Lactarius

{{Short description|Genus of fungi}} {{For|the genus of fish|Lactarius (fish)}} {{Automatic taxobox | image = Lactarius torminosus (3).JPG | image_alt = "Lactarius torminosus" | image_caption = ''Lactarius torminosus'' | taxon = Lactarius | authority = Pers. (1797) | type_species = ''Lactarius torminosus'' | type_species_authority = (Schaeff.) Gray (1821) | synonyms = {{plainlist| *''Lactaria'' Pers. (1797) *''Agaricus'' sect. ''Lactifluus'' Pers. (1801) *''Agaricus'' subdiv. ''Galorrheus'' Fr. (1818) *''Lactariella'' J.Schröt. (1898) *''Lactariopsis'' Henn. (1901) *''Gloeocybe'' Earle (1909) *''Hypophyllum'' Earle (1909) }} | synonyms_ref = <ref name=mycobank-lactarius/> | subdivision_ranks = Species | subdivision = | diversity = c. 583 species | diversity_ref = <ref name=Lee2019/> | diversity_link = List of Lactarius species }}

'''''Lactarius''''' is a genus of mushroom-producing, ectomycorrhizal fungi, containing several edible species. The species of the genus, commonly known as '''milk-caps''', are characterized by the milky fluid ("latex") they exude when cut or damaged. Like the closely related genus ''Russula'', their flesh has a distinctive brittle consistency. It is a large genus with over 500 known species,<ref name=Lee2019/> mainly distributed in the Northern hemisphere. Recently, the genus ''Lactifluus'' has been separated from ''Lactarius'' based on molecular phylogenetic evidence.

==Taxonomy== thumb|Small droplets of "latex" exuding from a damaged species of Lactarius.

The genus ''Lactarius'' was described by Christian Hendrik Persoon in 1797<ref name=Persoon1797/> with ''L. piperatus'' as the original type species. In 2011, ''L. torminosus'' was accepted as the new type of the genus after the splitting-off of ''Lactifluus'' as separate genus.<ref name=Buyck2010/><ref name="Barrie2011"/><ref name="Norvell2011"/>

The name "''Lactarius''" is derived from the Latin ''lac'', "milk".

===Placement within Russulaceae=== {{cladogram|align=right|title= |clade= {{clade |style=font-size:90%;line-height:90% |label1= |1={{clade |1={{clade |1='''''Lactarius''''' |2=''Multifurca'' }} |2=''Russula'' }} |2=''Lactifluus''}} |caption=Phylogenetic relationships of ''Lactarius'', ''Lactifluus'', ''Multifurca'', and ''Russula'' according to Buyck et al. 2008.<ref name=Buyck2008/> }}

Molecular phylogenetics uncovered that, while macromorphologically well-defined, milk-caps were in fact a paraphyletic genus; as a consequence, the genera ''Lactifluus'' was split from ''Lactarius'', and the species ''L. furcatus'' was moved to the new genus ''Multifurca'', together with some former ''Russula'' species.<ref name=Buyck2010/><ref name=Buyck2008/> ''Multifurca'' also represents the likely sister group of ''Lactarius'' (see phylogeny, right). In the course of these taxonomical rearrangements, the name ''Lactarius'' was conserved for the genus with the new type species ''Lactarius torminosus''; this way, the name ''Lactarius'' could be retained for the bigger genus with many well-known temperate species, while the name ''Lactifluus'' has to be applied only to a smaller number of species, containing mainly tropical, but also some temperate milk-caps such as ''Lactifluus volemus'' and ''Lf. vellereus''.<ref name=Buyck2010/><ref name="Barrie2011"/><ref name="Norvell2011"/>

===Relationships within ''Lactarius''===

Phylogenetic analyses have also revealed that ''Lactarius'', in the strict sense, contains some species with closed (angiocarpous) fruitbodies, e.g. ''L. angiocarpus'' described from Zambia.<ref name=Eberhardt2004/> The angiocarpous genera ''Arcangeliella'' and ''Zelleromyces'' are phylogenetically part of ''Lactarius''.<ref name=Eberhardt2004/><ref name="verbeken2014"/>

