because of a general decline of the genus in Europe. Species grow in a symbiotic mycorrhizal association with trees from the families Fagaceae (beeches and oaks) and Pinaceae (pines). Accurate DNA-based methods have been developed to determine the presence of Phellodon species in the soil, even in the extended absence of visible fruitbodies. Although Phellodon fruitbodies are considered inedible due to their fibrous flesh, the type species, P. niger , is used in mushroom dyeing.

Taxonomy

Phellodon was circumscribed in 1881 by Finnish mycologist Petter Karsten to contain white-toothed fungi. Karsten included three species: P. cyathiformis , P. melaleucus , and the type, P. niger (originally published with the epithet nigrum). [1] P. nigrum was originally described by Elias Fries in 1815 as a species of Hydnum . [2] Some early authors did not consider Phellodon distinct enough to be a separate genus, and folded species assigned to this genus into Hydnellum . [3] [4]

Hydnellum is classified in the family Bankeraceae, which was circumscribed by Marinus Anton Donk in 1961. Donk's original family concept included the genera Bankera and Phellodon, whose species produce hyaline (translucent) and echinulate spores (covered with small spines). Donk also noted that Bankeraceae species lacked clamp connections. [5] When clamp connections were discovered in Phellodon fibulatus and tuberculate spore ornamentation (the presence of small nodules on the spores) was found in P. niger, Kenneth Harrison thought the family Bankeraceae was superfluous, and placed Phellodon and Bankera in the family Hydnaceae. [6] This taxonomic rearrangement was rejected by Rudolph Arnold Maas Geesteranus in 1974, who showed that the tuberculate spores of P. niger were the result of an immature specimen. [7] Richard Baird and Saeed Khan investigated spore ornamentation in North American Phellodon species using scanning electron microscopy, and rejected the placement of Phellodon in the Bankeraceae, preferring to leave it and Bankera in the Hydnaceae. [8] Modern molecular phylogenetic analysis places Phellodon in the thelephoroid clade (roughly equivalent to the order Thelephorales) along with the related genera Bankera, Hydnellum, and Sarcodon . Although the status of the Bankeraceae has not been fully clarified with molecular genetic techniques, [9] Phellodon is classified in this family by authorities on fungal taxonomy. [10] [11] [12]

The generic name is derived from Greek phell-, meaning 'cork', and -don, meaning 'tooth'. In North America, Phellodon species are commonly known as "cork hydnums". [13] The British Mycological Society, in their recommended list of common names for fungi in the United Kingdom, name Phellodon species in the form "descriptor word" plus tooth: fused tooth (P. confluens), grey tooth (P. melaleucus), black tooth (P. niger), and woolly tooth (P. tomentosus). [14]

Description

The caps of some Phellodon species (P. tomentosus pictured) can fuse during growth. Phellodon tomentosus I Slavkovsky les CZ.jpg
The caps of some Phellodon species ( P. tomentosus pictured) can fuse during growth.

The fruit bodies of Phellodon species have caps and stipe, and thus fall into the general category of stipitate hydnoid fungi. The cap surfaces are initially velvety to tomentose, eventually becoming matted. The surface is rough, with pits and ridges, and sometimes with concentrical zones of color or texture. The color can vary considerably, from cream to yellowish, brownish, greenish, greyish or black. [15] Neighboring fruitbodies can fuse, forming intertwined caps and partially fused stipes. [16] Alexander H. Smith wrote of P. tomentosus, "It often occurs in large mats of fused caps almost producing a ceiling over large areas of the moss under conifers." [17] Phellodon fruitbodies can envelop nearby grass or twigs. The stipe is thickly tomentose or smooth, typically the same color as the cap or darker. [18] In P. niger, the outer covering of the stipe is a thick felty layer of mycelium that absorbs water like a sponge. [19] The hymenophore (the fertile, spore-bearing surface) is on the underside of the cap. The spines become grey at maturity. [18] In conditions of high humidity, P. niger can form striking drops of black liquid on the actively growing caps. [19]

Spines of Phellodon tomentosus Phellodon tomentosus (7).JPG
Spines of Phellodon tomentosus

The fibrous flesh is single to double-layered; duplex layering results from differences in compactness or in the alignment of the constituent hyphae. [15] Tough and leathery when fresh, the flesh develops a corklike texture when dry. In the dried state it often has an odor of fenugreek or curry powder. [16] Phellodon species are often free of insect damage, suggesting that they may have defensive chemicals that deter predation. [20] Fruitbodies are not considered edible due to their fibrous flesh. [17]

The hyphal system is monomitic, containing only generative hyphae. These hyphae are not less than 6  μm in diameter. All European species lack clamp connections, [16] but they are present in the North American species P. fibulatus [6] and P. mississippiensis . [21] The basidia (spore-bearing cells) are four-spored. [18] Cystidia are either absent, or present infrequently as incompletely differentiated cystidioles (sterile cells about the size of an immature basidium). [16] In mass, the spores are white. [13] Spores are broadly ellipsoid to roughly spherical, and echinulose (covered with small spines). They are also hyaline and inamyloid. [18]

