{{Short description|Species of fungus}} {{Speciesbox | image = Captura de Pantalla 2023-11-03 a la(s) 13.06.28.png | image_caption = | taxon = Ramaria subbotrytis | authority = (Coker) Corner (1950) | synonyms_ref = <ref name="urlMycoBank: Ramaria subbotrytis"/> | synonyms = *''Clavaria subbotrytis'' <small>Coker (1923)</small> }}

'''''Ramaria subbotrytis''''' is a species of coral fungus in the family Gomphaceae. It was previously classified in the family Ramariaceae, and before that in the genus Clavaria, family Clavariaceae. It was originally described as ''Clavaria subbotrytis'' by William Chambers Coker in 1923 from collections made in North Carolina.<ref name="Coker 1923"/> E.J.H. Corner transferred it to the genus ''Ramaria'' in 1950.<ref name="Corner 1950"/>

''Ramaria subbotrytis'' accumulates arsenic and besides arsenic species known to be found in macrofungi, such as arsenobetaine, contains a very unusual organoarsenic compound homoarsenocholine.<ref name=Brae>{{cite journal |vauthors=Braeuer S, Borovička J, Glasnov T, Guedes de la Cruz G, Jensen KB, Goessler W |year=2018 |title=Homoarsenocholine – A novel arsenic compound detected for the first time in nature |journal=Talanta |volume=188 |pages=107–110 |issn=0039-9140 |doi=10.1016/j.talanta.2018.05.065 |pmid=30029352 |pmc=6118324}}</ref> It grows in association with trees, especially conifers.<ref name="pubmed.ncbi.nlm.nih.gov">{{cite journal |last1=Nouhra |first1=Eduardo |title=Morphological and molecular characterization of selected Ramaria mycorrhizae |journal= Mycorrhiza|date=2005 |volume=15 |issue=1 |pages=55–59 |doi=10.1007/s00572-004-0294-5 |pmid=14745631 |hdl=11336/31987 |s2cid=2024353 |hdl-access=free }}</ref>

The species is listed as edible by some sources.<ref name="Miller 2006">{{cite book|last1=Miller Jr.|first1=Orson K.|title=North American Mushrooms: A Field Guide to Edible and Inedible Fungi|last2=Miller|first2=Hope H.|publisher=FalconGuide|year=2006|isbn=978-0-7627-3109-1|location=Guilford, CN|pages=349|author-link=Orson K. Miller Jr.}}</ref><ref>{{cite journal |last1=Hall |first1=Ian |title=A list of putative edible or medicinal ectomycorrhizal mushrooms |date=May 2016 |doi=10.13140/RG.2.1.2978.9048 |url=https://www.researchgate.net/publication/302319041}}</ref>

thumb|Coker's illustration of various Clavarias, with R. subbotrytis as figure 5

== Taxonomy and Phylogeny ==

''Ramaria subbotrytis'', first described as ''Clavaria subbotrytis'' by Coker in 1923, was moved to the genus ''Ramaria'' by Corner in 1950.<ref name="Coker 1923" /><ref name="Corner 1950" /> A synonym of ''R. subbotrytis'' is therefore ''C. subbotrytis''. The species has also undergone multiple other taxonomic changes, such as from the family Clavariaceae, to Ramariaceae, and most recently to the family Gomphaceae.

A recent phylogenetic study placed the genus ''Ramaria'' within the order Gomphales and the family Gomphaceae through ribosomal DNA analysis. References to unpublished analyses in the study theorize that ''Ramaria'' is monophyletic. The study places ''Ramaria'' as closely related to other hymenomycetes, including the genera ''Gomphus'' and ''Clavariadelphus'', which all form a sister clade to another monophyletic group which includes gasteromycetes in the genera ''Sphaerobolus'' and ''Geastrum'', and the order Phallales (represented in the study by genus ''Pseudocolus'').<ref>{{cite journal |last1=Hibbet |first1=David |title=Evolution of gilled mushrooms and puffballs inferred from ribosomal DNA sequences |journal=Proceedings of the National Academy of Sciences of the United States of America |date=October 1997 |volume=94 |issue=22 |pages=12002–12006 |doi=10.1073/pnas.94.22.12002 |pmid=9342352 |pmc=23683 |bibcode=1997PNAS...9412002H |doi-access=free }}</ref>

thumb|Coker's image of Clavaria subbotrytis holotype

== Morphology ==

''R. subbotrytis'' is coral pink when immature, and fades to a creamy color except at the tips of the branches when it reaches maturity. It ranges from 7.5 to 15 centimeters tall and 5.5 to 9 centimeters wide.