Systematics within the genus ''Lactarius'' is a subject of ongoing research. Three subgenera are currently accepted and supported by molecular phylogenetics:<ref name="verbeken2013"/> *''Piperites'': Northern temperate region, three species in tropical Africa. *''Russularia'': Northern temperate region and tropical Asia. *''Plinthogalus'': Northern temperate region, tropical Africa, and tropical Asia. Some additional species, all tropical, do not seem to fall into these subgenera and occupy more basal positions within ''Lactarius''.<ref name="verbeken2014"/> This includes for example ''L. chromospermus'' from tropical Africa with an odd brown spore color.<ref name="verbeken2014"/><ref name=Buyck1995/>

Currently, over 600 species of the genus ''Lactarius'' are recognised,<ref name=kirk2014/> while the angiocarpous genera ''Arcangeliella'' and ''Zelleromyces'' have not yet been synonymized with ''Lactarius''. It is estimated that a significant number of ''Lactarius'' species remain to be described.<ref name="verbeken2013"/>

=== Selected species === {{Main|List of Lactarius species}} *''Lactarius deliciosus'' - saffron milk-cap or red pine mushroom *''Lactarius deterrimus'' - false saffron milk-cap *''Lactarius indigo'' - indigo milk-cap *''Lactarius quietus'' - oak milk-cap *''Lactarius torminosus'' - woolly milk-cap *''Lactarius turpis'' - ugly milk-cap *''Lactarius trivialis'' - dark purple or creamy brown cap

==Description==

===Macromorphology===

[[Image:Lactarius quietus 2010 G1 crop.jpg|thumb|''Lactarius quietus'' exuding cream-colored latex from gills upon cut.]]

[[Image:Lactarius alnicola 6958.jpg|thumb|Spores of ''Lactarius alnicola'' showing a reticulate (net-like) ornament with an amyloid stain reaction.]]

[[Image:2013-08-06 Lactarius indigo (Schwein.) Fr 359776.jpg|thumb|''Lactarius indigo'' is one of the most strikingly colored ''Lactarius''.]]

The eponymous "milk" and the brittle consistency of the flesh are the most prominent field characters of milk-cap fruitbodies. The milk or latex emerging from bruised flesh is often white or cream, but more vividly coloured in some species; it can change upon exposition or remain unchanged. Fruitbodies are small to very large, gilled, rather fleshy, without veil, often depressed or even funnel-shaped with decurrent gills. Cap surface can be glabrous, velvety or pilose, dry, sticky or viscose and is often zonate. Several species have pits (scrobicules) on the cap or pileus surface. Dull colors prevail, but some more colorful species exist, e.g. the blue ''Lactarius indigo'' or the orange species of section ''Deliciosi''. Spore print color is white to ocher or, in some cases, pinkish. Some species have angiocarpous, i.e., closed fruitbodies.<ref name=Eberhardt2004/>

===Micromorphology===

Microscopically, ''Lactarius'' species have elliptical, rarely globoid spores with amyloid ornamentation in the form of more or less prominent warts or spines, connected by ridges, like other members of the family Russulaceae. The trama (flesh) contains spherical cells that cause the brittle structure. Unlike ''Russula'', ''Lactarius'' also have lactiferous, i.e. latex-carrying hyphae in their trama.

===Species identification=== Distinguishing ''Lactarius'' from ''Lactifluus'' based on morphology alone is difficult; there are no synapomorphic characters known so far that define both genera unequivocally but tendencies exist:<ref name="verbeken2013"/> zonate and viscose to glutinose caps are only found in ''Lactarius'', as well as closed (angiocarpous) and sequestrate fruitbodies. All known annulate and pleurotoid (i.e., laterally stiped) milk-caps, on the contrary, belong to ''Lactifluus''.

