Phellodon

{{Short description|Genus of tooth fungi in the family Bankeraceae}} {{Use dmy dates|date=November 2019}} {{Automatic taxobox | image = Phellodon niger 496570.jpg | image_caption = ''Phellodon niger'' | taxon = Phellodon | authority = P.Karst. (1881) | type_species = ''Phellodon niger'' | type_species_authority = (Fr.) P.Karst. (as 'nigrum') (1881) | subdivision_ranks = Species | subdivision = {{center|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.

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'').<ref name="Karsten 1881"/> ''P. nigrum'' was originally described by Elias Fries in 1815 as a species of ''Hydnum''.<ref name="Fries 1815"/> Some early authors did not consider ''Phellodon'' distinct enough to be a separate genus, and folded species assigned to this genus into ''Hydnellum''.<ref name="Miller 1933"/><ref name="Ragab 1953"/>

''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.<ref name="Donk 1961"/> 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.<ref name="Harrison 1972"/> 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.<ref name="Maas Geesteranus 1974"/> 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.<ref name="Baird 1987"/> 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,<ref name="Binder 2005"/> ''Phellodon'' is classified in this family by authorities on fungal taxonomy.<ref name="Cannon 2007"/><ref name="Kirk 2008"/><ref name="urlMycoBank: Phellodon"/>

The generic name is derived from Greek {{Lang|grc-latn|phell}}-, meaning 'cork', and -{{Lang|grc-latn|don}}, meaning 'tooth'. In North America, ''Phellodon'' species are commonly known as "cork hydnums".<ref name="McKnight 1998"/> 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'').<ref name="BMS"/>

==Description== [[File:Phellodon tomentosus I Slavkovsky les CZ.jpg|thumb|left|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.<ref name="Stalpers 1993"/> Neighboring fruitbodies can fuse, forming intertwined caps and partially fused stipes.<ref name="Pegler 1997"/> 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."<ref name="Smith 1975"/> ''Phellodon'' fruitbodies can envelop nearby grass or twigs. The stipe is thickly tomentose or smooth, typically the same color as the cap or darker.<ref name="Zhishu 1993"/> In ''P. niger'', the outer covering of the stipe is a thick felty layer of mycelium that absorbs water like a sponge.<ref name="Harrison 1961"/> The hymenophore (the fertile, spore-bearing surface) is on the underside of the cap. The spines become grey at maturity.<ref name="Zhishu 1993"/> In conditions of high humidity, ''P. niger'' can form striking drops of black liquid on the actively growing caps.<ref name="Harrison 1961"/> thumb|right|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.<ref name="Stalpers 1993"/> 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.<ref name="Pegler 1997"/> ''Phellodon'' species are often free of insect damage, suggesting that they may have defensive chemicals that deter predation.<ref name="Marren 2000"/> Fruitbodies are not considered edible due to their fibrous flesh.<ref name="Smith 1975"/>

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,<ref name="Pegler 1997"/> but they are present in the North American species ''P. fibulatus''<ref name="Harrison 1972"/> and ''P. mississippiensis''.<ref name="Baird 2013"/> The basidia (spore-bearing cells) are four-spored.<ref name="Zhishu 1993"/> Cystidia are either absent, or present infrequently as incompletely differentiated cystidioles (sterile cells about the size of an immature basidium).<ref name="Pegler 1997"/> In mass, the spores are white.<ref name="McKnight 1998"/> Spores are broadly ellipsoid to roughly spherical, and echinulose (covered with small spines). They are also hyaline and inamyloid.<ref name="Zhishu 1993"/>

==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.<ref name="Stalpers 1993"/> 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.<ref name="Agerer 1992"/> 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.<ref name="Högberg 1999"/><ref name="Taylor 2003"/>

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.<ref name="Baird 2013"/><ref name="Baird 2013b"/>

==Conservation== thumb|left|''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.<ref name="Stalpers 1993"/> 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.<ref name="ENRR 2004"/>

