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| genus = '''''Lactarius'''''
| genus = '''''Lactarius'''''
| genus_authority = [[Christian Hendrik Persoon|Pers.]] 1797
| genus_authority = [[Christian Hendrik Persoon|Pers.]] 1797
| synonyms_ref = <ref name=mycobank-lactarius/>
| diversity = c.400 species
| synonyms =
{{plainlist|
*''Lactaria'' <small>[[Pers.]] (1797)</small>
*''Agaricus'' sect. ''Lactifluus'' <small>[[Pers.]] (1801)</small>
*''Agaricus'' subdiv. ''Galorrheus'' <small>[[Fr.]] (1818)</small>
*''Lactariella'' <small>[[J. Schröt.]] (1898)</small>
*''Lactariopsis'' <small>[[Henn.]] (1901)</small>
*''Gloeocybe'' <small>Earle (1909)</small>
*''Hypophyllum'' <small>Earle (1909)</small>
}}
| diversity = c. 350 species
| diversity_link = List of Lactarius species
| diversity_link = List of Lactarius species
}}
}}
'''''Lactarius''''' is a [[genus]] of [[mushroom]]-producing [[fungi]]. The genus, collectively known [[common name|commonly]] as '''milk-caps''', are characterized by the fact that they exude a milky fluid ('latex') if cut or damaged. Like the genus ''[[Russula]]'', with which they are grouped in the family '''[[Russulaceae]]''', their flesh has a distinctive brittle consistency.


'''''Lactarius''''' is a [[genus]] of [[mushroom]]-producing, [[Ectomycorrhiza|ectomycorrhizal]] [[fungi]], containing several [[Edible mushroom|edible]] species. The species of the genus, [[common name|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 roughly 350 known species, mainly distributed in the Northern hemisphere. Recently, the genus ''[[Lactifluus]]'' has been separated from ''Lactarius'' based on [[Molecular phylogenetics|molecular phylogenetic]] evidence.
Often the [[Gill (mushroom)|gills]] are decurrent (starting to run down the stem) and the cap is depressed or even funnel-shaped when older. The stem and cap sometimes show 'strobicules' (or 'strobiculi'), which are flat-based shallow oval pits.


==Nomenclature==
To identify to which of the roughly 400 ''Lactarius'' species a given specimen belongs, note whether the cap is bald/greasy or velvety or hairy/shaggy at the rim, particularly in young individuals. Also the initial colour of the milk (white, cream, orange, violet, ...) and the final colour on drying are determining characteristics.


The genus was described by [[Christian Hendrik Persoon]] in 1797. ''Lactarius'' is derived from the [[Latin]] ''[[wikt:lac#Latin|lac]]'' '[[milk]]'.
The genus ''Lactarius'' was described by [[Christian Hendrik Persoon|C. H. Persoon]] in 1797<ref name=Persoon1797/> with ''[[Lactifluus piperatus|L. piperatus]]'' as the original type species. In 2010, ''[[Lactarius torminosus|L. torminosus]]'' was accepted as the new type of the genus after the splitting-off of ''Lactifluus'' as separate genus (see Systematics section).

The name "''Lactarius''" is derived from the [[Latin]] ''[[wikt:lac#Latin|lac]]'', "[[milk]]".

==Description==

===Macromorphology===

[[Image:Lactarius_quietus_2010_G1_crop.jpg|thumb|''[[Lactarius quietus]]'' exuding cream-colored latex from gills upon cut.|left]]

[[Image:2013-08-06_Lactarius_indigo_(Schwein.)_Fr_359776.jpg|thumb|''[[Lactarius indigo]]'' is one of the most strikingly colored ''Lactarius''.|right]]

The eponymous "milk" and the brittle consistency of the flesh are the most prominent field characters of milk-cap [[Sporocarp (fungi)|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 (mycology)|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 pink ''[[Lactarius chrysorrheus|L. chrysorrheus]]''. [[Spore print]] color is white to ocre or, in some cases, pinkish. Some species have angiocarp, i.e., closed fruitbodies.<ref name=Eberhardt2004/>

===Micromorphology===

[[Image:Lactarius_alnicola_6958.jpg|thumb|Spores of ''[[Lactarius alnicola]]'' showing a reticulate (net-like) ornament with an amyloid stain reaction.|right]]