Characters important for identification of milk-caps (''Lactarius'' and ''Lactifluus'') are:<ref name=Courtecuisse2013/><ref name=Eyssartier2011/><ref name=Kuo2011/> initial colour of the latex and color change, texture of cap surface, taste (mild, peppery, or bitter) of latex and flesh, odor, and microscopical features of the spores and the cap cuticle (pileipellis). The habitat and especially the type of host tree can also be critical. While there are some easily recognizable species, other species can be quite hard to determine without microscopical examination.<ref name=Kuo2011/>

==Distribution and habitat== ''Lactarius'' is one of the most prominent genera of mushroom-forming fungi in the Northern hemisphere. It also occurs natively in Northern Africa,<ref name=Courtecuisse2013/> tropical Africa,<ref name=Verbeken2002/> tropical Asia,<ref name="verbeken2014"/><ref name="Le2007"/> Central America,<ref name=Halling2002/> and Australia.<ref name="miller1986"/> Its possible native distribution in South America and different parts of Australasia is unclear, as many species in those regions, poorly known, might in fact belong to ''Lactifluus'', which has a more tropical distribution than ''Lactarius''.<ref name=ugent-lactifluus/> Several species have also been introduced with their host trees outside their native range,<ref name="vellinga2009"/> e.g. in South America,<ref name="Sa2013"/> Southern Africa,<ref name=Verbeken2002/> Australia,<ref name="Dunstan 1998"/> and New Zealand.<ref name="mcnabb1971"/>

==Ecology== ''Lactarius'' belongs to a lineage of ectomycorrhiza obligate symbionts.<ref name=Rinaldi2008/> As such, they are dependent on the occurrence of possible host plants. Confirmed habitats apart from temperate forests include arctic tundra and boreal forest,<ref name=Geml2009/> mediterranean maquis,<ref name=Courtecuisse2013/><ref name=Comandini2006/><ref name="Nuytinck2004"/> tropical African shrubland,<ref name=Verbeken2002/> tropical Asian rainforest,<ref name="verbeken2014"/><ref name="Le2007"/> mesoamerican tropical oak forests,<ref name=Halling2002/> and Australian ''Eucalyptus'' forests.<ref name="miller1986"/>

[[Image:Lactarius pyrogallus.jpg|thumb|''Lactarius pyrogalus'' mainly associates with common hazel.|left]]

While most species display a preference towards either broadleaf or coniferous hosts,<ref name=Courtecuisse2013/><ref name=Eyssartier2011/> some are more strictly associated with certain genera or species of plant hosts. A well-studied example is that of alders, which have several specialized ''Lactarius'' symbionts (e.g. ''L. alpinus'', ''L. brunneohepaticus'', ''L. lilacinus''), some of which even evolved specificity to one of the ''Alnus'' subgenera.<ref name=Rochet2011/> Other examples of specialized associations of ''Lactarius'' are with ''Cistus'' shrubs (''L. cistophilus'' and ''L. tesquorum''),<ref name=Comandini2006/><ref name="Nuytinck2004"/> beech (e.g. ''L. blennius''), birches (e.g. ''L. pubescens''), hazel (e.g. ''L. pyrogalus''), oak (e.g. ''L. quietus''), pines (e.g. ''L. deliciosus''), or fir (e.g. ''L. deterrimus''). For most tropical species, host plant range is poorly known, but species in tropical Africa seem to be rather generalist.<ref name=Verbeken2002/>

''Lactarius'' species are considered late-stage colonizers, that means, they are generally not present in early-colonizing vegetation, but establish in later phases of succession.<ref name="visser1995"/> However, species symbiotic with early colonizing trees, such as ''L. pubescens'' with birch, will rather occur in early stages.<ref name=Twieg2007/> Several species have preferences regarding soil pH and humidity,<ref name=Courtecuisse2013/><ref name=Eyssartier2011/> which will determine the habitats in which they occur.

