Hydnellum

{{Short description|Genus of fungi in the family Bankeraceae}} {{Good article}} {{Automatic taxobox | image = Hydnellum peckii2.jpg | image_caption = The bleeding tooth fungus – ''Hydnellum peckii'' | taxon = Hydnellum | authority = P.Karst. (1879) | type_species = ''Hydnellum suaveolens'' | type_species_authority = Scop. : Fr. (1772) | synonyms_ref = <ref name="urlMycoBank: Hydnellum"/> | synonyms = *''Calodon'' Quél. ex P.Karst. (1881) *''Phaeodon'' J.Schröt. (1888) | subdivision_ranks = Species | subdivision = }}

'''''Hydnellum''''' is a genus of tooth fungi in the family Bankeraceae (order Thelephorales). Widely distributed in the Northern Hemisphere, the genus contains around 120 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.

''Hydnellum'' species produce pigments that have been used to dye textiles. Several chemical compounds—some with unique biological activity—have been isolated and identified from the genus.

One of the better-known species is the unusual pinkish-white ''Hydnellum peckii'', also known as "strawberries and cream" or as the "bleeding tooth fungus" due to the red droplets that appear on the pinkish or whitish fruitbodies. Another species, ''H. suaveolens'', has a strong odor of anise or peppermint.

==Taxonomy== ''Hydnellum'' was circumscribed by Finnish mycologist Petter Adolf Karsten in 1879 with what was then known as ''Hydnum suaveolens'' as the type species.<ref name="Karsten 1879"/> Before then, fungi with spines (hydnoid fungi) had been grouped in ''Hydnum'' by Elias Fries in his 1821 work ''Systema mycologicum''.<ref>Harrison (1961), [https://www.biodiversitylibrary.org/page/39011840 p. 16.]</ref> Karsten defined ''Hydnellum'' as having fruitbodies with a corky or leathery, tough cap, and a centrally attached stipe.<ref name="Karsten 1879"/> Synonyms of ''Hydnellum'' include ''Calodon'' (Karsten, 1881<ref name="Karsten 1881"/>), and ''Phaeodon'' (Joseph Schröter, 1888<ref name="Schröter 1888"/>).<ref name="urlMycoBank: Hydnellum"/>

''Hydnellum'' is classified in the family Bankeraceae, which was circumscribed by Marinus Anton Donk in 1961. The genus was not in Donk's original family concept, which included only ''Bankera'' and ''Phellodon'', genera whose species produce hyaline (translucent), and ornamented spores. Donk left ''Hydnellum'' in the tribe Hydnelleae of the family Thelephoraceae, along with ''Sarcodon'' and ''Hydnodon''.<ref name="Donk 1961"/> In 1981,<ref name="Jülich 1981"/> however, Walter Jülich emended Donk's concept of the Bankeraceae, adding hydnoid genera that produced brown, lobed spores—''Hydnellum'' and ''Sarcodon''.<ref>Stalpers (1993), p. 22.</ref>

The name originates from the Greek word ''ύδνο'' (=''hydnum'') meaning "spongy plant" or "fungus".<ref name="url BC NTFP Mushrooms"/> The British Mycological Society, in their recommended list of common names for fungi in the United Kingdom, name ''Hydnellum'' fungi in the form "descriptor word" plus "tooth", such as "gold tooth" (''H. auratile''), "zoned tooth" (''H. concrescens''), and "velvet tooth" (''H. spongiosipes'').<ref name="BMS"/>

==Phylogeny== Phylogenetic studies have shown that, in its current circumscription, the genus ''Hydnellum'' is polyphyletic and only distantly related to the white-spored genera ''Bankera'' and ''Phellodon'', currently grouped together in the family Bankeraceae.<ref name="Binder 2005"/><ref name="Tedersoo 2010"/><ref name="Baird 2013"/> Analyses of the ITS region by Parfitt and colleagues<ref name="Parfitt 2007"/> and Ainsworth and colleagues,<ref name="Ainsworth 2010"/> have further revealed multiple cryptic lineages within the genus. Most of the cryptic species belong to the ''H. concrescens''/''H. scrobiculatum'' species complex, resulting in frequent misinterpretations of these old and widely applied names.<ref name="Ainsworth 2010"/><ref name="Loizides 2016"/> At the moment, the number of phylogenetically distinct lineages confirmed by molecular sequencing exceeds the number of described taxa in the genus, making further taxonomic revisions inevitable in the future.

==Morphology== [[File:Gezoneerde stekelzwam (Hydnellum concrescens) Locatie, Hortus (Haren, Groningen) 02.JPG|thumb|right|The caps of some ''Hydnellum'' species (''H. concrescens'' pictured) can fuse together during growth.]] ''Hydnellum'' fruitbodies have caps and stipes, often with indeterminant growth forms, that may grow in spurts and decay over several weeks.<ref name="Baird 1986"/><ref name="Baird 2013"/> Neighboring fruitbodies can coalesce, forming intricately intertwined caps and partially fused stipes.<ref name="Harrison p.13"/> The flesh has a zoned appearance and is fibrous when fresh, but becomes hard and woody when dry.<ref name="Baird 1986"/> Zones in the flesh reflect differences in growth during periods of low daytime and high nighttime humidity, and give a fairly accurate record of daily growth.<ref name="Harrison 1968"/> The spines are crowded closely together and are typically decurrent (extending down the length of the stipe). Fruitbodies may display a variety of colors, from white to yellow, olive green, shades of orange, light brown, or dark brown in age.<ref name="Baird 1986"/>