Microscopically, ''Lactarius'' species have elliptical, rarely globoid spores with [[Amyloid (mycology)|amyloid]] ornamentation in the form of more or less prominent warts or spines, connected by ridges, like other members of the [[Russulaceae]] family. The [[Trama (mycology)|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===

Separating ''Lactarius'' from ''[[Lactifluus]]'' based on morphology alone is difficult; there are no [[Synapomorphy|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 adn 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 curticle ([[Pileipellis|pileipellis]]). The habitat and especially the type of host tree can also be critical. While there are some easily recognizable species, others can be quite hard to determine without microscopical examination.<ref name=Kuo2011/>

==Distribution==

''Lactarius'' is one of the most prominent genera of mushroom-forming fungi in the Northern hemisphere. It also occcurs [[Indigenous (ecology)|natively]] in Northern Africa,<ref name=Courtecuisse2013/> tropical Africa,<ref name=Verbeken2002/> tropical Asia,<ref name="Le2007"/><ref name="verbeken2014"/> and Central America.<ref name=Halling2002/> Its possible native distribution in South America and Oceania 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=australia-deliciosus/> and New Zealand.<ref name="mcnabb1971"/>


==Ecology==
==Ecology==


''Lactarius'' belongs to a lineage of obligate [[Ectomycorrhiza|ectomycorrhizal]] [[Symbiosis|symbionts]].<ref name=Rinaldi2008/> As such, they are dependent on the occurrence of possible host plants. Confirmed habitats apart from [[Temperate forest|temperate forests]] include arctic [[Tundra|tundra]] and [[Taiga|boreal forest]],<ref name=Geml2009/> mediterranean [[Maquis shrubland|maquis]],<ref name=Comandini2006/><ref name="Nuytinck2004"/><ref name=Courtecuisse2013/> [[Miombo|tropical African shrubland]],<ref name=Verbeken2002/> [[Tropical rainforest|tropical Asian rainforest]],<ref name="Le2007"/><ref name="verbeken2014"/> mesoamerican tropical oak forests,<ref name=Halling2002/> and Australian ''[[Eucalyptus]]'' forests.<ref name="miller1986"/>
Lactarius is one of the most prominent genera of ectomycorrhizal fungi.<ref>{{cite journal |last1=Rinaldi |first1=A.C. |last2=Comandini |first2=O.|last3=Kuyper |first3=T.W. |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> With more than 400 species described worldwide, Lactarius taxa play a significant role as late-stage colonizers of trees and shrubs in a vast range of ecosystems, from boreal coniferous forests to temperate Mediterranean-type maquis. An interesting case is that of the strict association of selected ''Lactarius'' species - such as ''L. tesquorum'' and ''L. cistophilus'' - with ''[[Cistus]]'' spp., a group of about 20 woody, evergreen or semideciduous shrub species found in wide semi-arid areas from the Canary Islands throughout the Mediterranean region to the Caucasus. ''[[Cistus]]'' species are obligate seeding, early colonizers that follow disturbance, particularly fire, in low maquis-type Mediterranean ecosystems, rendering their ectomycorrhizal ecology particularly intriguing.<ref>{{cite journal |last1=Nuytinck |first1=J. |last2=Verbeken |first2=A.|last3=Rinaldi |first3=A.C. |last4=Leonardi |first4=M.|last5=Pacioni |first5=G. |last6=Comandini |first6=O. |year=2004 |title=Characterization of ''Lactarius tesquorum'' ectomycorrhizae on ''Cistus'' sp. and molecular phylogeny of related European ''Lactarius'' taxa |journal=Mycologia |volume=96 |pages=272–282 |pmid=21148854 |url=http://www.mycologia.org/cgi/content/full/96/2/272 |issue=2 |doi=10.2307/3762063}}</ref><ref>{{cite journal |last1=Comandini |first1=O. |last2=Rinaldi |first2=A.C. |year=2008 |title=''Lactarius cistophilus'' Bon & Trinbach + ''Cistus'' sp |journal=Descriptions of Ectomycorrhizae |volume=96 |pages=83–88 }}</ref>


[[Image:Lactarius_pyrogallus.jpg|thumb|''[[Lactarius pyrogalus]]'' mainly associates with [[Corylus avellana|common hazel]].|left]]
==A selection of well-known European species==
[[File:Gruzd.jpg|thumb|Lactarius]]
{{Main|List of Lactarius species}}
*''[[Lactarius turpis]]'' - ugly milk-cap
*''[[Lactarius torminosus]]'' - woolly milk-cap
*''[[Lactarius piperatus]]'' - peppery milk-cap
*''[[Lactarius uvidus]]''


