Editing Truffle (section)

==Taxonomic details==

=== Species ===

==== Black ====
{{Main|Tuber melanosporum}}
[[File:Tuber brumale - Vue sur la tranche coupée.jpg|alt=|thumb|Black Périgord truffle, cross-section]]

The black truffle or black Périgord truffle (''Tuber melanosporum''), the second-most commercially valuable species, is named after the [[Périgord]] region in France.<ref name="Trappe-2009">{{Cite journal|last=Trappe|first=Jim|date=2009|title=Taming the truffle—the history, lore, and science of the ultimate mushroom |journal=Gastronomica|volume=9|issue=1|pages=116–117|doi=10.1525/gfc.2009.9.1.116|issn=1529-3262}}</ref> Black truffles associate with [[oak]]s, [[hazel]]nut, cherry, and other deciduous trees and are harvested in late autumn and winter.<ref name="Trappe-2009" /><ref name="Carluccio032">{{cite book|title=The Complete Mushroom Book|author=Carluccio|first=Antonio|publisher=Quadrille|year=2003|isbn=978-1-84400-040-1|author-link=Antonio Carluccio}}</ref> The genome sequence of the black truffle was published in March 2010.<ref name="truffle genome2">{{cite journal | last1 = Martin | first1 = F. | last2 = Kohler | first2 = A. | last3 = Murat | first3 = C. | last4 = Balestrini | first4 = R. | last5 = Coutinho | first5 = P.M. | last6 = Jaillon | first6 = O. | last7 = Montanini | first7 = B. | last8 = Morin | first8 = E. | last9 = Noel | first9 = B. | last10 = Percudani | first10 = R. | last11 = Porcel | first11 = B. | year = 2010 | title = Périgord black truffle genome uncovers evolutionary origins and mechanisms of symbiosis | journal = Nature | volume = 464 | issue = 7291| pages = 1033–1038 | doi=10.1038/nature08867 | pmid=20348908| bibcode = 2010Natur.464.1033M | doi-access = free | hdl = 2318/100278 | hdl-access = free }}</ref>

==== Summer or burgundy ====
{{Main|Tuber aestivum}}
[[File:Black.summer.truffle.arp.jpg|thumb|Summer truffles in a shop in Rome]]

The black summer truffle (''Tuber aestivum'') is found across Europe and is prized for its culinary value.<ref name="Paolocci-2004">{{Cite journal|last1=Paolocci|first1=Francesco|last2=Rubini|first2=Andrea|last3=Riccioni|first3=Claudia|last4=Topini|first4=Fabiana|last5=Arcioni|first5=Sergio|date=2004|title=Tuber aestivum and Tuber uncinatum: two morphotypes or two species?|journal=FEMS Microbiology Letters|volume=235|issue=1|pages=109–115|doi=10.1111/j.1574-6968.2004.tb09574.x|issn=0378-1097|pmid=15158269|doi-access=free}}</ref> Burgundy truffles (designated ''[[Tuber uncinatum]]'', but the same species) are harvested in autumn until December and have aromatic flesh of a darker colour. These are associated with various trees and shrubs.<ref name="Paolocci-2004" />

==== White ====
{{main|Tuber magnatum}}
[[File:Truffle washed and cut.jpg|alt=|thumb|A white truffle washed and with a corner cut to show the interior]] 
Tuber magnatum, the high-value ''white truffle'' ({{langx|it|tartufo bianco}}) is found mainly in the [[Langhe]] and [[Montferrat]] areas<ref>{{cite web|url=http://www.lifeinitaly.com/food/truffle.asp|title=White truffles from Alba|last=Demetri|first=Justin|date=2012|website=Life in Italy|publisher=Lifeinitaly.com|access-date=2012-06-16|archive-date=2009-10-16|archive-url=https://web.archive.org/web/20091016180401/http://www.lifeinitaly.com/food/truffle.asp|url-status=live}}</ref> of the [[Piedmont]] region in northern [[Italy]], and most famously, in the countryside around the cities of [[Alba, Piedmont|Alba]] and [[Asti]].<ref>{{Cite journal|last1=Bencivenga  |first1=M. |last2=Di Massimo |first2=G. |last3=Donnini |first3=D. |last4=Baciarelli Falini |first4=L. |date=2009|title=The cultivation of truffles in Italy |journal=Acta Botanica Yunnanica |volume=16 |issue=Suppl 16 |pages=100–102}}</ref> A large percentage of Italy's white truffles also come from [[Molise]].