The spores are produced on basidia lining the outer part of the branch tips. The basidia measure 6.5 to 9.5 micrometers, and are irregular and 4-spored. The hymenium is 55 to 65 micrometers thick with hyphae varying in size.

The spores are “nearly smooth, elliptic, cinnamon-ocraceous,” with hints of a rose color in the spore print. Coker measured the spores of one specimen between 3 to 3.7 by 7.4 to 9 micrometers, and the basidia of another specimen at 6.5 to 9.5 micrometers. He did not observe any clamp connections.<ref name="Coker 1923" /> Corner, however, described ''R. subbotrytis'' as having clamp connections in the basidia, but not in the hyphae.<ref>{{cite journal |last1=Corner |first1=E. J. H. |title=Species of Ramaria (clavariaceae) without clamps |journal=Botany School, University of Cambridge, UK |date=15 June 1965 |volume=49 |pages=101–113 |doi=10.1016/S0007-1536(66)80040-2 |url=https://www.sciencedirect.com/science/article/abs/pii/S0007153666800402|url-access=subscription }}</ref>

The texture of ''R. subbotrytis'' is tender but not brittle. It has a short, clean, smooth, and glabrous stipe with no aborted branches. It tapers to a point on the base. Coker noted the flesh to sliver or peel when damaged.<ref name="Coker 1923" />

{{mycomorphbox | name = ''Ramaria subbotrytis'' | hymeniumType = smooth | capShape = no | stipeCharacter = bare | sporePrintColor = brown | ecologicalType = mycorrhizal | howEdible = edible | howEdible2 = caution }}

== Chemical composition ==

Coker noted ''R. subbotrytis'' to have a “krauty” taste and odor, though only slightly.<ref name="Coker 1923" />

In a 2018 study, ''R. subbotrytis'' was determined through HPLC and mass spectrometry to contain several rare arsenic species, including trimethylarsoniopropanate and dimethylarsinoylacetate, which had only been found previously in marine samples. The study also noted the presence of homoarsenocholine, a compound which had never before been observed in nature.<ref name="Brae" />

== Ecological relationships ==

As a member of ''Ramaria'', ''R. subbotrytis'' forms ectomycorrhizal relationships with trees. Coker found his holotype at the base of a hickory tree.<ref name="Coker 1923" /> ''Ramaria'' are notable symbionts of conifers, especially in the Pacific Northwest, where their mycorrhiza have been studied in-depth. A study in this region on the identification of different ''Ramaria'' mycorrhiza showed that the mycorrhiza of different species in the genus are very morphologically similar and difficult to distinguish from one another. Furthermore, it characterized the mycorrhiza of ''Ramaria'' as forming mycelial mats below the soil which are made up of hyphae, mycorrhizal roots, and substrate.<ref name="pubmed.ncbi.nlm.nih.gov"/>

''R. subbotrytis'' has been described to be distributed throughout North America and Europe. The social network iNaturalist displays observations (both research grade and unverified) in both regions.<ref>{{cite web |title=Observations of Ramaria subbotrytis |url=https://www.inaturalist.org/observations?taxon_id=352197 |website=iNaturalist |access-date=3 November 2023}}</ref> It has been described in scientific literature as appearing in Slovakia, the Czech Republic,<ref name="Brae" /> the Urals of Russia,<ref>{{cite journal |last1=Shiryaev |first1=Anton |title=Clavarioid fungi of the Urals. II. The nemoral zone |journal=Karstenia |date=2007 |volume=47 |pages=5–16 |doi=10.29203/ka.2007.417 |url=http://www.funga.fi/Karstenia/Karstenia_47-1_2007-2.pdf |issn=0453-3402}}</ref> the Jilotzingo region of Mexico,<ref name="colposdigital.colpos.mx">{{cite journal |last1=Bautista |first1=Bautista |title=Conocimiento micológico tradicional y mesofauna asociada a los hongos comestibles silvestres de Santa Ana Jilotzingo, Estado de México. |journal=Colegio de Posgrados |date=February 2023 |url=http://colposdigital.colpos.mx:8080/xmlui/handle/10521/5093 |access-date=3 November 2023}}</ref> throughout the Iberian Peninsula,<ref>{{cite journal |last1=Parra Becerra |first1=Manuel |title=http://sociedadgaditanahistorianatural.com/wp-content/uploads/2012/05/01_Becerra_web.pdf |journal=Sociedad gitana historia natural |date=18 August 2011 |issue=3 May 2012 |pages=1–2 |url=http://sociedadgaditanahistorianatural.com/wp-content/uploads/2012/05/01_Becerra_web.pdf}}</ref> and throughout the United States and Canada, making the species widely geographically distributed. It was described in 2011 in Andalucía, Spain for the first time.<ref>{{cite journal |last1=Parra Becerra |first1=Manuel |title=http://sociedadgaditanahistorianatural.com/wp-content/uploads/2012/05/01_Becerra_web.pdf |journal=Sociedad gitana historia natural |date=18 August 2011 |issue=3 May 2012 |pages=1–2 |url=http://sociedadgaditanahistorianatural.com/wp-content/uploads/2012/05/01_Becerra_web.pdf}}</ref>