Spores of ''Hydnellum'' are almost spherical to oblong and tuberculate, and are brown in mass.<ref name="Baird 1986"/><ref name="Ainsworth 2010"/> The basidia (spore-bearing cells) are narrowly club-shaped and usually four-spored; there are no cystidia in ''Hydnellum''.<ref name="Pegler 1997"/> Three types of hyphae are found in the flesh of ''Hydnellum'': generative hyphae (thin-walled, not inflated); skeletal hyphae (thick-walled and narrow); and thin-walled gloeoplerous-like hyphae, which stain with methyl blue.<ref>Stalpers (1997), p. 9.</ref> thumb|left|Spines of ''H. ferrugineum'' In conditions of high humidity, several species can form striking colored drops on the actively growing caps: red drops in ''H. peckii'', ''H. diabolus'', ''H. ferrugineum'', and ''H. cruentum'', yellow drops in ''H. caeruleum'', and coffee-colored drops in ''H. mirabile''.<ref name="Harrison p.15">Harrison (1961), [https://www.biodiversitylibrary.org/page/39011841 p. 15.]</ref> The common names of ''H. peckii'' reflect its appearance: "strawberries and cream" and "bleeding tooth fungus".<ref name="Arora 1986"/> Some ''Hydnellum'' species have a mealy odor (e.g. ''H. mirabile'' and ''H. pineticola'') similar to freshly ground flour. ''H. zonatum'' smells like melilot,<ref name="Rumack 1994"/> while ''H. suaveolens'' has a sweet odor resembling anise or peppermint. All are too tough and woody to be edible, and many have an acrid taste anyway.<ref name="Arora 1986"/>

Differences between ''Hydnellum'' species tend to be more distinguishable in younger specimens. Fruitbody development is greatly influenced by environmental factors such as levels of rainfall, drying winds, and temperature.<ref name="Harrison p.14"/> The blue tooth (''H. caeruleum''), for example, develops a deeper blue color when it grows during cooler autumn weather.<ref name="Harrison p.14"/> Optimal growth occurs during periods of frequent light rains and high humidity; if the habitat dries out, growth will stop, but may resume after further precipitation. This intermittent growth affects the fruitbodies of different species to variable extents, leading to large variations in form, surface texture, and color.<ref name="Harrison p.14"/> The morphological variability of fruitbodies and the dependence of their appearance upon environmental conditions has made ''Hydnellum'' a difficult group to study. Canadian mycologist Kenneth A. Harrison, who described several new species from North America, noted "[t]he remarkable longevity of individual sporophores of many species and the changes in appearance that occur during the long period of their development have confused all workers studying this group."<ref>Harrison (1961), [https://www.biodiversitylibrary.org/page/39011844 p. 12.]</ref> For example, ''H. aurantiacum'', initially white, becomes in turn shades of orange, rusty-brown, and brownish-black. Its fruitbody initially has a turbinate (cushion-like) shape with a lumpy surface, later becoming flattened to funnel-shaped with a smooth to corrugated surface texture.<ref name="Harrison p.13">Harrison (1961), [https://www.biodiversitylibrary.org/page/39011843 p. 13.]</ref> {{multiple image | header = Immature fruitbodies | align = center | image1 = Hydnellum compactum 01.JPG | width1 = {{#expr: (150 * 2048 / 1536) round 0}} | caption1 = ''H. compactum'' | image2 = Scharfer Korkstacheling Hydnellum peckii jung.jpg | width2 = {{#expr: (150 * 3000 / 2026) round 0}} | caption2 = ''H. peckii'' | image3 = Hydnellum ferrugineum 59273.jpg | width3 = {{#expr: (150 * 750 / 500) round 0}} | caption3 = ''H. ferrugineum'' }} The caps form from the top of the short stipe by the growth and expansion of a blunt margin and later as a thickening of the upper surface.<ref name="Harrison p.13"/> Spines start to form when the cap hangs over the stipe slightly. They are white in many species, but become brown in maturity as the brown-colored spores accumulate on the surface.<ref name="Harrison p.14">Harrison (1961), [https://www.biodiversitylibrary.org/page/39011842 p. 14.]</ref>

==Habitat and distribution== ''Hydnellum'' fungi are mycorrhizal, and are usually found in coniferous and mixed woods.<ref name="Ainsworth 2010"/><ref name="Pegler 1997"/> Favored tree hosts include members of the Fagaceae and the Pinaceae.<ref>Stalpers (1997), p. 34.</ref> The genus is widely distributed in the Northern Hemisphere, particularly Europe and North America,<ref name="Banker 1906"/> but some species are also found in tropical Asia.<ref name="Pegler 1997"/> Harrison identified a dozen new species from North America in the 1960s.<ref name="Harrison 1968"/><ref>Harrison (1961).</ref><ref name="Harrison 1964"/> Rudolph Arnold Maas Geesteranus recognized 16 European species in his 1975 treatment of the genus,<ref name="Maas Geesteranus 1975"/> to which ''H. dianthifolium'' has been recently added by Loizides and colleagues.<ref name="Loizides 2016"/>

Some ''Hydnellum'' species, including ''H. ferrugineum'' and ''H. scleropodium'', form a tough mat of mycelia in the humus and upper soil of pine forests. This mycelial mat grows larger with old trees, and can cover an area of several square meters. These areas generally lack dwarf shrubs and promote the vigorous growth of mosses; reindeer lichens often occur in the center of large mats. The presence of the fungus changes the nature of the soil, resulting in a thinner humus layer, decreased groundwater penetration, decreased soil pH, and increases in the level of root respiration as well as the quantity of roots. The fungus also decreases the organic carbon and nitrogen concentrations. Soil with the mycelium becomes more podzolized than the surrounding soil.<ref name="Hintikka 1967"/><ref name="Fisher 1972"/>