==== Whitish ====
{{Main|Tuber borchii}}
The "[[whitish truffle]]" (''Tuber borchii'') is a similar species native to [[Tuscany]], [[Abruzzo]], [[Romagna]], [[Umbria]], the [[Marche]], and [[Molise]]. It is reportedly not as aromatic as those from Piedmont, although those from [[Città di Castello]] are said to come quite close.<ref name="Carluccio032" />

==== Other ''Tuber'' ====
A less common truffle is "garlic truffle" (''[[Tuber macrosporum]]'').

In the U.S. [[Pacific Northwest]], several species of truffle are harvested both recreationally and commercially, most notably, the ''[[Leucangium carthusianum]]'', Oregon black truffle; ''[[Tuber gibbosum]]'', Oregon spring white truffle; and ''[[Tuber oregonense]]'', Oregon winter white truffle. ''[[Kalapuya brunnea]]'', the Oregon brown truffle, has also been commercially harvested and is of culinary note. The Oregon white truffle is increasingly harvested due to its high quality and also exported to other countries. Oregon celebrates its traditional truffle harvesting with a 'truffle festival', combined with culinary shows and wine tastings.<ref>[https://knowyourforest.org/sites/default/files/documents/Oregon_Native_Truffles.pdf "Oregon Native Truffles: A primer for commercial production on small forestlands in the Pacific Northwest"]. ''Know Your Forest''. Accessed 8 October 2023.</ref>

The [[pecan truffle]] (''Tuber lyonii'')<ref>{{cite journal|first=Fred K. |last=Butters|year=1903|title=A Minnesota Species of Tuber|journal=Botanical Gazette|volume=35|issue=6|pages=427–431|doi=10.1086/328364|jstor=2556357|s2cid=84500806}}</ref> syn. ''texense''<ref>{{cite journal|first1=J. M. |last1=Trappe |first2=A. M. |last2=Jumpponen |first3=E. |last3=Cázares|name-list-style=amp|year=1996|title=NATS truffle and truffle-like fungi 5: ''Tuber lyonii'' (=''T. texense''), with a key to the spiny-spored ''Tuber'' species groups|url=http://www.cybertruffle.org.uk/cyberliber/59575/0060/0365.htm|journal=Mycotaxon|volume=60|pages=365–372|access-date=2010-06-04|archive-date=2015-09-23|archive-url=https://web.archive.org/web/20150923232916/http://www.cybertruffle.org.uk/cyberliber/59575/0060/0365.htm|url-status=live}}</ref> is found in the [[Southern United States]], usually associated with [[pecan]] trees. Chefs who have experimented with them agree "they are very good and have potential as a food commodity".<ref>{{cite web|url=http://www.caes.uga.edu/commodities/fruits/pecantruffles/index.html|title=Pecan Truffles|author=Tim Brenneman|date=2010|archive-url=https://web.archive.org/web/20100609195108/http://www.caes.uga.edu/commodities/fruits/pecantruffles/index.html|archive-date=2010-06-09|url-status=dead|access-date=2010-06-03}}</ref> Although pecan farmers used to find them along with pecans and discard them, considering them a nuisance, they sell for about $160 a pound and have been used in some gourmet restaurants.<ref name="pecanmag2">{{cite magazine|last1=Smith |first1=M. E. |display-authors=etal |date=2012|title=Pecan Truffles ( ''Tuber lyonii'' ): What We Know and What We Need to Know|magazine=Georgia Pecan Magazine|issue=Spring 2012|pages=52–58}}</ref>

==== Beyond ''Tuber'' {{anchor|Similar organisms}} ====
{{missing information|section|taste and flavor|date=February 2022}}
The term "truffle" has been applied to several other genera of similar underground fungi. The genera ''[[Terfezia]]'' and ''[[Tirmania]]'' of the family [[Terfeziaceae]] are known as the "desert truffles" of Africa and the Middle East. ''[[Pisolithus tinctorius]]'', which was historically eaten in parts of [[Germany]], is sometimes called "Bohemian truffle".<ref name="Ramsbottom53">{{cite book |last=Ramsbottom |first=J. |title=Mushrooms & Toadstools |publisher=Collins |year=1953}}</ref>