==Edibility== [[File:2005-10-29 market stall with Lactarius deliciosus.jpg|thumb|right|''Lactarius deliciosus'' for sale on a market in Barcelona, Spain]] Several ''Lactarius'' species are edible. ''L. deliciosus'' notably ranks among the most highly valued mushrooms in the Northern hemisphere, while opinions vary on the taste of other species, such as ''L. indigo'' or ''L. deterrimus''. Several species are reported to be regularly collected for food in Russia, Tanzania and Hunan, China.<ref name=Harkonen2002/> Some ''Lactarius'' are considered toxic, for example ''L. turpis'', which contains the mutagenic compound necatorin,<ref name=Suortti1983/> or ''L. helvus''. There are, however, no deadly poisonous mushrooms in the genus. Bitter or peppery species, for example ''L. torminosus'', are generally not considered edible, at least raw, but are nevertheless consumed in some regions, e.g. in Finland.<ref name=Vetelainen2008/> Some small, fragrant species, such as the "candy caps", are sometimes used as flavoring.

''L. deliciosus'' is one of the few ectomycorrhizal mushrooms that has been successfully cultivated.<ref name=Guerin-Laguette2014/><ref name=effnz/>

==Chemistry== Different bioactive compounds have been isolated from ''Lactarius'' species, such as sesquiterpenoids,<ref name=Anke1989/> aromatic volatiles,<ref name="Rapior2000"/><ref name=wood2012/> and mutagenic substances.<ref name=Suortti1983/> Pigments have been isolated from colored ''Lactarius'' species, such as ''L. deliciosus''<ref name="yang2006"/> or ''L. indigo''.<ref name=Harmon1979/>

==Pharmacology== An extract of ''Lactarius badiosanguineus'' exhibits inhibitory activity on thrombin.<ref>{{cite journal |last1=Doljak |first1=B. |last2=Stegnar |first2=M. |last3=Urleb |first3=U. |last4=Kreft |first4=S. |last5=Umek |first5=A. |last6=Ciglarič |first6=M. |last7=Štrukelj |first7=B. |last8=Popovič |first8=T. |title=Screening for selective thrombin inhibitors in mushrooms |journal=Blood Coagulation and Fibrinolysis |volume=12 |issue=2 |pages=123–8 |year=2001 |pmid=11302474 |doi=10.1097/00001721-200103000-00006 |s2cid=28411589 }}</ref>

==See also== *List of ''Lactifluus'' species

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

<ref name=mycobank-lactarius>{{Cite web |title=MycoBank: ''Lactarius'' |url=http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=56791&Fields=All |access-date=2014-09-28}}</ref>

<ref name=Persoon1797>{{cite book |author=Persoon CH. |year=1797 |title=Tentamen dispositionis methodicae Fungorum |language=la}}</ref>

<ref name=Buyck2008>{{Cite journal |vauthors=Buyck B, Hofstetter V, Eberhardt U, Verbeken A, Kauff F |title=Walking the thin line between ''Russula'' and ''Lactarius'': the dilemma of ''Russula'' sect. ''Ochricompactae'' |journal=Fungal Diversity |volume=28 |year=2008 |pages=15–40 |url=http://www.fungaldiversity.org/fdp/sfdp/28-2.pdf }}</ref>

<ref name=Buyck2010>{{Cite journal |vauthors=Buyck B, Hofstetter V, Verbeken A, Walleyn R |title=Proposal to conserve ''Lactarius'' nom. cons. (Basidiomycota) with conserved type |journal=Taxon |volume=59 |year=2010 |pages=447–453 |doi=10.1002/tax.591031 |doi-access=free }} {{open access}}</ref>

<ref name="Barrie2011">{{Cite journal |author=Barrie F. |title=Report of the General Committee: 11 |journal=Taxon |volume=60 | issue=4 | year=2011 |pages=1211–1214 |doi=10.1002/tax.604026 |doi-access=free }} {{open access}}</ref>

<ref name="Norvell2011">{{Cite journal |author=Norvell LL. |title=Report of the Nomenclature Committee for Fungi: 16 |journal=Taxon |volume=60 |year=2011 |pages=223–226 |doi=10.1002/tax.601023 |doi-access=free }} {{open access}}</ref>