==Conservation== thumb|left|''Hydnellum ferrugineum'' is of conservation concern in Europe. Some ''Hydnellum'' species have been shown to be in decline in Europe, including the Czech Republic,<ref name="Hrouda 1999"/> the Netherlands,<ref name="Arnolds 1989"/> Norway,<ref name="Gulden 1992"/> and Scotland.<ref name="Newton 2002"/> In the United Kingdom, several are listed in the biodiversity action plan for stiped hydnoid fungi: ''H. aurantiacum'' is classified as critically endangered; ''H. caeruleum'', ''H. ferrugineum'' are listed as endangered, while ''H. concrescens'', ''H. spongiosipes'', ''H. peckii'', and ''H. scrobiculatum'' are considered vulnerable.<ref name="ENRR 2004"/> ''H. ferrugineum'' and ''H. peckii'' are sensitive to the increased nitrogen deposition resulting from clear-cutting, a forestry practice used in some areas of Europe.<ref name="Arnolds 2010"/>

Conservation efforts for ''Hydnellum'' are hindered by the fact that some species are difficult to discriminate in the field, making it hard to determine an appropriate conservation status.<ref name="Ainsworth 2010"/><ref name="Parfitt 2007"/> Techniques based on species-specific PCR primers and DNA extraction from soil have been developed to detect the mycelia of various ''Hydnellum'' species without having to rely on the presence of fruitbodies, which may help conservation efforts as well as improve understanding of below-ground ecology.<ref name="van der Linde 2008"/> Similar techniques have been used to show that, in the case of ''H. aurantiacum'' and ''H. caeruleum'', the fungus can persist below the ground for at least four years without producing fruitbodies.<ref name="van der Linde 2012"/>

==Bioactive compounds== thumb|right|250px|alt=2,5-dihydroxy-3,6-bis(4-hydroxyphenyl)-1,4-benzoquinone|Structural formula of the anticoagulant and antibacterial compound atromentin, found in ''H. peckii'' Several chemical compounds—some with unique biological activity—have been isolated and identified from ''Hydnellum'' species. For example, ''Hydnellum peckii'' contains atromentin, a pigment with anticoagulant properties similar to heparin.<ref name="Khanna 1965"/> 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''.<ref name="Zheng 2006"/>

Some species are used as dyes. ''Hydnellum caeruleum'', used in North America and Scandinavia to dye silk and wool, produces a range of colors including tan, blue, and forest-green, depending on the mordant that is used. ''Hydnellum peckii'' produces gray, brown, and olive colors.<ref name="Roberts 2011"/> Hydnuferrugin and hydnuferruginin are pigments responsible for the coloration of ''H. ferrugineum'' and ''H. zonatum''.<ref name="Gripenberg 1981"/> Geogenin is a yellow pigment found in ''H. geogenium''.<ref name="Steglich 1979"/>

Thelephoric acid is present in several ''Hydnellum'' species.<ref>Stalpers (1997), p. 7.</ref> This compound, derived metabolically from the shikimic acid pathway, inhibits the enzyme prolyl endopeptidase, which is involved in deteriorating certain neuropeptides that are believed to contribute to memory and learning.<ref name="Kwak 1999"/> ''Hydnellum caeruleum'' and ''H. concrescens'' have several ''para''-terphenyl derivatives named thelephantins (specifically, variants thelephantins I–N),<ref name="Quang 2004"/> some of which can inhibit the enzyme alpha-glucosidase.<ref name="Wang 2014"/> The compounds hydnellins A and B are terphenyls found in ''H. suaveolens'' and ''Hydnellum geogerirum''.<ref name="Hashimoto 2006"/> The chemicals responsible for the fragrant anise-like aroma of ''H. suaveolens'' have been identified as coumarin and ''para''-anisaldehyde.<ref name="DeShazer 1988"/>

==Species== Karsten's original 1879 circumscription of ''Hydnellum'' contained 19 species.<ref name="Karsten 1879"/> Joost Stalpers included 34 ''Hydnellum'' species in his 1993 monograph on the Thelephorales.<ref>Stalpers (1993).</ref> The tenth edition of the ''Dictionary of the Fungi'' (2008) indicated 38 species in the genus.<ref name="Kirk 2008"/> Larsson ''et al.'' included 12 species transferred to this genus from ''Sarcodon'' in 2019.<ref name="Larsson 2019">{{Cite journal|author=Larsson, Svantesson|author2=Miscevic, Kõljalg|author3=Larsson|date=2019|title=Reassessment of the generic limits for Hydnellum and Sarcodon (Thelephorales, Basidiomycota)|journal=MycoKeys|issue=54|pages=31–47|doi=10.3897/mycokeys.54.35386|pmid=31231164|pmc=6579789 |doi-access=free }}</ref> In 2024, Douch ''et al''. described four more species.<ref name="Douch 2024">{{Cite journal |last1=Douch |first1=James K. |last2=Vaughan |first2=Luke J. |last3=Cooper |first3=Jerry A. |last4=Holmes |first4=Gareth D. |last5=Robinson |first5=Richard |last6=Stefani |first6=Franck |last7=Idnurm |first7=Alexander |last8=May |first8=Tom W. |date=2024-08-29 |title=Taxonomic revision of fleshy species of ''Hydnellum'', ''Neosarcodon'', and ''Sarcodon'' (Thelephorales) from Australasia |url=https://www.tandfonline.com/doi/full/10.1080/00275514.2024.2363211 |journal=Mycologia |language=en |volume=116 |issue=6 |pages=965–992 |doi=10.1080/00275514.2024.2363211 |pmid=39208239 |issn=0027-5514 |url-access=subscription}}</ref>