{{anchor|Geopora}}''[[Geopora]]'' spp. are important ectomycorrhizal partners of trees in woodlands and forests throughout the world.<ref name="Læssøe 2007" /> ''Pinus edulis'', a widespread pine species of the Southwest US, is dependent on ''Geopora'' for nutrient and water acquisition in arid environments.<ref name="Flores-Rentería-2014">{{Cite journal|last1=Flores-Rentería|first1=Lluvia|last2=Lau|first2=Matthew K.|last3=Lamit|first3=Louis J.|last4=Gehring|first4=Catherine A.|date=2014|title=An elusive ectomycorrhizal fungus reveals itself: a new species of Geopora (Pyronemataceae) associated with ''Pinus edulis'' |journal=Mycologia|volume=106|issue=3 |pages=553–563 |doi=10.3852/13-263|pmid=24871594|s2cid=207630013|issn=0027-5514}}</ref> Like other truffle fungi, ''Geopora'' produces subterranean [[Sporocarp (fungi)|sporocarps]] as a means of sexual reproduction.<ref name="Flores-Rentería-2014" /> ''[[Geopora cooperi]]'', also known as pine truffle or fuzzy truffle, is an edible species of this genus.<ref name="Læssøe 2007" />

[[File:Rhizopogon rubescens.jpg|thumb|''Rhizopogon'' truffle]]
''[[Rhizopogon]]'' spp. are ectomycorrhizal members of the Basidiomycota and the order [[Boletales]], a group of fungi that typically form mushrooms.<ref name="Molina-1994">{{Cite journal|last1=Molina |first1=Randy|last2=Trappe |first2=James M.|date=April 1994 |title=Biology of the ectomycorrhizal genus, Rhizopogon. I. Host associations, host-specificity and pure culture syntheses |journal=New Phytologist |volume=126 |issue=4 |pages=653–675|doi=10.1111/j.1469-8137.1994.tb02961.x |issn=0028-646X|doi-access=free }}</ref> Like their ascomycete counterparts, these fungi can create truffle-like fruiting bodies.<ref name="Molina-1994" /> ''Rhizopogon'' spp. are ecologically important in coniferous forests where they associate with various pines, firs, and [[Douglas fir]].<ref name="Griffiths">{{Cite journal |last1=Griffiths |first1=Robert P. |last2=Caldwell |first2=Bruce A. |last3=Cromack |first3=Kermit Jr. |last4=Morita |first4=Richard Y. |date=February 1990 |title=Douglas-fir forest soils colonized by ectomycorrhizal mats. I. Seasonal variation in nitrogen chemistry and nitrogen cycle transformation rates  |journal=Canadian Journal of Forest Research |volume=20 |issue=2 |pages=211–218 |doi=10.1139/x90-030 |bibcode=1990CaJFR..20..211G |issn=0045-5067}}</ref> In addition to their ecological importance, these fungi hold economic value, as well. ''Rhizopogon'' spp. are commonly used to inoculate coniferous seedlings in nurseries and during reforestation.<ref name="Molina-1994" />

''[[Hysterangium]]'' spp. are ectomycorrhizal members of the Basidiomycota and the order [[Hysterangiales]] that form sporocarps similar to true truffles.<ref name="Kluber-2010">{{Cite journal|last1=Kluber|first1=Laurel A.|last2=Tinnesand|first2=Kathryn M.|last3=Caldwell|first3=Bruce A.|last4=Dunham|first4=Susie M.|last5=Yarwood|first5=Rockie R.|last6=Bottomley|first6=Peter J.|last7=Myrold|first7=David D.|date=2010|title=Ectomycorrhizal mats alter forest soil biogeochemistry |journal=Soil Biology and Biochemistry|volume=42|issue=9|pages=1607–1613|doi=10.1016/j.soilbio.2010.06.001 |bibcode=2010SBiBi..42.1607K |s2cid=85720492 |issn=0038-0717}}</ref> These fungi form mycelial mats of vegetative hyphae that may cover 25–40% of the forest floor in Douglas fir forests, thereby contributing to a significant portion of the biomass present in soils.<ref name="Kluber-2010" /> Like other ectomycorrhizal fungi, ''Hysterangium'' spp. play a role in nutrient exchange in the nitrogen cycle by accessing nitrogen unavailable to host plants and acting as nitrogen sinks in forests.<ref name="Griffiths" />

''[[Glomus (fungus)|Glomus]]'' spp. are [[arbuscular mycorrhiza]]e of the phylum Glomeromycota within the order [[Glomerales]].<ref name="Trappe 2009">{{Cite work |last1=Trappe |first1=James M. |title=Diversity, ecology, and conservation of truffle fungi in forests of the Pacific Northwest |last2=Molina |first2=Randy |last3=Luoma |first3=Daniel L. |last4=Cázares |first4=Efren |last5=Pilz |first5=David |last6=Smith |first6=Jane E. |last7=Castellano |first7=Michael A. |last8=Miller |first8=Steven L. |last9=Trappe |first9=Matthew J. |date=2009 |work=General Technical Report PNW-GTR-772 |publisher=USDA Forest Service |location=Portland, Oregon |doi=10.2737/pnw-gtr-772 |hdl=2027/umn.31951d02938269i}}</ref> Members of this genus have low host specificity, associating with a variety of plants including hardwoods, forbs, shrubs, and grasses.<ref name="Trappe 2009" /> These fungi commonly occur throughout the Northern Hemisphere.<ref name="Trappe 2009" />

Members of the genus ''[[Elaphomyces]]'' are commonly mistaken for truffles.