<ref name=Eberhardt2004>{{ cite journal |vauthors=Eberhardt U, Verbeken A |year=2004 |title=Sequestrate ''Lactarius'' species from tropical Africa: ''L. angiocarpus'' sp. nov. and ''L. dolichocaulis'' comb. nov. |journal=Mycological Research |volume=108 |issue=Pt 9 |pages=1042–1052 |doi=10.1017/S0953756204000784 |pmid=15506016}}</ref>

<ref name="verbeken2014">{{Cite journal |author1=Verbeken A |author2=Stubbe D |author3=van de Putte K |author4=Eberhardt U |author5=Nuytinck J. |title=Tales of the unexpected: angiocarpous representatives of the Russulaceae in tropical South East Asia |journal=Persoonia |volume=32 |year=2014 |pages=13–24 |doi=10.3767/003158514X679119|pmid=25264381 |pmc=4150074 }} {{open access}}</ref>

<ref name="verbeken2013">{{Cite journal |vauthors=Verbeken A, Nuytinck J |title=Not every milkcap is a ''Lactarius'' |journal=Scripta Botanica Belgica |volume=51 |year=2013 |pages=162–168 |url=http://fungi.fr/Html/VerbekenNuytinckLactarius.pdf }}</ref>

<ref name=Buyck1995>{{Cite journal |vauthors=Buyck B, Verbeken A |title=Studies in tropical African Lactarius species, 2: ''Lactarius chromospermus'' Pegler |journal=Mycotaxon |volume=56 |year=1995 |pages=427–442}}</ref>

<ref name=kirk2014>{{Cite web |author=Kirk PM. |title=Species Fungorum (version September 2014). In: Species 2000 & ITIS Catalogue of Life |url=http://www.catalogueoflife.org/col |access-date=2014-09-27 |archive-date=2020-05-13 |archive-url=https://web.archive.org/web/20200513162655/http://www.catalogueoflife.org/col/ |url-status=dead }}</ref>

<ref name=ugent-lactifluus>{{Cite web|title=Contrasting evolutionary patterns in two sister genera of macrofungi: ''Lactarius'' and ''Lactifluus'' |url=https://www.ugent.be/we/biology/en/research/mycology/projects/lactarius-lactifluus.htm |access-date=2014-09-27 |url-status=dead |archive-url=https://web.archive.org/web/20141006140713/https://www.ugent.be/we/biology/en/research/mycology/projects/lactarius-lactifluus.htm |archive-date=2014-10-06 }}</ref>

<ref name=Courtecuisse2013>{{cite book |vauthors=Courtecuisse R, Duhem B |title=Champignons de France et d'Europe |publisher=Delachaux & Niestlé |series=Guide Delachaux |year=2013 |location=Paris |isbn=978-2-603-02038-8 |language=fr}}</ref>

<ref name=Eyssartier2011>{{cite book |vauthors=Eyssartier G, Roux P |title=Le guide des champignons: France et Europe |publisher=Editions Belin |year=2011 |location=Paris |isbn=978-2-7011-5428-2 |language=fr}}</ref>

<ref name=Kuo2011>{{Cite web |author=Kuo M. |title=MushroomExpert.com: The genus ''Lactarius'' |date=2011 |url=http://www.mushroomexpert.com/lactarius.html |access-date=2014-09-28}}</ref>

<ref name=Verbeken2002>{{cite book |vauthors=Verbeken A, Buyck B |year=2002 |title=Diversity and ecology of tropical ectomycorrhizal fungi in Africa. In: Tropical Mycology: Macromycetes (eds. Watling R, Frankland JC, Ainsworth AM, Isaac S, Robinson CH.) |url=http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=26 |format=PDF |pages=11–21 |access-date=2014-09-28 |archive-url=https://web.archive.org/web/20141006165405/http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=26 |archive-date=2014-10-06 |url-status=dead }}</ref>

<ref name="Le2007">{{Cite journal |vauthors=Le HT, Stubbe D, Verbeken A, Nuytinck J, Lumyong S, Desjardin DE |title=''Lactarius'' in Northern Thailand: 2. ''Lactarius'' subgenus ''Plinthogali'' |journal=Fungal Diversity |volume=27 |year=2007 |pages=61–94|url=https://www.researchgate.net/publication/259599758 |format=PDF}}</ref>