{{As of|2025|August}}, Index Fungorum lists approximately 120 species of ''Hydnellum.''<ref name="Kirk 2014" /> Some of the species are listed below. {| class="wikitable sortable collapsible" |- ! Image !! Name !! Distribution |- |120px|| ''Hydnellum amygdaliolens'' (Rubio Casas, Rubio Roldán & Català) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum aurantiacum'' (Batsch) P.Karst. (1879) || Asia, Europe, North America<ref name="Pegler 1997"/> |- |120px|| ''Hydnellum auratile'' (Britzelm.) Maas Geest. (1959) || Europe, North America<ref name="Maas Geesteranus 1959"/> |- |120px|| ''Hydnellum caeruleum'' (Hornem.) P.Karst. (1879) || Asia, Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum chrysinum'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- |120px|| ''Hydnellum coactum'' (Y.H. Mu & H.S. Yuan) Della Magg. (2023) || China<ref name="Hu 2020"/> |- ||| ''Hydnellum coalitum'' Maas Geest. (1975) || Europe<ref name="Maas Geesteranus 1975"/> |- |120px|| ''Hydnellum compactum'' (Pers.) P.Karst. (1879) || Europe<ref name="Maas Geesteranus 1971"/> |- ||| ''Hydnellum complicatum'' Banker (1906) || North America<ref name="Banker 1906"/> |- |120px|| ''Hydnellum concrescens'' (Pers.) Banker (1906) || Asia, Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum conigenum'' (Peck) Banker (1906) || North America<ref name="McKnight 1998"/> |- |120px|| ''Hydnellum cristatum'' (Bres.) Stalpers (1993) || Europe, North America |- ||| ''Hydnellum cruentum'' K.A.Harrison (1961) || Nova Scotia, Canada<ref name="Harrison p.37">Harrison (1961), [https://www.biodiversitylibrary.org/page/39011819 p. 37.]</ref> |- ||| ''Hydnellum crustulinum'' Maas Geest. (1971) || Punjab, India<ref name="Maas Geesteranus 1971"/> |- ||| ''Hydnellum cumulatum'' K.A.Harrison (1964) || Europe,<ref name="Ainsworth 2011"/> North America<ref name="Harrison 1964"/> |- ||| ''Hydnellum cyanodon'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- |120px|| ''Hydnellum cyanopodium'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- |120px|| ''Hydnellum dianthifolium'' Loizides, Arnolds & P.-A. Moreau (2016) || Southern Europe<ref name="Loizides 2016"/> |- ||| ''Hydnellum earlianum'' Banker (1906) || North America<ref name="Banker 1906"/> |- |120px|| ''Hydnellum fennicum'' (P.Karst.) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum ferrugineum'' (Fr.) P.Karst. (1879) || North Africa, Asia, Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum floriforme'' (Schaeff.) Banker (1906) || North America<ref name="Banker 1906"/> |- ||| ''Hydnellum fraudulentum'' Maas Geest. (1971) || Australia<ref name="Maas Geesteranus 1971"/> |- ||| ''Hydnellum frondosum'' K.A.Harrison (1961) || Nova Scotia, Canada<ref name="Harrison p.45">Harrison (1961), [https://www.biodiversitylibrary.org/page/39011798 p. 45.]</ref> |- |120px|| ''Hydnellum fuligineoviolaceum'' (Kalchbr.) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum fuscoindicum'' (K.A.Harrison) E.Larss., K.H.Larss. & Kõljalg || North America<ref name="Larsson 2019"/> |- ||| ''Hydnellum gatesiae'' Douch, L.J. Vaughan & T.W. May || Australia<ref name="Douch 2024"/> |- |120px|| ''Hydnellum geogenium'' (Fr.) Banker (1913) || Europe, North America<ref name="urlRogers: H.geogenium"/> |- |120px|| ''Hydnellum glaucopus'' (Maas Geest. & Nannf.) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum gracilipes'' (P.Karst.) P.Karst. (1879) || Europe<ref name="Kõljalg 2000"/> |- |120px|| ''Hydnellum grosselepidotum'' Y.H. Mu & H.S. Yuan (2020) || China<ref name="Hu 2020"/> |- | |''Hydnellum hangzhouense'' (X.P. Fan, Y.F. Sun & B.K. Cui) |Eastern China<ref>{{Cite journal |last1=Fan |first1=Xiang-Ping |last2=Song |first2=Chang-Ge |last3=Xu |first3=Yi-Hua |last4=Sun |first4=Yi-Fei |last5=Cui |first5=Bao-Kai |date=2025-07-22 |title=Morphology and molecular phylogeny reveal one new species of Hydnellum (Thelephorales) from Eastern China |url=https://phytotaxa.mapress.com/pt/article/view/phytotaxa.711.1.3 |journal=Phytotaxa |language=en |volume=711 |issue=1 |pages=43–60 |doi=10.11646/phytotaxa.711.1.3 |issn=1179-3163|url-access=subscription }}</ref> |- |120px|| ''Hydnellum joeides'' (Pass.) E.Larss., K.H.Larss. & Kõljalg || Europe, North America<ref name="Larsson 2019"/> |- ||| ''Hydnellum lepidum'' (Maas Geest.) E. Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum lidongensis'' (Y.H. Mu & H.S. Yuan) Della Magg. (2023) || China<ref name="Hu 2020"/> |- ||| ''Hydnellum longidentatum'' Coker (1939) || United States<ref name="Coker 1939"/> |- ||| ''Hydnellum lundellii'' (Maas Geest. & Nannf.) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum martioflavum'' (Snell, K.A.Harrison & H.A.C.Jacks.) E.Larss., K.H.Larss. & Kõljalg || Europe, North America<ref name="Larsson 2019"/> |- |120px|| ''Hydnellum mirabile'' (Fr.) P.Karst. (1879) || Europe, North America<ref name="urlRogers: H. mirabile"/> |- ||| ''Hydnellum multiceps'' K.A.Harrison (1961) || Nova Scotia, Canada<ref name="Harrison p.45"/> |- ||| ''Hydnellum nigellum'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- ||| ''Hydnellum nothofagacearum'' Douch & J.A. Cooper || New Zealand<ref name="Douch 2024"/> |- ||| ''Hydnellum papuanum'' Maas Geest. (1971) || Papua New Guinea<ref name="Maas Geesteranus 1971"/> |- ||| ''Hydnellum parvum'' Banker (1913) || North America<ref name="Baird 2013"/> |- |120px|| ''Hydnellum peckii'' Banker (1912) || Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum pseudoioeides'' Douch & J.A. Cooper || Oceania<ref name="Douch 2024"/> |- |120px|| ''Hydnellum regium'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- ||| ''Hydnellum rickeri'' Banker (1913) || North America<ref name="Banker 1913"/> |- |120px|| ''Hydnellum scabrosum'' (Fr.) E.Larss., K.H.Larss. & Kõljalg || Europe<ref name="Larsson 2019"/> |- ||| ''Hydnellum scleropodium'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- |120px|| ''Hydnellum scrobiculatum'' (Fr.) P.Karst. (1879) || Asia, Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum septentrionale'' K.A.Harrison (1964) || North America<ref name="Harrison 1964"/> |- ||| ''Hydnellum singeri'' Maas Geest. (1969) || Colombia<ref name="Maas Geesteranus 1969"/> |- |120px|| ''Hydnellum spongiosipes'' (Peck) Pouzar (1960) || Europe, North America<ref name="Pegler 1997"/> |- ||| ''Hydnellum staurastrum'' Maas Geest. (1971) || Malaysia<ref name="Maas Geesteranus 1971"/> |- |120px|| ''Hydnellum suaveolens'' (Scop.) P.Karst. (1879) || Asia, Europe, North America<ref name="urlRogers: H.suaveolens"/> |- ||| ''Hydnellum subzonatum'' K.A.Harrison (1961) || Nova Scotia, Canada<ref name="Harrison p.37"/> |- ||| ''Hydnellum tardum'' Maas Geest. (1975) || Europe<ref name="Maas Geesteranus 1975"/> |- |120px|| ''Hydnellum underwoodii'' (Banker) E.Larss., K.H.Larss. & Kõljalg || North America<ref name="Larsson 2019"/> |- ||| ''Hydnellum variisporum'' Douch, Robinson & L.J. Vaughan || Australia<ref name="Douch 2024"/> |- |120px|| ''Hydnellum versipelle'' (Fr.) E.Larss., K.H.Larss. & Kõljalg || Europe, North America<ref name="Larsson 2019"/> |- |}