A study conducted about the mesofauna associated with different fungi in the province of Santa Ana Jilotzingo, Mexico found that collembolans and oribatid mites are the most prominent inhabitants of ''R. subbotrytis'' sporomes. Analysis of the mites’ and collembolans’ gut contents suggested a symbiotic relationship between the fungus and the arthropods, possibly involving spore dispersal due to the presence of spores in their digestive tracts.<ref name="colposdigital.colpos.mx"/>

== Edibility and traditional use == Many ''Ramaria'' species are used in traditional medicinal practices, and are traded commercially for these purposes or as food sources.<ref>{{cite book |last1=Boa |first1=Eric |title=Edible Fungi: A Global Overview of Their Use and Importance to People |date=2004 |publisher=Food and Acgriculture Organization of the United Nations |location=Rome |page=115 |url=https://www.researchgate.net/publication/284221866 |access-date=3 November 2023}}</ref> Some sources consider ''Ramaria subbotrytis'' to be edible,<ref name="Miller 2006" /> though others list it as “unknown” and cite its known arsenic contents.<ref name="Brae" />

== Similar species == Coker lists several morphologically similar species to ''R. subbotrytis'', including ''R. botrytis'', ''R. conjunctipes'' var. ''odora'', and ''R. formosa''. It is distinguished from ''R. botrytis'' because ''R. subbotrytis'' has much darker spores and lacks red branch tips in its immature stages. ''R. conjunctipes'' var. ''odora'' is distinguishable by its more brittle, solid flesh, different odor, deeper colored flesh with tips that are the same color as the rest of the fruiting body, and a wider and less compound base. It differs from ''R. formosa'' which is much more brittle and has yellow tips in immature stages, as well as a deeper pink color and narrower and smoother cinnamon colored spores than ''R. subbotrytis''.

Coker also described ''R. subbotrytis'' var. ''intermedia'', which is an intermediate species between ''R. botrytis'' and ''R. subbotrytis''. It has a similar body and base morphology to ''R. botrytis'', but the same spore color and shape as ''R. subbotrytis''. For that reason, Coker chose to classify it as a variant of ''R. subbotrytis''.<ref name="Coker 1923" />

==References== {{Reflist|refs=

<ref name="Coker 1923">{{cite book |author=Coker WC. |title=The Clavarias of the United States and Canada |publisher=The University of North Carolina Press |year=1923 |location=Chapel Hill, North Carolina |page=116 |url=https://www.biodiversitylibrary.org/page/4597619}}</ref>

<ref name="Corner 1950">{{cite book |author=Corner EJH. |title=A Monograph of ''Clavaria'' and Allied Genera |year=1950 |location=Cambridge, UK |publisher=Cambridge University Press |page=625}}</ref>

<ref name="urlMycoBank: Ramaria subbotrytis">{{cite web |title=''Ramaria subbotrytis'' (Coker) Corner :625, 1950 |url=http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=62565&Fields=All |publisher=MycoBank. International Mycological Association |accessdate=2013-02-11}}</ref>

}}

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Category:Gomphaceae Category:Fungi described in 1923 Category:Edible fungi Category:Fungi of North America Category:Fungus species