<ref name=Halling2002>{{cite book |vauthors=Halling RE, Mueller GM |year=2002 |title=Agarics and boletes of neotropical oakwoods. In: Tropical Mycology: Macromycetes (eds. Watling R, Frankland JC, Ainsworth AM, Isaac S, Robinson CH.) |url=http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=16 |format=PDF |pages=1–10 |access-date=2014-09-28 |archive-url=https://web.archive.org/web/20141006165405/http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=16 |archive-date=2014-10-06 |url-status=dead }}</ref>

<ref name="miller1986">{{Cite journal |author1=Miller OK Jr |author2=Hilton RN. |title=New and interesting agarics from Western Australia |journal=Sydowia |volume=39 |year=1986 |pages=126–137 |url=http://www.landesmuseum.at/pdf_frei_remote/Sydowia_39_0126-0137.pdf }}</ref>

<ref name="vellinga2009">{{Cite journal |vauthors=Vellinga EC, Wolfe BE, Pringle A |title=Global patterns of ectomycorrhizal introductions |journal=New Phytologist |volume=181 |year=2009 |pages=960–973 |doi=10.1111/j.1469-8137.2008.02728.x |pmid=19170899 |issue=4|doi-access= }}</ref>

<ref name="Sa2013">{{Cite journal |vauthors=Sà MC, Baseia IG, Wartchow F |title=Checklist of Russulaceae from Brazil |journal=Mycotaxon |year=2013 |pages=online 125: 303 |url=http://www.mycotaxon.com/resources/checklists/Sa-v125-checklist.pdf }}</ref>

<ref name="Dunstan 1998">{{cite journal |author=Dunstan WA, Dell B, Malajczuk. |title=The diversity of ectomycorrhizal fungi associated with introduced ''Pinus'' spp. in the Southern Hemisphere, with particular reference to Western Australia |journal=Mycorrhiza |year=1998 |volume=8 |issue=2 |pages=71–79 |doi=10.1007/s005720050215|s2cid=32374410 }}</ref>

<ref name="mcnabb1971">{{Cite journal |author=McNabb RFR. |title=The Russulaceae of New Zealand 1. ''Lactarius'' DC ex S. F. Gray |journal=New Zealand Journal of Botany |volume=9 |year=1971 |pages=46–66 |doi=10.1080/0028825X.1971.10430170|doi-access=free }}</ref>

<ref name=Rinaldi2008>{{cite journal |vauthors=Rinaldi AC, Comandini O, Kuyper TW |year=2008 |title=Ectomycorrhizal fungal diversity: separating the wheat from the chaff |journal=Fungal Diversity |volume=33 |pages=1–45 |url=http://www.fungaldiversity.org/fdp/sfdp/33-1.pdf }}</ref>

<ref name=Geml2009>{{cite journal |vauthors=Geml J, Laursen GA, Timling I, McFarland JM, Booth MG, Lennon N, Nusbaum C, Tayler DL |year=2009 |title=Molecular phylogenetic biodiversity assessment of arctic and boreal ectomycorrhizal ''Lactarius'' Pers. (Russulales; Basidiomycota) in Alaska, based on soil and sporocarp DNA |journal=Molecular Ecology |volume=18 |issue=10 |pages=2213–2227 |url=http://www.lter.uaf.edu/pdf/1320_Geml_Laursen_2009.pdf |doi=10.1111/j.1365-294X.2009.04192.x |pmid=19389163 |s2cid=8274584 |access-date=2014-09-28 |archive-url=https://web.archive.org/web/20150910064227/http://www.lter.uaf.edu/pdf/1320_Geml_Laursen_2009.pdf |archive-date=2015-09-10 |url-status=dead }}</ref>