==References== {{Reflist|30em|refs=

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<ref name="Baird 1986">{{cite journal |vauthors=Baird RE, Khan SR |year=1986 |title=The stipitate Hydnums (Thelephoraceae) of Florida USA |journal=Brittonia |volume=38 |issue=2 |pages=171–84 |jstor=2807273 |doi=10.2307/2807273|s2cid=85360191 }}</ref>

<ref name="Baird 2013">{{cite journal |vauthors=Baird R, Wallace L, Baker G, Scruggs M |title=Stipitate hydnoid fungi of the temperate southeastern United States |journal=Fungal Diversity|year=2013 |volume=62 |pages=41–114 |doi=10.1007/s13225-013-0261-6|s2cid=16846274 }}</ref>

<ref name="Banker 1906">{{cite journal |author=Banker HJ |title=A contribution to a revision of the North American Hydnaceae. |journal=Memoirs of the Torrey Botanical Club.|year=1906 |volume=12 |issue=2 |pages=99–194}}</ref>

<ref name="Banker 1913">{{cite journal |author=Banker HJ |title=Type studies in the Hydnaceae – V. The genus ''Hydnellum'' |journal=Mycologia |year=1913 |volume=5 |issue=4 |pages=194–205 |url=http://www.cybertruffle.org.uk/cyberliber/59350/0005/004/0194.htm |doi=10.2307/3753385|jstor=3753385 |url-access=subscription }}</ref>

<ref name="Binder 2005">{{cite journal |vauthors=Binder M, Hibbett DS, Larsson KH, Larsson E, Langer E, Langer G |year=2005 |title=The phylogenetic distribution of resupinate forms across the major clades of mushroom-forming fungi (Homobasidiomycetes). |journal=Systematics and Biodiversity |volume=3 |issue=2 |pages=113–157 |doi=10.1017/s1477200005001623|s2cid=13102957 }}</ref>

<ref name="BMS">{{cite web |url=http://www.fungi4schools.org/Reprints/ENGLISH_NAMES.pdf |title=Recommended English Names for Fungi in the UK |publisher=British Mycological Society |url-status=dead |archive-url=https://web.archive.org/web/20110716083053/http://www.fungi4schools.org/Reprints/ENGLISH_NAMES.pdf |archive-date=2011-07-16 }}</ref>

<ref name="Coker 1939">{{cite journal |author=Coker WC |title=New or noteworthy Basidiomycetes |journal=Journal of the Elisha Mitchell Scientific Society |year=1939 |volume=55 |pages=373–86}}</ref>

<ref name="DeShazer 1988">{{cite journal |vauthors=Wood WF, DeShazer DA, Largent DL |title=The identify and metabolic fate of volatiles responsible for the odor of ''Hydnellum suaveolens'' |journal=Mycologia |year=1988 |volume=80 |issue=2 |pages=252–5 |url=http://www.cybertruffle.org.uk/cyberliber/59350/0080/002/0252.htm | doi = 10.2307/3807803 |jstor=3807803 |url-access=subscription }}</ref>

<ref name="Donk 1961">{{cite journal |author=Donk MA |title=Four new families of Hymenomycetes |year=1961 |journal=Persoonia |volume=1 |issue=4 |pages=405–7 |url=http://www.cybertruffle.org.uk/cyberliber/61056/0001/004/0405.htm}}</ref>

<ref name="ENRR 2004">{{cite report |vauthors=Bridge PD, Panchal G |title=Number 557. Population diversity and speciation in ''Hydnellum'' and ''Phellodon'' species |series=English Nature Research Reports |year=2004 |publisher=English Nature |url=http://publications.naturalengland.org.uk/file/6167520566312960 |format=PDF |issn=0967-876X}}</ref>