Habitat and distribution

Fruit bodies grow on the ground. Phellodon species, like all members of the order Thelephorales, are thought to be mycorrhizal, forming symbiotic relationships with trees. Usual hosts include species from the families Fagaceae (beeches and oaks) and Pinaceae. [15] The ectomycorrhizae that P. niger forms with Norway spruce (Picea abies) has been comprehensively described. It is distinguished from the ectomycorrhizae of other Thelephorales species by the unique shape of its chlamydospores. [22] Stable isotope ratio analysis of the abundance of the stable isotope carbon-13 shows that P. niger has a metabolic signature close to that of saprotrophic fungi, indicating that it may be able to obtain carbon from sources other than a tree host. [23] [24]

Many Phellodon species are known from the southeastern United States, where they have been extensively researched. According to Baird and colleagues, there are nine distinct Phellodon species from Great Smoky Mountains and the surrounding southern Appalachian Mountains. Three additional species from this area, P. mississippiensis , P. brunneo-olivaceus , and P. fuligineoalbus were added to the genus in 2013–14. [21] [25]

Conservation

Phellodon melaleucus Phellodon melaleucus 264419.jpg
Phellodon melaleucus

Phellodon species, like other members of the family Bankeraceae, are sensitive to air pollution and soil pollution, and are in general decline in western Europe. [15] In a preliminary assessment for a red list of threatened British Fungi, P. confluens, P. tomentosus, and P. melaleucus are considered vulnerable, and P. niger is rare. [26]

Conservation efforts for stipitate hydnoid fungi are hampered by a dearth of information about their basic ecology, and so molecular genetic techniques are increasingly employed in attempts to better understand these fungi. In the case of Phellodon tomentosus, for example, there is little correlation between fruitbody appearance and below-ground mycelium, making it hard to determine the distribution and rarity of the fungus with standard surveying techniques. [27] Phellodon melaleucus and P. niger were included in a Scottish study to develop species-specific PCR primers that can be used to detect the mycelia of stipitate hydnoids in soil. [28] DNA testing of collections labelled as P. melaleucus and P. niger from the United Kingdom revealed additional cryptic species. [29] [30] PCR analysis can be used to determine the presence of a Phellodon species up to four years after the appearance of fruitbodies, allowing a more accurate determination of their possible decline and threat of extinction. [31]

Chemistry

Phellodon species contain thelephoric acid, a metabolite of the shikimic acid pathway. Thelephoric acid is a terphenyl quinone—a 1,4-benzoquinone wherein positions carbon-2 and carbon-5 are substituted with phenyl groups. The hirsutane derivative phellodonic acid is found in P. melaleucus. Phellodonic acid, which exhibits antibiotic activity towards bacteria and other fungi, was the first bioactive compound reported from any member of the order Thelephorales. [32] A total synthesis was described for phellodonic acid in 2008 using cis-1,2-dihydrocatechol as the starting material. [33] P. niger has been a source for several bioactive compounds: the cyathane-type diterpenoids, nigernin A and B; a nitrogenous terphenyl derivative, phellodonin; 2',3'-diacetoxy-3,4,5',6',4''-pentahydroxy-p-terphenyl; grifolin; and 4-O-methylgrifolic acid. [34] P. niger has also been used for mushroom dyeing, in which it produces gray-blue and green colors. [35]

Species

Phellodon was originally circumscribed with three species. Joost Stalpers included 13 Phellodon species in his 1993 monograph on the Thelephorales. [15] The tenth edition of the Dictionary of the Fungi (2008) indicated 16 species in the genus. [11] As of September 2015, Index Fungorum lists 18 species of Phellodon, [36] not including the three eastern United States species added in 2013–14. [21] [25]

ImageScientific nameDistribution
2009-12-23 Phellodon atratus 30824 cropped.jpg Phellodon atratus K.A.Harrison (1964) California, United States [37]
Phellodon brunneo-olivaceus R.E.Baird (2013)United States [25]
Phellodon confluens (Pers.) Pouzar (1956)China, [38] eastern United States, [39] Europe [40]
Phellodon excentrimexicanus R.E.Baird (1985)Mexico [41]
Phellodon fibulatus K.A.Harrison (1972) North Carolina, United States [6]
Phellodon fuligineoalbus (J.C.Schmidt) Baird (2013)United States [25]
Phellodon implicatus R.E.Baird & S.R.Khan (1986) Florida, United States [42]
Phellodon indicus Khara (1978) Himachal Pradesh, India [43]
Phellodon maliensis (Lloyd) Maas Geest. (1966)Australia, New Zealand [44]
Phellodon melaleucus 264419.jpg Phellodon melaleucus (Sw. ex Fr.) P.Karst. (1881)Europe, North America [16]
Phellodon mississippiensis R.Baird (2014) Mississippi, United States [21]
2013-10-24 Phellodon niger (Fr.) P. Karst 379497.jpg Phellodon niger (Fr.) P.Karst. (1881)Europe, North America [16]
Phellodon nothofagi McNabb (1971)New Zealand [45]
Phellodon plicatus (Lloyd) Maas Geest. (1966)Australia [46]
Phellodon putidus (G.F.Atk.) Banker (1906)North America [47]
Phellodon radicatus R.E.Baird (1985)North America [41]
Phellodon rufipes Maas Geest. (1971)Japan [44]
Phellodon secretus by caspar s.jpg Phellodon secretus Niemelä & Kinnunen (2003) –Finland [48]
Phellodon sinclarii (13142242875).jpg Phellodon sinclairii (Berk.) G.Cunn. (1958)New Zealand [49]
Phellodon tenuis R.E.Baird (1988)Brazil [50]
Phellodon tomentosus 120380.jpg Phellodon tomentosus (L.) Banker (1906)Europe, [16] North America [47]

Related Research Articles

<span class="mw-page-title-main">Hydnellum peckii</span> Species of fungus

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.