<ref name=Comandini2006>{{cite journal |vauthors=Comandini O, Contu M, Rinaldi AC |year=2006 |title=An overview of ''Cistus'' ectomycorrhizal fungi |journal=Mycorrhiza |volume=16 |pages=381–395 |url=http://mycorrhiza.ag.utk.edu/reviews/rev_coman1.pdf |doi=10.1007/s00572-006-0047-8 |pmid=16896800 |issue=6 |s2cid=195074078 |url-status=dead |archive-url=https://web.archive.org/web/20160304073409/http://mycorrhiza.ag.utk.edu/reviews/rev_coman1.pdf |archive-date=2016-03-04 }}</ref>

<ref name="Nuytinck2004">{{Cite journal |author1=Nuytinck J |author2=Verbeken A |author3=Rinaldi AC |author4=Leonardi M |author5=Pacioni G |author6=Comandini O. |title=Characterization of ''Lactarius tesquorum'' ectomycorrhizae on ''Cistus'' sp. and molecular phylogeny of related European ''Lactarius'' taxa |journal=Mycologia |volume=96 |year=2004 |pages=272–282 |url=http://www.mycologia.org/content/96/2/272.short |issue=2 |doi=10.2307/3762063 |pmid=21148854|jstor=3762063 |url-access=subscription }}</ref>

<ref name=Rochet2011>{{cite journal |vauthors=Rochet J, Moreau PA, Manzi S, Gardes M |year=2011 |title=Comparative phylogenies and host specialization in the alder ectomycorrhizal fungi ''Alnicola'', ''Alpova'' and ''Lactarius'' (Basidiomycota) in Europe |journal=BMC Evolutionary Biology |volume=11 |pages=40 |doi=10.1186/1471-2148-11-40 |pmid=21306639 |pmc=3045908 |doi-access=free }}</ref>

<ref name="visser1995">{{Cite journal |author=Visser S. |title=Ectomycorrhizal fungal succession in jack pine stands following wildfire |journal=New Phytologist |volume=129 |issue=3 |year=1995 |pages=389–401 |doi=10.1111/j.1469-8137.1995.tb04309.x|s2cid=85061542 |doi-access=free }}</ref>

<ref name=Twieg2007>{{cite journal |vauthors=Twieg BD, Durall DM, Simard SW |year=2007 |title=Ectomycorrhizal fungal succession in mixed temperate forests |journal=New Phytologist |volume=176 |pages=437–447 |doi=10.1111/j.1469-8137.2007.02173.x |pmid=17888121 |issue=2|doi-access= }}</ref>

<ref name=Harkonen2002>{{cite book |author=Härkonen M. |year=2002 |title=Mushroom collecting in Tanzania and Hunan (Southern China): Inherited wisdom and folklore of two different cultures. In: Tropical Mycology: Macromycetes (eds. Watling R, Frankland JC, Ainsworth AM, Isaac S, Robinson CH.) |url=http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=149 |format=PDF |pages=149–165 |access-date=2014-09-28 |archive-url=https://web.archive.org/web/20141006165405/http://fmedicine.ajums.ac.ir/_fmedicine/documents/Tropical%20mycology%20volume%201,%20macromycetes_20130417_083537.pdf#page=149 |archive-date=2014-10-06 |url-status=dead }}</ref>

<ref name=Suortti1983>{{cite journal |vauthors=Suortti T, von Wright A, Koskinen A |year=1983 |title=Necatorin, a highly mutagenic compound from ''Lactarius necator'' |journal=Phytochemistry |volume=22 |issue=12 |pages=2873–2874 |doi=10.1016/S0031-9422(00)97723-9|bibcode=1983PChem..22.2873S }}</ref>

<ref name=Vetelainen2008>{{cite report |vauthors=Veteläinen M, Huldén M, Pehu T |title=State of plant genetic resources for food and agriculture in Finland. Second Finnish National Report |work=Country Report on the State of Plant Genetic Resources for Food and Agriculture |date=2008 |publisher=Ministry of Agriculture and Forestry |location=Sastamala, Finland |page=14 |url=http://www.fao.org/docrep/013/i1500e/Finland.pdf }}</ref>