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.<ref name="van der Linde 2009"/> ''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.<ref name="van der Linde 2008"/> DNA testing of collections labelled as ''P. melaleucus'' and ''P. niger'' from the United Kingdom revealed additional cryptic species.<ref name="Parfitt 2007"/><ref name="Ainsworth 2010"/> 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.<ref name="van der Linde 2012"/>

==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.<ref name="Stadler 1993"/> A total synthesis was described for phellodonic acid in 2008 using ''cis''-1,2-dihydrocatechol as the starting material.<ref name="Reekie 2008"/> ''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<nowiki>''</nowiki>-pentahydroxy-''p''-terphenyl; grifolin; and 4-''O''-methylgrifolic acid.<ref name="Fang 2010"/> ''P. niger'' has also been used for mushroom dyeing, in which it produces gray-blue and green colors.<ref name="Roberts 2011"/>

==Species== ''Phellodon'' was originally circumscribed with three species. Joost Stalpers included 13 ''Phellodon'' species in his 1993 monograph on the Thelephorales.<ref name="Stalpers 1993"/> The tenth edition of the Dictionary of the Fungi (2008) indicated 16 species in the genus.<ref name="Kirk 2008"/> {{As of|2015|September}}, Index Fungorum lists 18 species of ''Phellodon'',<ref name="Kirk 2015"/> not including the three eastern United States species added in 2013–14.<ref name="Baird 2013"/><ref name="Baird 2013b"/> {| class="wikitable" |- ! Image !! Scientific name !! Distribution

|- |120px||''Phellodon atratus'' K.A.Harrison (1964) ||California, United States<ref name="Harrison 1964"/> |- |||''Phellodon brunneo-olivaceus'' R.E.Baird (2013) ||United States<ref name="Baird 2013b"/> |- |||''Phellodon confluens'' (Pers.) Pouzar (1956) ||China,<ref name="Dai 2011"/> eastern United States,<ref name="Baird 1986b"/> Europe<ref name="Pouzar 1956"/> |- |||''Phellodon excentrimexicanus'' R.E.Baird (1985) ||Mexico<ref name="Baird 1985"/> |- |||''Phellodon fibulatus'' K.A.Harrison (1972) ||North Carolina, United States<ref name="Harrison 1972"/> |- |||''Phellodon fuligineoalbus'' (J.C.Schmidt) Baird (2013) ||United States<ref name="Baird 2013b"/> |- |||''Phellodon implicatus'' R.E.Baird & S.R.Khan (1986) ||Florida, United States<ref name="Baird 1986"/> |- |||''Phellodon indicus'' Khara (1978) ||Himachal Pradesh, India<ref name="Khara 1978"/> |- |||''Phellodon maliensis'' (Lloyd) Maas Geest. (1966) ||Australia, New Zealand<ref name="Maas Geesteranus 1971"/> |- |120px||''Phellodon melaleucus'' (Sw. ex Fr.) P.Karst. (1881) ||Europe, North America<ref name="Pegler 1997"/> |- |||''Phellodon mississippiensis'' R.Baird (2014) ||Mississippi, United States<ref name="Baird 2013"/> |- |120px||''Phellodon niger'' (Fr.) P.Karst. (1881) ||Europe, North America<ref name="Pegler 1997"/> |- |||''Phellodon nothofagi'' McNabb (1971) ||New Zealand<ref name="McNabb 1971"/> |- |||''Phellodon plicatus'' (Lloyd) Maas Geest. (1966) ||Australia<ref name="Maas Geesteranus 1966"/> |- |||''Phellodon putidus'' (G.F.Atk.) Banker (1906) ||North America<ref name="Banker 1906"/> |- |||''Phellodon radicatus'' R.E.Baird (1985) ||North America<ref name="Baird 1985"/> |- |||''Phellodon rufipes'' Maas Geest. (1971) ||Japan<ref name="Maas Geesteranus 1971"/> |- |120px||''Phellodon secretus'' Niemelä & Kinnunen (2003) –Finland<ref name="Niemelä 2003"/> |- |120px||''Phellodon sinclairii'' (Berk.) G.Cunn. (1958) ||New Zealand<ref name="Cunningham 1958"/> |- |||''Phellodon tenuis'' R.E.Baird (1988) ||Brazil<ref name="Baird 1988"/> |- |120px||''Phellodon tomentosus'' (L.) Banker (1906) ||Europe,<ref name="Pegler 1997"/> North America<ref name="Banker 1906"/> |- |}