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 [[Alnus|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"/> [[Fagus sylvatica|beech]] (e.g. ''[[Lactarius blennius|L. blennius]]''), [[Betula|birches]] (e.g. ''[[Lactarius pubescens|L. pubescens]]''), [[Corylus|hazel]] (e.g. ''[[Lactarius pyrogalus|L. pyrogalus]]''), [[Quercus|oak]] (e.g. ''[[Lactarius quietus|L. quietus]]''), [[Pinus|pines]] (e.g. ''[[Lactarius deliciosus|L. deliciosus]]''), or [[Picea abies|fir]] (e.g. ''[[Lactarius deterrimus|L. deterrimus]]''). For most tropical species, host plant range is poorly known, but species seem to be rather generalist in tropical Africa.<ref name=Verbeken2002/>
===Section ''Deliciosi''===
*''[[Lactarius deliciosus]]''
*''[[Lactarius deterrimus]]''
*''[[Lactarius salmonicolor]]''
{{Clear}}
[[Image:Lactarius salmonicolor.jpg|thumb|110px|left|''Lactarius salmonicolor'']]
This group of mushrooms has orange or red milk and carrot-orange coloration, in many cases becoming flecked with green or blue-green with age. They are all generally considered to be edible and tasty, the appetizingly named ''Lactarius deliciosus'' reputedly being much better than the others. They grow in coniferous woods.


''Lactarius'' species are considered late-stage colonizers, that means, they are generally not present in early-colonizing vegetation, but establish in later phases of [[Ecological succession|succession]].<ref name="visser1995"/> However, species symbiotic with early colonizing trees, such as ''[[Lactarius pubescens|L. pubescens]]'' with [[Betula|birch]], will rather occur in early successional 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.
''L. deliciosus'' is the only one with an English name: 'saffron milk-cap'. It is found under pine or spruce. ''L. deterrimus'' was formerly considered to be a variety of ''L. deliciosus'' and is distinguished by the milk turning wine-red on drying - but only after up to 30 minutes. ''L. salmonicolor'' is associated with spruce and remains a pure orange, unlike the other two which discolour green.


==Systematics and taxonomy==
===''Lactarius chrysorrheus''===
[[Image:Lactarius chrysorreus 041113w.jpg|120px|right]]
The milk of this mushroom emerges white but immediately changes to a deep yellow colour. The top of the cap is a pinkish red with concentric zoning on a background showing yellow tints. It is common under broad-leaved trees, especially oak. There are conflicting reports as to its edibility.


===Placement within Russulaceae===
===''Lactarius decipiens''===
[[Image:Lactarius decipiens 031111w.jpg|120px|left]]
This flesh-coloured Lactarius is closely related to ''L. chrysorrheus'', and again the milk is initially white and turns yellow, but only after several seconds. To allow the milk to contact the air this is best tested on a paper tissue (a cloth handkerchief is liable to be permanently stained).


{{cladogram|align=right|title=
===''Lactarius quietus''===
|clade=
[[Image:Lactarius quietus 051106w.jpg|120px|right]]
{{clade
''L. quietus'', which is commonly found under oak trees, is often thought to be a non-descript mushroom, and in fact that is the significance of its Latin epithet ''quietus''. It has a distinctive smell said to be of bedbugs or of wet laundry. The milk goes a little cream but does not change further.
|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. 2010.<ref name=Buyck2008/>
}}

[[Molecular phylogenetics]] uncovered that, while macromorphologically well-defined, milk-caps were in fact a [[Paraphyly|paraphyletic]] genus; as a consequence, the genera ''[[Lactifluus]]'' was split from ''Lactarius'', and the species ''[[Multifurca furcata|L. furcatus]]'' was moved to the new genus ''[[Multifurca]]'', together with some former ''Russula'' species.<ref name=Buyck2008/><ref name=Buyck2010/> ''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 (biology)|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 ''[[Lactifluus vellereus|Lf. vellereus]]''.<ref name=Buyck2010/>

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

[[Image:2011-07-25_Arcangeliella_crassa_Singer_%26_A.H._Smith_159495_crop.jpg|thumb|''[[Arcangiella crassa]]'' is one of the milk-caps with closed fruitbodies that are phylogenetically nested within ''Lactarius''.|left]]