<ref name=Guerin-Laguette2014>{{cite journal |author1=Guerin-Laguette A. |author2=Cummings N. |author3=Butler R.C. |author4=Willows A. |author5=Hesom- Williams N. |author6=Li S. |author7=Wang Y. | year=2014|title=''Lactarius deliciosus'' and ''Pinus radiata'' in New Zealand: towards the development of innovative gourmet mushroom orchards|journal=Mycorrhiza |volume=24 |issue=7 |pages=511–523 |doi=10.1007/s00572-014-0570-y|pmid=24676792 |s2cid=13077838 }}</ref>

<ref name=effnz>{{Cite web |title=Edible Forest Fungi New Zealand |url=http://www.effnz.co.nz/options.htm |access-date=2014-09-28}}</ref>

<ref name=Anke1989>{{Cite journal |vauthors=Anke H, Bergendorff O, Sterner O |title=Assays of the biological activities of guaiane sesquiterpenoids isolated from the fruit bodies of edible ''Lactarius'' species |journal=Food and Chemical Toxicology |volume=27| year=1989 |pages=393–397 |doi=10.1016/0278-6915(89)90145-2 |pmid=2792969 |issue=6}}</ref>

<ref name="Rapior2000">{{Cite journal |vauthors=Rapior S, Fons F, Bessière JM |title=The fenugreek odor of ''Lactarius helvus'' |journal=Mycologia |volume=92 |issue=2 |year=2000 |pages=305–308 |jstor=3761565 |doi=10.2307/3761565}}</ref>

<ref name=wood2012>{{Cite journal |author1=Wood WF |author2=Brandes JA |author3=Foy BD |author4=Morgan CG |author5=Mann TD |author6=DeShazer DA. |title=The maple syrup odour of the "candy cap" mushroom, ''Lactarius fragilis'' var. ''rubidus'' |journal=Biochemical Systematics and Ecology |volume=43 |year=2012 |pages=51–53 |doi=10.1016/j.bse.2012.02.027}}</ref>

<ref name=Harmon1979>{{cite journal |vauthors=Harmon AD, Weisgraber KH, Weiss U |year=1979 |title=Preformed azulene pigments of ''Lactarius indigo'' (Schw.) Fries (Russulaceae, Basidiomycetes) |journal=Cellular and Molecular Life Sciences |volume=36 |pages=54–56 |doi=10.1007/BF02003967|s2cid=21207966 }}</ref> subject

<ref name="yang2006">{{Cite journal |vauthors=Yang XL, Luo DQ, Dong ZJ, Liu JK |title=Two new pigments from the fruiting bodies of the basidiomycete ''Lactarius deliciosus'' |journal=Helvetica Chimica Acta |volume=89 |issue=5 |year=2006 |pages=988–990 |url=http://www.mycochem.cn/Article/UploadFiles/200705/20070517093926795.pdf |doi=10.1002/hlca.200690103 |access-date=2014-09-28 |archive-url=https://web.archive.org/web/20141006103107/http://www.mycochem.cn/Article/UploadFiles/200705/20070517093926795.pdf |archive-date=2014-10-06 |url-status=dead }}</ref>

<ref name=Lee2019>{{Cite journal |last1=Lee |first1=Hyun |last2=Wissitrassameewong |first2=Komsit |last3=Smythe |first3=Jim |last4=Myung Soo |first4=Park |last5=Verbeken |first5=Annemieke |last6=Eimes |first6=John |last7=Lim |first7=Young Woon |title=Taxonomic revision of the genus Lactarius (Russulales, Basidiomycota) in Korea |journal=Fungal Diversity |date=2019-05-10 |volume=95 |pages=275–335 |doi=10.1007/s13225-019-00425-6 |s2cid=149443885 |quote=To date, 583 Lactarius species have been recorded globally |doi-access=free |hdl=1854/LU-8620961 |hdl-access=free }}</ref>

}}

==External links== * [http://www.hti.umich.edu:80/cgi/t/text/text-idx?c=fung1tc;cc=fung1tc;view=toc;idno=AAC3719.0001.001 ''North American species of Lactarius''] by L. R. Hesler and Alexander H. Smith, 1979 (full text of monograph).

Category:Lactarius Category:Russulales genera