==References== {{Reflist|refs=

<ref name="Agerer 1992">{{cite journal |author=Agerer R. |title=Ectomycorrhizae of ''Phellodon niger'' on Norway spruce and their chlamydospores |journal=Mycorrhiza |year=1992 |volume=2 |issue=1 |pages=47–52 |doi=10.1007/BF00206283|s2cid=11608006 }}</ref>

<ref name="Ainsworth 2010">{{cite journal |vauthors=Ainsworth AM, Parfitt D, Rogers HJ, Boddy L |title=Cryptic taxa within European species of ''Hydnellum'' and ''Phellodon'' revealed by combined molecular and morphological analysis |journal=Fungal Ecology |year=2010 |volume=3 |issue=2 |pages=65–80 |doi=10.1016/j.funeco.2009.07.001}}</ref>

<ref name="Baird 1985">{{cite journal |author=Baird RE. |title=New species of stipitate hydnums from southeastern United States and Mexico |journal=Mycotaxon |year=1985 |volume=23 |pages=297–304 |url=http://www.cybertruffle.org.uk/cyberliber/59575/0023/0297.htm}}</ref>

<ref name="Baird 1986">{{cite journal |vauthors=Baird RE, Khan SR |title=The stipitate hydnums (Thelephoraceae) of Florida |journal=Brittonia |year=1986 |volume=30 |issue=2 |pages=171–184 |jstor=2807273 |doi=10.2307/2807273|s2cid=85360191 }}</ref>

<ref name="Baird 1986b">{{cite book |author=Baird RE. |title=Study of the stipitate Hydnums from the Southern Appalachian Mountains. Genera: ''Bankera'', ''Hydnellum'', ''Phellodon'', ''Sarcodon'' |series=Bibliotheca Mycologica |year=1986 |volume=104 |publisher=Lubrecht & Cramer |location=Berlin |isbn=978-3-443-59005-5}}</ref>

<ref name="Baird 1987">{{cite journal |vauthors=Baird RE, Khan SR |title=Spore ornamentation of North American ''Phellodon'' species |journal=Mycologia |year=1987 |volume=79 |issue=2 |pages=334–337 |doi=10.2307/3807670 |jstor=3807670 |url=http://www.cybertruffle.org.uk/cyberliber/59350/0079/002/0334.htm|url-access=subscription }}</ref>

<ref name="Baird 1988">{{cite journal |author=Baird RE. |title=Notes on the stipitate hydnums I. A new species of ''Phellodon'' |journal=Nova Hedwigia |year=1988 |volume=47 |issue=3–4 |pages=429–431}}</ref>

<ref name="Baird 2013">{{cite journal |vauthors=Baird RE, Wallace LE, Baker G |title=Stipitate hydnums of the southern United States 1: ''Phellodon mississippiensis'' sp. nov. |journal=Mycotaxon |year=2013 |volume=123 |pages=183–191 |doi=10.5248/123.183|doi-access=free }}</ref>

<ref name="Baird 2013b">{{cite journal |vauthors=Baird RE, Wallace LE, Baker G, Scruggs M |title=Stipitate hydnoid fungi of the temperate southeastern United States |journal=Fungal Diversity |year=2013 |volume=62 |issue=1 |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 |pages=99–194 (see p. 161) |url=https://www.biodiversitylibrary.org/page/31682597}}</ref>