<ref name="Fisher 1972">{{cite journal |author=Fisher RF |title=Spodosol development and nutrient distribution under Hydnaceae fungal mats |journal= Soil Science Society of America Journal|year=1972 |volume=36 |issue=3 |pages=492–5 |doi=10.2136/sssaj1972.03615995003600030035x|bibcode=1972SSASJ..36..492F }}</ref>

<ref name="Gripenberg 1981">{{cite journal |author=Gripenberg J. |title=Fungus pigments. XXIX. The pigments of ''Hydnellum ferrugineum'' (Fr.) Karsten and ''H. zonatum'' (Batsch) Karsten |journal=Acta Chemica Scandinavica B |year=1981 |volume=35 |pages=513–9 |url=http://actachemscand.org/pdf/acta_vol_35b_p0513-0519.pdf|doi=10.3891/acta.chem.scand.35b-0513 |doi-access=free }}</ref>

<ref name="Gulden 1992">{{cite journal |vauthors=Gulden G, Hanssen EW |author-link=Gro Gulden |title=Distribution and ecology of stipitate hydnaceous fungi in Norway, with special reference to the question of decline |journal=Sommerfeltia |year=1992 |volume=13 |pages=1–58|doi=10.2478/som-1992-0001 |s2cid=236991359 |doi-access=free }}</ref>

<ref name="Harrison 1964">{{cite journal |author=Harrison KA |title=New or little known North American stipitate Hydnums |journal=Canadian Journal of Botany |year=1964 |volume=42 |issue=9 |pages=1205–33 |doi=10.1139/b64-116}}</ref>

<ref name="Harrison 1968">{{cite journal |author=Harrison KA |title=Studies on the Hydnums of Michigan I. Genera ''Phellodon'', ''Bankera'', ''Hydnellum'' |journal=Michigan Botanist |year=1968 |volume=7 |pages=212–64 |url=http://babel.hathitrust.org/cgi/pt?id=mdp.39015005837763;view=1up;seq=646}}</ref>

<ref name="Hashimoto 2006">{{cite journal |vauthors=Hashimoto T, Quang DN, Kuratsune M, Asakawa Y |title=Hydnellins A and B, nitrogen-containing terphenyls from the mushrooms ''Hydnellum suaveolens'' and ''Hydnellum geogerirum'' |journal=Chemical and Pharmaceutical Bulletin |year=2006 |volume=54 |issue=6 |pages=912–4 |doi=10.1248/cpb.54.912|pmid=16755070 |doi-access=free }}{{open access}}</ref>

<ref name="Hintikka 1967">{{cite journal |vauthors=Hintikka V, Näykki O |title=Tutkimuksia ruosteorakkaan, ''Hydnellum ferrugineum'', vaikutuksesta metsämaaperään ja-kasvillisuuteen [Notes on the effects of the fungus ''Hydnellum ferrugineum'' (Fr.) Karst. on forest soil and vegetation] |journal=Communicationes Instituti Forestalis Fenniae |year=1967 |volume=62 |issue=2 |pages=1–23}}</ref>

<ref name="Hrouda 1999">{{cite journal |author=Hrouda P. |title=Hydnaceous fungi of the Czech Republic and Slovakia |journal=Czech Mycology |year=1999 |volume=51 |issue=2–3 |pages=99–155 |url=http://www.sci.muni.cz/botany/mycology/hydna.htm|doi=10.33585/cmy.51202 |url-access=subscription }}</ref>

<ref name="Hu 2020">{{cite journal | last1=Mu | first1=Yan-Hong | last2=Hu | first2=Ya-Ping | last3=Wei | first3=Yu-Lian | last4=Yuan | first4=Hai-Sheng | title=Hydnaceous fungi of China 8. Morphological and molecular identification of three new species of Sarcodon and a new record from southwest China | journal=MycoKeys | date=2020 | issue=66 | pages=83–103 | doi=10.3897/mycokeys.66.49910 | doi-access=free | pmid=32296287 | pmc=7148386 }}</ref>

<ref name="Jülich 1981">{{cite book |author=Jülich W. |year=1981 |title=Higher taxa of Basidiomycetes |series=Bibliotheca Mycologica |volume=85 |location=Vaduz, Liechtenstein |publisher=J. Cramer |isbn=978-3-7682-1324-0}}</ref>

<ref name="Karsten 1879">{{cite journal |author=Karsten PA |title=Symbolae ad mycologiam Fennicam. VI |journal=Meddelanden Af Societas Pro Fauna et Flora Fennica |year=1879 |volume=5 |pages=5–46 (see p. 41) |url=https://www.biodiversitylibrary.org/page/3768952}}</ref>

<ref name="Karsten 1881">{{cite journal |author=Karsten PA |title=Enumeratio Hydnearum Fr. Fennicarum, systemate novo dispositarum |year=1881 |journal=Revue Mycologique Toulouse |volume=3 |issue=9 |pages=19–21 |url=https://www.biodiversitylibrary.org/page/11599314 |language=la}}</ref>

<ref name="Khanna 1965">{{cite journal |vauthors=Khanna JM, Malone MH, Euler KL, Brady LR |year=1965 |title=Atromentin – anticoagulant from '' Hydnellum diabolus'' |journal=Journal of Pharmaceutical Sciences |volume=54 |issue=7 |pages=1016–20 |pmid=5862512 |doi=10.1002/jps.2600540714}}</ref>

<ref name="Kirk 2008">{{cite book |vauthors=Kirk PM, Cannon PF, Minter DW, Stalpers JA |title=Dictionary of the Fungi |edition=10th |publisher=CAB International |location=Wallingford, UK |year=2008 |page=324 |isbn=978-0-85199-826-8}}</ref>