=== Phylogeny ===
[[File:Truffle Evolution.jpg|thumb|upright=1.2|Evolution of subterranean fruiting bodies from above-ground mushrooms.]]
Phylogenetic analysis has demonstrated the [[convergent evolution]] of the ectomycorrhizal trophic mode in diverse fungi. The subphylum [[Pezizomycotina]], containing the order Pezizales, is approximately 400 million years old.<ref name="Kohler">{{Cite journal |last=Kohler |first=Annegre |year=2015 |title=Convergent losses of decay mechanisms and rapid turnover of symbiosis genes in mycorrhizal mutualists |journal=Nature Genetics |location=Nature Genetics |publisher=Lawrence Berkeley National Laboratory, United States Department of Energy |volume=47 |issue=4 |pages=410–5 |doi=10.1038/ng.3223 |oclc=946824824 |pmid=25706625 |s2cid=20914242 |doi-access=free|hdl=1942/18722 |hdl-access=free }}</ref> Within the order Pezizales, subterranean fungi evolved independently at least fifteen times.<ref name="Kohler" /> Contained within Pezizales are the families [[Tuberaceae]], [[Pezizaceae]], [[Pyronemataceae|Pyronematacae]], and [[Morchellaceae]]. All of these families contain lineages of subterranean or truffle fungi.<ref name="Læssøe 2007" />

The oldest ectomycorrhizal fossils are from the [[Eocene]] about 50 million years ago. The specimens are preserved [[permineralized]] in-situ in the [[Eocene Okanagan Highlands]] [[Princeton chert]] site. This indicates that the soft bodies of ectomycorrhizal fungi do not easily fossilize.<ref>{{cite journal |last1=LePage |first1=B. A. |last2=Currah |first2=R. S. |last3=Stockey |first3=R. A. |last4=Rothwell |first4=G. W. |year=1997 |title=Fossil ectomycorrhizae from the middle Eocene |journal=American Journal of Botany |volume=84 |issue=3 |pages=410–412 |doi=10.2307/2446014 |jstor=2446014 |pmid=21708594 |s2cid=29913925}}</ref> Molecular clockwork has suggested the evolution of ectomycorrhizal fungi occurred approximately 130 million years ago.<ref>{{Cite journal |last1=Berbee |first1=Mary L. |last2=Taylor |first2=John W. |date=August 1993 |title=Dating the evolutionary radiations of the true fungi |journal=Canadian Journal of Botany |volume=71 |issue=8 |pages=1114–1127 |doi=10.1139/b93-131 |bibcode=1993CaJB...71.1114B |issn=0008-4026}}</ref>

The evolution of subterranean fruiting bodies has occurred numerous times within the [[Ascomycota]], [[Basidiomycota]], and [[Glomeromycota]].<ref name="Læssøe 2007" /> For example, the genera ''[[Rhizopogon]]'' and ''[[Hysterangium]]'' of Basidiomycota both form subterranean fruiting bodies and play similar ecological roles as truffle forming ascomycetes. The ancestors of the Ascomycota genera ''Geopora'', ''Tuber'', and ''Leucangium'' originated in Laurasia during the Paleozoic era.<ref name="Trappe 2009" />

Phylogenetic evidence suggests that most subterranean fruiting bodies evolved from above-ground mushrooms. Over time mushroom stipes and caps were reduced, and caps began to enclose reproductive tissue. The dispersal of sexual spores then shifted from wind and rain to utilising animals.<ref name="Trappe 2009" />