<ref name="Binder 2005">{{cite journal |vauthors=Binder M, Hibbett DS, Larsson KH, Larsson E, Langer E, Langer G |title=The phylogenetic distribution of resupinate forms across the major clades of mushroom-forming fungi (Homobasidiomycetes) |journal=Systematics and Biodiversity |year=2005 |volume=3 |issue=2 |pages=113–157 |doi=10.1017/S1477200005001623 |s2cid=13102957 |url=http://www.clarku.edu/faculty/dhibbett/reprints%20pdfs/binder%20et%20al.%202005%20syst%20biodiv.pdf |access-date=9 September 2015 |archive-url=https://web.archive.org/web/20160304051657/http://www.clarku.edu/faculty/dhibbett/reprints%20pdfs/binder%20et%20al.%202005%20syst%20biodiv.pdf |archive-date=4 March 2016 |url-status=dead }}</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="Cannon 2007">{{cite book |vauthors=Cannon PF, Kirk PM |title=Fungal Families of the World |publisher=CAB International |location=Wallingford, UK |year=2007 |pages=33–34 |isbn=978-0-85199-827-5}}</ref>

<ref name="Cunningham 1958">{{cite journal |author=Cunningham GH. |title=Hydnaceae of New Zealand. Part I. The pileate genera ''Beenakia'', ''Dentinum'', ''Hericium'', ''Hydnum'', ''Phellodon'' and ''Steccerinum'' |journal=Transactions and Proceedings of the Royal Society of New Zealand |volume=85 |issue=4 |year=1957–58 |url=http://rsnz.natlib.govt.nz/volume/rsnz_85/rsnz_85_04_006410.html |pages=585–601}}</ref>

<ref name="Dai 2011">{{cite journal |author=Dai Y-C. |title=A revised checklist of corticioid and hydnoid fungi in China for 2010 |journal=Mycoscience |year=2011 |volume=52 |issue=1 |pages=69–79 |doi=10.1007/s10267-010-0068-1|s2cid=86049359 |doi-access=free }}</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=405–407 |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="Fang 2010">{{cite journal |vauthors=Fang ST, Zhang L, Li ZH, Li B, Liu JK |title=Cyathane diterpenoids and nitrogenous terphenyl derivative from the fruiting bodies of basidiomycete ''Phellodon niger'' |journal=Chemical and Pharmaceutical Bulletin |year=2010 |volume=58 |issue=9 |pages=1176–1179 |pmid=20823596 |doi=10.1248/cpb.58.1176 |doi-access=free }}</ref>

<ref name="Fries 1815">{{cite book |author=Fries EM. |title=Observationes mycologicae |year=1815 |publisher=Gerhard Bonnier |location=Copenhagen |volume=1 |page=134 |language=Latin |url=http://www.cybertruffle.org.uk/cyberliber/01532/0134.htm}}</ref>

<ref name="Harrison 1961">{{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 (see p. 15) |url=https://www.biodiversitylibrary.org/page/39011849}} {{open access}}</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=1205–1233 |doi=10.1139/b64-116}}</ref>

<ref name="Harrison 1972">{{cite journal |author=Harrison KA. |title=A new species of ''Phellodon'' possessing clamp connections |journal=Canadian Journal of Botany |year=1972 |volume=50 |issue=6 |pages=1219–1221 |doi=10.1139/b72-147}}</ref>

<ref name="Högberg 1999">{{cite journal |vauthors=Högberg P, Plamboeck AH, Taylor AF, Fransson PM |year=1999 |title=Natural C-13 abundance reveals trophic status of fungi and host-origin of carbon in mycorrhizal fungi in mixed forests |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=96 |issue=15 |pages=8534–8539 |doi=10.1073/pnas.96.15.8534|pmc=17551 |pmid=10411910|bibcode=1999PNAS...96.8534H |doi-access=free }} {{open access}}</ref>