<ref name="Kirk 2014">{{cite web |author=Kirk PM |title=Species Fungorum (version 22nd December 2014). In: Species 2000 & ITIS Catalogue of Life |url=http://www.catalogueoflife.org/col/browse/tree/id/20915520 |access-date=2015-01-03 |archive-date=2015-01-04 |archive-url=https://web.archive.org/web/20150104071923/http://www.catalogueoflife.org/col/browse/tree/id/20915520 |url-status=dead }}</ref>

<ref name="Kõljalg 2000">{{cite journal |vauthors=Kõljalg U, Renvall P |title=''Hydnellum gracilipes'' – a link between stipitate and resupinate Hymenomycetes |journal=Karstenia |volume=40 |issue=1–2 |pages=71–77|doi=10.29203/ka.2000.355 |year=2000 |url=http://www.funga.fi/Karstenia/Karstenia%2040-1%202000-14.pdf |doi-access=free }}</ref>

<ref name="Kwak 1999">{{cite journal |vauthors=Kwak JY, Rhee IK, Lee KB, Hwang JS, Yoo ID, Song KS |title=Thelephoric acid and kynapcin-9 in mushroom ''Polyozellus multiplex'' inhibit prolyl endopeptidase in vitro |journal=Journal of Microbiology and Biotechnology |year=1999 |volume=9 |issue=6 |pages=798–803}}</ref>

<ref name="Loizides 2016">{{cite journal |vauthors=Loizides M, Alvarado P, Assyov B, Arnolds E, Moreau PA |title=''Hydnellum dianthifolium'' sp. nov. (Basidiomycota, Thelephorales), a new tooth-fungus from southern Europe with notes on ''H. concrescens'' and ''H. scrobiculatum'' |journal=Phytotaxa |year=2016 |volume=280 |issue=1 |pages=23–35 |doi=10.11646/phytotaxa.280.1.2}}</ref>

<ref name="Maas Geesteranus 1959">{{cite journal |author=Maas Geesteranus RA. |title=Sur un ''Hydnellum'' méconnu |journal=Persoonia |year=1959 |volume=1 |issue=1 |pages=111–4 |url=http://www.cybertruffle.org.uk/cyberliber/61056/0001/001/0111.htm |language=fr}}</ref>

<ref name="Maas Geesteranus 1969">{{cite journal |author=Maas Geesteranus RA. |title=Notes on Hydnums, VIII |journal=Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen Section C |year=1969 |volume=72 |pages=213–21}}</ref>

<ref name="Maas Geesteranus 1971">{{cite journal |author=Maas Geesteranus RA. |title=Hydnaceous fungi of the eastern old world |journal=Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen Afdeling Natuurkunde |year=1971 |volume=60 |issue=3 |pages=1–176 (see pp. 91–106)}}</ref>

<ref name="Maas Geesteranus 1975">{{cite journal |author=Maas Geesteranus RA. |title=Die terrestrischen Stachelpilze Europas |journal=Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen Afdeling Natuurkunde |year=1975 |volume=64 |issue=2 |pages=1–127 (see pp. 95–98) |language=de}}</ref>

<ref name="McKnight 1998">{{cite book |author=McKnight KH |title=A Field Guide to Mushrooms: North America |url=https://books.google.com/books?id=kSdA3V7Z9WcC&pg=PA94 |year=1998 |publisher=Houghton Mifflin Harcourt |isbn=978-0-395-91090-0 |page=94}}</ref>

<ref name="Newton 2002">{{cite journal |vauthors=Newton AC, Holden E, Davy LM, Ward SD, Fleming LV, Watling R |year=2002 |title=Status and distribution of stipitate hydnoid fungi in Scottish coniferous forests |journal=Biological Conservation |volume=107 |issue=2 |pages=181–92 |doi=10.1016/S0006-3207(02)00060-5}}</ref>

<ref name="Parfitt 2007">{{cite journal |vauthors=Parfitt D, Ainsworth AM, Simpson D, Rogers HJ, Boddy L |year=2007 |title=Molecular and morphological discrimination of stipitate hydnoids in the genera ''Hydnellum'' and ''Phellodon'' |journal=Mycological Research |volume=111 |issue=7 |pages=761–77 |pmid=17681224 |doi=10.1016/j.mycres.2007.05.003}}</ref>

<ref name="Pegler 1997">{{cite book |vauthors=Pegler DN, Roberts PJ, Spooner BM |title=British Chanterelles and Tooth Fungi |year=1997 |publisher=Royal Botanic Gardens |location=Kew, UK |isbn=978-1-900347-15-0 |pages=74–88}}</ref>

<ref name="Quang 2004">{{cite journal |vauthors=Quang DN, Hashimoto T, Hitaka Y, Tanaka M, Nukada M, Yamamoto I, Asakawa Y |title=Thelephantins I-N; ''p''-terphenyl derivatives from the inedible mushroom ''Hydnellum caeruleum'' |journal=Phytochemistry |year=2004 |volume=65 |issue=8 |pages=1179–84 |pmid=15110701 |doi=10.1016/j.phytochem.2004.02.018|bibcode=2004PChem..65.1179Q }}</ref>

<ref name="Roberts 2011">{{cite book |vauthors=Roberts P, Evans S |title=The Book of Fungi |year=2011 |publisher=University of Chicago Press |location=Chicago, Illinois |pages=469–70 |isbn=978-0-226-72117-0}}</ref>

<ref name="Rumack 1994">{{cite book |vauthors=Rumack BH, Spoerke DG |title=Handbook of Mushroom Poisoning: Diagnosis and Treatment |url=https://books.google.com/books?id=WPWsZNvOqVAC&pg=PA413 |year=1994 |publisher=CRC Press |isbn=978-0-8493-0194-0 |page=413}}</ref>

<ref name="Schröter 1888">{{cite book |author=Schröter J. |title=Kryptogamen-Flora von Schlesien |year=1888 |volume=3–1(4) |publisher=Cramer |location=Lehre, Germany |page=458}}</ref>