The [[phylogeny]] and [[biogeography]] of the genus ''Tuber'' was investigated in 2008<ref>{{cite journal |last1=Jeandroz |first1=S. |last2=Murat |first2=C. |last3=Wang |first3=Y. |last4=Bonfante |first4=P. |last5=Le Tacon |first5=F. |year=2008 |title=Molecular phylogeny and historical biogeography of the genus Tuber, the true truffles |journal=Journal of Biogeography |volume=35 |issue=5 |pages=815–829 |doi=10.1111/j.1365-2699.2007.01851.x |bibcode=2008JBiog..35..815J |s2cid=84381208}}</ref> using [[Internal transcribed spacer|internal transcribed spacers (ITS)]] of [[nuclear DNA]] and revealed five major [[clade]]s (Aestivum, Excavatum, Rufum, Melanosporum and Puberulum); this was later improved and expanded in 2010 to nine major clades using 28S [[Large subunit|large subunits (LSU)]] rRNA of [[mitochondrial DNA]]{{citation needed|date=October 2023}}. The Magnatum and Macrosporum clades were distinguished as distinct from the Aestivum clade. The Gibbosum clade was resolved as distinct from all other clades, and the Spinoreticulatum clade was separated from the Rufum clade.<ref>{{cite journal |vauthors=Bonito GM, Gryganskyi AP, Trappe JM, Vilgalys R |year=2010 |title=A global meta-analysis of Tuber ITS rDNA sequences: species diversity, host associations and long-distance dispersal |journal=PLOS ONE |volume=8 |issue=1 |pages=e52765 |bibcode=2013PLoSO...852765B |doi=10.1371/journal.pone.0052765 |pmc=3534693 |pmid=23300990 |doi-access=free}}</ref>

The truffle habit has evolved independently among several basidiomycete genera.<ref name="Bruns-1989">{{Cite journal |last1=Bruns |first1=Thomas D. |last2=Fogel |first2=Robert |last3=White |first3=Thomas J. |last4=Palmer |first4=Jeffrey D. |date=1989 |title=Accelerated evolution of a false-truffle from a mushroom ancestor |url=https://deepblue.lib.umich.edu/bitstream/2027.42/62545/1/339140a0.pdf |url-status=live |journal=Nature |volume=339 |issue=6220 |pages=140–142 |bibcode=1989Natur.339..140B |doi=10.1038/339140a0 |issn=0028-0836 |pmid=2716834 |s2cid=4312286 |archive-url=https://web.archive.org/web/20210827065108/https://deepblue.lib.umich.edu/bitstream/handle/2027.42/62545/339140a0.pdf;jsessionid=6C2FB0F2AE5763E76E2D55E2708812C9?sequence=1 |archive-date=2021-08-27 |access-date=2019-09-01 |hdl-access=free |hdl=2027.42/62545}}</ref><ref>{{Cite journal |last=Hibbett |first=David S. |date=2007 |title=After the gold rush, or before the flood? Evolutionary morphology of mushroom-forming fungi (Agaricomycetes) in the early 21st century |journal=Mycological Research |volume=111 |issue=9 |pages=1001–1018 |doi=10.1016/j.mycres.2007.01.012 |issn=0953-7562 |pmid=17964768}}</ref><ref>{{Cite journal |last=Albee-Scott |first=Steven |date=2007 |title=The phylogenetic placement of the Leucogastrales, including Mycolevis siccigleba (Cribbeaceae), in the Albatrellaceae, using morphological and molecular data |journal=Mycological Research |volume=111 |issue=6 |pages=653–662 |doi=10.1016/j.mycres.2007.03.020 |issn=0953-7562 |pmid=17604150}}</ref> Phylogenetic analysis has revealed that basidiomycete subterranean fruiting bodies, like their ascomycete counterparts, evolved from above ground mushrooms. For example, ''Rhizopogon'' species likely arose from an ancestor shared with ''[[Suillus]]'', a mushroom-forming genus.<ref name="Bruns-1989" /> Studies have suggested that selection for subterranean fruiting bodies among ascomycetes and basidiomycetes occurred in water-limited environments.<ref name="Trappe 2009" /><ref name="Bruns-1989" />

=== Etymology ===
Most sources agree that the term "truffle" is derived from the [[Latin]] term {{lang|la|tūber}} by way of the Vulgar Latin {{lang|la|tufera}}, meaning "swelling" or "lump".<ref>{{cite web |date=3 July 2008 |title=Truffle – Podictionary Word of the Day, July 3rd 2008 |url=https://blog.oup.com/2008/07/truffle/ |access-date=12 March 2023 |website=OUP Blog |publisher=Oxford University Press}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.merriam-webster.com/dictionary/truffle |access-date=12 March 2023 |dictionary=Merriam-Webster}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.dictionary.com/browse/truffle |access-date=12 March 2023 |dictionary=Dictionary.com}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.etymonline.com/word/truffle |access-date=12 March 2023 |dictionary=Online Etymology Dictionary}}</ref> This then entered other languages through [[Old French]] dialects.