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

<ref name="Khara 1978">{{cite journal |author=Khara HS. |title=Some stipitate hydnums from north western Himalayas |journal=Indian Journal of Mycology and Plant Pathology |year=1978 |volume=7 |issue=2 |pages=127–134}}</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=520 |isbn=978-0-85199-826-8}}</ref>

<ref name="Kirk 2015">{{cite web |author=Kirk PM. |title=Species Fungorum (version 26th August 2015). In: Species 2000 & ITIS Catalogue of Life |url=http://www.catalogueoflife.org/col/browse/tree/id/3824ba8ee911ca405f1b73ccf6041d29 |access-date=2015-09-08 |archive-date=8 August 2020 |archive-url=https://web.archive.org/web/20200808234015/http://www.catalogueoflife.org/col/browse/tree/id/3824ba8ee911ca405f1b73ccf6041d29 |url-status=dead }}</ref>

<ref name="Maas Geesteranus 1966">{{cite journal |author=Maas Geesteranus RA. |title=Notes on Hydnums, IV |journal=Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen Section C |year=1966 |volume=69 |pages=317–333}}</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 1974">{{cite journal |author=Maas Geesteranus RA. |title=Notes on hydnums. IX |journal=Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen Afdeling Natuurkunde, Series C |year=1974 |volume=77 |pages=215–222}}</ref>

<ref name="Marren 2000">{{cite report |author=Marren P.|author-link=Peter Marren|year=2000 |title=Stipitate hydnoid fungi in Britain. English Nature Research Report No. 420. |publisher=English Nature |location=Peterborough, UK |url=http://publications.naturalengland.org.uk/file/6292262992478208 |format=PDF}}</ref>

<ref name="McNabb 1971">{{cite journal |author=McNabb RFR. |title=Some new and revised taxa of New Zealand Basidiomycetes (Fungi) |journal=New Zealand Journal of Botany |year=1971 |volume=9 |issue=2 |pages=355–370 |doi=10.1080/0028825X.1971.10429148|doi-access=free }}</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=PA97 |year=1998 |publisher=Houghton Mifflin Harcourt |isbn=978-0-395-91090-0 |page=97}}</ref>

<ref name="Miller 1933">{{cite journal |author=Miller LW. |title=The genera of Hydnaceae |journal=Mycologia |year=1933 |volume=25 |issue=4 |pages=286–301 |url=http://www.cybertruffle.org.uk/cyberliber/59350/0025/004/0286.htm |doi=10.2307/3754097|jstor=3754097 |url-access=subscription }}</ref>

<ref name="Niemelä 2003">{{cite journal |vauthors=Niemelä T, Kinnunen J, Renvell P, Schigel D |title=''Phellodon secretus'' (Basidiomycota), a new hydnaceous fungus from northern pine woodlands |journal=Karstenia |year=2003 |volume=43 |issue=2 |pages=37–44 |doi=10.29203/ka.2003.391 |url=http://karstenia.fi/phellodon-secretus-basidiomycota-a-new-hydnaceous-fungus-from-northern-pine-woodlands/ |format=abstract|doi-access=free }}</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=761–777 |doi=10.1016/j.mycres.2007.05.003 |pmid=17681224}}</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 |page=45}}</ref>

<ref name="Pouzar 1956">{{cite journal |author=Pouzar Z. |title=Príspevek k poznání nasich kloboukatých losáku |journal=Ceská Mykologie |year=1956 |volume=10 |issue=2 |pages=65–76 |language=Czech}}</ref>

<ref name="Ragab 1953">{{cite journal |author=Ragab MA. |title=Taxonomic studies in the Hydnaceae with reference to their hyphal systems |journal=Mycologia |year=1955 |volume=45 |issue=6 |pages=941–946 |doi=10.1080/00275514.1953.12024326 |url=http://www.cybertruffle.org.uk/cyberliber/59350/0045/006/0941.htm| jstor=4547774|url-access=subscription }}</ref>