<ref name="Steglich 1979">{{cite journal |vauthors=Steglich W, Jendrny B |title=Geogenin, ein neuartiger Pyronfarbstoff aus ''Hydnellum geogenium'' |trans-title=Geogenin, a new kind of pyrone pigment from ''Hydnellum geogenium'' |journal=Beihefte zur Sydowia Annales Mycologici |series=Series II |year=1979 |pages=378–80 |language=de}}</ref>

<ref name="Tedersoo 2010">{{cite journal |vauthors=Tedersoo L, May T, Smith ME |title=Ectomycorrhizal lifestyle in fungi: global diversity, distribution, and evolution of phylogenetic lineages. |journal=Mycorrhiza |year=2010 |volume=20 |issue=4 |pages=217–263 |doi=10.1007/s00572-009-0274-x |pmid=20191371|s2cid=3351967 }}</ref>

<ref name="url BC NTFP Mushrooms">{{cite web |url=http://bcmushrooms.forrex.org/ntfp/pages/hydnumrepan/hydnumrepan_image.html |title=Welcome to BC NTFP Mushrooms |vauthors=Gamiet S, Berch SM, Kroeger P, Roberts C, Winder R, MacKinnon A |work= BC NTFP mushrooms |publisher=Southern Interior Forest Extension and Research Partnership|access-date=2010-02-12}}</ref>

<ref name="urlMycoBank: Hydnellum">{{cite web |title=''Hydnellum'' P. Karst. 1879 |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=17781 |publisher=MycoBank. International Mycological Association |access-date=2012-03-25}}</ref>

<ref name="urlRogers: H.geogenium">{{cite web|author=Phillips R |title=''Hydnellum geogenium'' |publisher=RogersMushrooms |url=http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6117~gid~~source~gallerydefault.asp |access-date=2015-01-06 |url-status=dead |archive-url=https://web.archive.org/web/20150122080910/http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6117~gid~~source~gallerydefault.asp |archive-date=2015-01-22 }}</ref>

<ref name="urlRogers: H. mirabile">{{cite web|author=Phillips R |title=''Hydnellum mirabile'' |publisher=RogersMushrooms |url=http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6118~gid~~source~gallerydefault.asp |access-date=2015-01-06 |url-status=dead |archive-url=https://web.archive.org/web/20141129034610/http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6118~gid~~source~gallerydefault.asp |archive-date=2014-11-29 }}</ref>

<ref name="urlRogers: H.suaveolens">{{cite web|author=Phillips R |title=''Hydnellum suaveolens'' |publisher=RogersMushrooms |url=http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6124~gid~~source~gallerydefault.asp |access-date=2015-01-06 |url-status=dead |archive-url=https://web.archive.org/web/20150402200708/http://www.rogersmushrooms.com/gallery/DisplayBlock~bid~6124~gid~~source~gallerydefault.asp |archive-date=2015-04-02 }}</ref>

<ref name="van der Linde 2008">{{cite journal |vauthors=van der Linde S, Alexander I, Anderson IC |year=2008 |title=A PCR-based method for detecting the mycelia of stipitate hydnoid fungi in soil |journal=Journal of Microbiological Methods |volume=75 |issue=1 |pages=40–46 |doi=10.1016/j.mimet.2008.04.010 |pmid=18586344}}</ref>

<ref name="van der Linde 2012">{{cite journal |vauthors=van der Linde S, Holden E, Parkin PI, Alexander IJ, Anderson IC |title=Now you see it, now you don't: The challenge of detecting, monitoring and conserving ectomycorrhizal fungi |journal=Fungal Ecology |year=2012 |volume=5 |issue=5 |pages=633–40 |doi=10.1016/j.funeco.2012.04.002}}</ref>

<ref name="Wang 2014">{{cite journal |vauthors=Wang SM, Han JJ, Ma K, Jin T, Bao L, Pei YF, Liu HW |title=New α-glucosidase inhibitors with ''p''-terphenyl skeleton from the mushroom ''Hydnellum concrescens'' |journal=Fitoterapia |year=2014 |volume=98 |pages=149–55 |doi=10.1016/j.fitote.2014.07.019 |pmid=25088970}}</ref>

<ref name="Zheng 2006">{{cite journal |vauthors=Zheng CJ, Sohn MJ, Kim WG |title=Atromentin and leucomelone, the first inhibitors specific to enoyl-ACP reductase (FabK) of ''Streptococcus pneumoniae'' |journal=Journal of Antibiotics |year=2006 |volume=59 |issue=12 |pages=808–12 |doi=10.1038/ja.2006.108 |pmid=17323650|doi-access=free }}</ref>

}}

===Cited works=== *{{cite report |author=Harrison KA |title=The Stipitate Hydnums of Nova Scotia |work=Publications of the Department of Agriculture Canada |location=Ottawa, Canada |publisher=Research Branch, Canada Department of Agriculture |year=1961 |volume=1099 |pages=1–60 |url=https://www.biodiversitylibrary.org/page/39011849}} {{open access}} *{{cite journal |author=Stalpers JA |year=1993 |title=The Aphyllophoraceous fungi I. Keys to the species of the Thelephorales |journal=Studies in Mycology |volume=35 |pages=1–168 |url=http://www.cbs.knaw.nl/publications/Sim35/full%20text.htm |access-date=2015-01-05 |archive-date=2015-02-21 |archive-url=https://web.archive.org/web/20150221225508/http://www.cbs.knaw.nl/publications/Sim35/full%20text.htm |url-status=dead }}

==External links== * {{Commons category-inline|Hydnellum|''Hydnellum''}} * {{Wikispecies-inline|Hydnellum|''Hydnellum''}}

Category:Hydnellum Category:Thelephorales genera Category:Taxa named by Petter Adolf Karsten Category:Taxa described in 1879