<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>Bankera</i> Genus of fungi

Bankera is a genus of four species of tooth fungi in the family Bankeraceae. The genus was first circumscribed in 1951 by William Chambers Coker and Alma Holland Beers, but this publication was invalid according to the rules of botanical nomenclature. It was later published validly by Zdeněk Pouzar in 1955. The type species is B. fuligineoalba. The genus is ectomycorrhizal with gymnosperms, usually with trees from the pine family.

<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>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>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.

Hydnellum nigellum is a tooth fungus in the family Bankeraceae. Found in Nova Scotia, Canada, where it grows under spruce, it was described as new to science in 1964 by Canadian mycologist Kenneth A. Harrison. The fruitbodies of this fungus are small, measuring 3–6 cm (1.2–2.4 in) in diameter, and black.

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

Hydnellum regium is a tooth fungus in the family Bankeraceae. Found in western North America, it was described as new to science in 1964 by Canadian mycologist Kenneth A. Harrison, who reported collections from Oregon, Idaho, and Colorado. It fruits singly or in groups under pine and spruce trees.

<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 fuligineoviolaceum</i> Species of fungus

Hydnellum fuligineoviolaceum is a species of tooth fungus in the family Bankeraceae. It was originally described in 1874 by Hungarian mycologist Károly Kalchbrenner as Hydnum fuligineoviolaceum, in Elias Fries's work Hymenomycetes europaei. Narcisse Théophile Patouillard transferred it to the genus Sarcodon in 1900. Sarcodon talpa, published by Rudolph Arnold Maas Geesteranus in 1967, is a synonym.

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

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

<i>Sarcodon thwaitesii</i> Species of fungus

Sarcodon thwaitesii is a species of tooth fungus in the family Bankeraceae. It is found in Asia, Europe, and New Zealand, where it fruits on the ground in mixed forest.

<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.

Kenneth A. Harrison was a Canadian mycologist. He was for many years a plant pathologist at what is now the Atlantic Food and Horticulture Research Centre in Nova Scotia. After retirement, he contributed to the taxonomy of the Agaricomycotina, particularly the tooth fungi of the families Hydnaceae and Bankeraceae, in which he described several new species.

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

Hydnellum versipelle is a species of tooth fungus in the family Bankeraceae. It was originally described by Elias Fries in 1861 as Hydnum versipelle. Taisiya Lvovna Nikolayeva transferred it to the genus Sarcodon in 1961. Hydnum crassum, published by Kenneth A. Harrison in 1961, is a synonym. The species is found in Europe and North America.

References

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  2. ↑ Fries EM. (1815). Observationes mycologicae (in Latin). Vol. 1. Copenhagen: Gerhard Bonnier. p. 134.
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  5. ↑ Donk MA. (1961). "Four new families of Hymenomycetes". Persoonia. 1 (4): 405–407.
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  7. ↑ Maas Geesteranus RA. (1974). "Notes on hydnums. IX". Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen Afdeling Natuurkunde, Series C. 77: 215–222.
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  17. 1 2 Smith AH. (1975). A Field Guide to Western Mushrooms. Ann Arbor, Michigan: University of Michigan Press. p. 57. ISBN   978-0-472-85599-5.
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  22. ↑ Agerer R. (1992). "Ectomycorrhizae of Phellodon niger on Norway spruce and their chlamydospores". Mycorrhiza. 2 (1): 47–52. doi:10.1007/BF00206283. S2CID   11608006.
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  25. 1 2 3 4 Baird RE, Wallace LE, Baker G, Scruggs M (2013). "Stipitate hydnoid fungi of the temperate southeastern United States". Fungal Diversity. 62 (1): 41–114. doi:10.1007/s13225-013-0261-6. S2CID   16846274.
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  30. ↑ Ainsworth AM, Parfitt D, Rogers HJ, Boddy L (2010). "Cryptic taxa within European species of Hydnellum and Phellodon revealed by combined molecular and morphological analysis". Fungal Ecology. 3 (2): 65–80. doi:10.1016/j.funeco.2009.07.001.
  31. ↑ van der Linde S, Holden E, Parkin PI, Alexander IJ, Anderson IC (2012). "Now you see it, now you don't: The challenge of detecting, monitoring and conserving ectomycorrhizal fungi". Fungal Ecology. 5 (5): 633–640. doi:10.1016/j.funeco.2012.04.002.
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