<ref name="Reekie 2008">{{cite journal |vauthors=Reekie TA, Austin KA, Banwell MG, Willis AC |title= The chemoenzymatic total synthesis of phellodonic acid, a biologically active and highly functionalized hirsutane derivative isolated from the Tasmanian fungus ''Phellodon melaleucus'' |journal=Australian Journal of Chemistry |year=2008 |volume=61 |issue=2 |pages=94–106 |doi=10.1071/CH07403|hdl=1885/50449 |s2cid= 96560493 |hdl-access=free }}</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 |page=474 |isbn=978-0-226-72117-0}}</ref>

<ref name="Smith 1975">{{cite book |author=Smith AH. |title=A Field Guide to Western Mushrooms |publisher=University of Michigan Press |location=Ann Arbor, Michigan |year=1975 |page=57 |isbn=978-0-472-85599-5 |url=https://books.google.com/books?id=q7eXGZQG7ksC&pg=PA57}}</ref>

<ref name="Stadler 1993">{{cite journal |vauthors=Stadler M, Anke T, Dasenbrock J, Steglich W |title=Phellodonic acid, a new biologically active hirsutane derivative from ''Phellodon melaleucus'' (Thelephoraceae, Basidiomycetes) |journal=Zeitschrift für Naturforschung C |year=1993 |volume=48 |issue=7–8 |pages=545–549 |doi=10.1515/znc-1993-7-803 |pmid=8216611 |s2cid=13176756 |url=http://zfn.mpdl.mpg.de/xtf/data/Reihe_C/48/ZNC-1993-48c-0545.pdf}}</ref>

<ref name="Stalpers 1993">{{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 (see pp. 18; 44–48) |url=http://www.cbs.knaw.nl/publications/Sim35/full%20text.htm |access-date=9 September 2015 |archive-date=21 February 2015 |archive-url=https://web.archive.org/web/20150221225508/http://www.cbs.knaw.nl/publications/Sim35/full%20text.htm |url-status=dead }}</ref>

<ref name="Taylor 2003">{{cite journal |vauthors=Taylor AF, Fransson PM, Högberg P, Högberg MN, Plamboeck AH |title=Species level patterns in C-13 and N-15 abundance of ectomycorrhizal and saprotrophic fungal sporocarps |journal=New Phytologist |year=2003 |volume=159 |issue=3 |pages=757–774 |doi=10.1046/j.1469-8137.2003.00838.x|pmid=33873595 |doi-access=free }} {{open access}}</ref>

<ref name="urlMycoBank: Phellodon">{{cite web |title=''Phellodon'' P. Karst. |url=http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=57133&Fields=All |publisher=MycoBank. International Mycological Association |access-date=2015-09-19}}</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 |pmid=18586344 |doi=10.1016/j.mimet.2008.04.010}}</ref>

<ref name="van der Linde 2009">{{cite journal |vauthors=van der Linde S, Alexander IJ, Anderson IC |title=Spatial distribution of sporocarps of stipitate hydnoid fungi and their belowground mycelium |journal=FEMS Microbiology Ecology |year=2009 |volume=69 |issue=3 |pages=344–352 |doi=10.1111/j.1574-6941.2009.00716.x |pmid=19558589|doi-access=free }} {{open access}}</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=633–640 |doi=10.1016/j.funeco.2012.04.002}}</ref>

<ref name="Zhishu 1993">{{cite book |vauthors=Zhishu B, Zheng G, Taihui L |title=The Macrofungus Flora of China's Guangdong Province |publisher=Columbia University Press |location=New York, New York |year=1993 |page=111 |isbn=9789622015562 |url=https://books.google.com/books?id=0cAered-vqYC&pg=PA111}}</ref>

}}

== External links == * {{Commons category-inline}}

Category:Phellodon Category:Thelephorales genera Category:Taxa named by Petter Adolf Karsten