Phylogenetic analyses have also revealed that ''Lactarius'', in the strict sense, contains some species with closed (angiocarpous) fruitbodies, e.g. ''[[Lactarius angiocarpus|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 ''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 more 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 ''[[Lactarius chromospermus|L. chromospermus]]'' from tropical Africa with an odd brown spore color.<ref name=Buyck1995/><ref name="verbeken2014"/>

Currently, around 600 Lactarius species are described,<ref name=kirk2014/> but roughly one fourth or 150 of these are believed to belong to ''Lactifluus'',<ref name=ugent-lactifluus/> while the angiocarpous genera ''Arcangeliella'' and ''Zelleromyces'' have not yet been [[Synonym (taxonomy)|synonymized]] with ''Lactarius''. It is estimated that a significant number of ''Lactarius'' species remain to be described.<ref name="verbeken2013"/>

==Edibility==

[[Image:2005-10-29_market_stall_with_Lactarius_deliciosus.jpg|thumb|''[[Lactarius deliciosus]]'' for sale on a market in Barcelona, Spain.|left]]

[[Image:081117_hongos.JPG|thumb|''Lactarius'' fruitbodies, including the blue ''[[L. indigo]]'', on a market in Guatemala.|right]]

Several ''Lactarius'' species are edible. ''[[Lactarius deliciosus|L. deliciosus]]'' notably ranks among the most highly valued mushrooms in the Northern hemisphere, while opinions vary on the taste of others, such as ''[[Lactarius indigo|L. indigo]]'' or ''[[Lactarius deterrimus|L. deterrimus]]''. Several species are reported to be regularly collected for food in Tanzania and Hunan, China.<ref name=Harkonen2002/> Some ''Lactarius'' are considered toxic, for example ''[[Lactarius turpis]]'', which contains a mutagenic compound,<ref name=Suortti1983/> or ''[[Lactarius helvus|L. helvus]]''. There are, however, no deadly poisonous mushrooms in the genus. Bitter or peppery species, for example ''[[Lactarius torminosus|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 [[Sesquiterpene|sesquiterpenoids]],<ref name=Anke1989/> aromatic [[Volatile organic compound|volatiles]]<ref name="Rapior2000"/><ref name=wood2012/>, and mutagenic substances.<ref name=Suortti1983/> [[Pigment|Pigments]] have been isolated from colored ''Lactarius'' species, such as ''L. deliciosus''<ref name="yang2006"/> or ''[[L. indigo]]''.<ref name=Harmon1979/>

==A selection of well-known 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

==Some ''Lactifluus'' species formerly classified in ''Lactarius''==
{{Main|List of Lactifluus species}}
*''[[Lactifluus glaucescens]]''
*''[[Lactifluus hygrophoroides]]''
*''[[Lactifluus gerardii]]''
*''[[Lactifluus piperatus]]''
*''[[Lactifluus vellereus]]'' - fleecy milk-cap
*''[[Lactifluus volemus]]'' - weeping milk-cap


==See also==
==See also==
*[[List of Lactarius species|List of ''Lactarius'' species]]
*[[List of Lactarius species|List of ''Lactarius'' species]]
*''[[Lactifluus]]''
*''[[Multifurca]]''
*''[[Russula]]''


==References==
==References==

{{Reflist}}
{{Reflist|colwidth=30em|refs=
*Partly from Dutch Wikipedia pages.

*Meinhard Moser, translated by Simon Plant: ''Keys to Agarics and Boleti'' (Roger Phillips 1983)
<ref name=Anke1989>{{Cite journal
*Régis Courtecuisse, Bernard Duhem : ''Guide des champignons de France et d'Europe'' (Delachaux & Niestlé, 1994–2000). ISBN 2-603-00953-2
|author=Anke H., Bergendorff O. & Sterner O.
*Roger Phillips : ''Mushrooms and other fungi of Great Britain and Europe'' (Pan Books Ltd. 1981 / Book Club Associates 1981) - for the English names.
|title=Assays of the biological activities of guaiane sesquiterpenoids isolated from the fruit bodies of edible ''Lactarius'' species
*{{cite journal | last1 = Verbeken | first1 = Annemieke | year = 2001 | title = Worldwide systematics of ''Lactarius'': a state of art | url = | journal = Micologia e vegetazione mediterranea | volume = 16 | issue = 1| pages = 71–88 }}
|journal=[[Food and Chemical Toxicology]]
{{Commons|Lactarius}}
|volume=27
|year=1989
|pages=393–397
|url=http://www.sciencedirect.com/science/article/pii/0278691589901452
|doi=10.1016/0278-6915(89)90145-2
}}</ref>

<ref name=australia-deliciosus>{{Cite website
|author=
|title=Australian Fungi - A blog: #15 Lactarius deliciosus
|url=http://australianfungi.blogspot.fr/2007/05/15-lactarius-deliciosus.html
|accessdate=2014-09-28
}}</ref>

<ref name=Buyck1995>{{Cite journal
|author=Buyck B. & Verbeken A.
|title=Studies in tropical African Lactarius species, 2: ''Lactarius chromospermus'' Pegler
|journal=[[Mycotaxon]]
|volume=56
|year=1995
|pages=427-442
|url=https://biblio.ugent.be/publication/250973
}}</ref>

<ref name=Buyck2008>{{Cite journal
|author=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
|author=Buyck B., Hofstetter V., Verbeken A. & Walleyn R.
|title=Proposal to conserve ''Lactarius'' nom. cons. (Basidiomycota) with conserved type
|journal=[[International_Association_for_Plant_Taxonomy#Taxon|Taxon]]
|volume=118
|year=2010
|pages=447–453
}}</ref>

<ref name=Buyck2010>{{Cite journal
|author=Buyck B., Hofstetter V., Verbeken A. & Walleyn R.
|title=Proposal to conserve ''Lactarius'' nom. cons. (Basidiomycota) with conserved type
|journal=[[International_Association_for_Plant_Taxonomy#Taxon|Taxon]]
|volume=118
|year=2010
|pages=447–453
}}</ref>

<ref name=Comandini2006>{{cite journal
|author=Comandini O., Contu M. & Rinaldi A.C.
|year=2006
|title=An overview of ''Cistus'' ectomycorrhizal fungi
|journal=[[Mycorrhiza (journal)|Mycorrhiza]]
|volume=16
|pages=381-395
|url=http://mycorrhiza.ag.utk.edu/reviews/rev_coman1.pdf
}}</ref>

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

<ref name=Eberhardt2004>{{cite journal
|author=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
|pages=1042–1052
|url=http://www.sciencedirect.com/science/article/pii/S0953756208604244
|doi=10.1017/S0953756204000784
}}</ref>

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

<ref name=Eyssartier2011>{{cite book
| author = Eyssartier G. & Roux P.
| title = Le guide des champignons: France et Europe
| publisher = Editions Belin
| series =
| volume =
| edition =
| date = 2011
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| isbn = 978-2-7011-5428-2
}}</ref>

<ref name=Geml2009>{{cite journal
|author= Geml J., Laursen G.A., Timling I., McFarland J.M., Booth M.G., Lennon N., Nusbaum C. & Tayler D.L.
|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 (journal)|Molecular Ecology]]
|volume=18
|pages=2213–2227
|url=http://www.lter.uaf.edu/pdf/1320_Geml_Laursen_2009.pdf
|doi=10.1111/j.1365-294X.2009.04192.x
}}</ref>

<ref name=Guerin-Laguette2014>{{cite journal
|author=Guerin-Laguette A., Cummings N., Butler R.C., Willows A., Hesom-Williams N., Li S. & Wang Y.
|year=2014
|title=''Lactarius deliciosus'' and ''Pinus radiata'' in New Zealand: towards the development of innovative gourmet mushroom orchards
|journal=[[Mycorrhiza (journal)|Mycorrhiza]]
|volume=24
|pages=511-523
|url=http://link.springer.com/article/10.1007%2Fs00572-014-0570-y
|doi=10.1007/s00572-014-0570-y
}}</ref>

<ref name=Halling2002>{{cite book
|author=Halling R.E. & Mueller G.M.
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<ref name=Harmon1979>{{cite journal
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<ref name=ugent-lactifluus>{{Cite website
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<ref name=Suortti1983>{{cite journal
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<ref name=Twieg2007>{{cite journal
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|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1469-8137.2007.02173.x/full
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<ref name=Verbeken2002>{{cite book
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<ref name="verbeken2014">{{Cite journal
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|author=Yang X.-L., Luo D.-Q., Dong Z.-J. & Liu J.-K.
|title=Two new pigments from the fruiting bodies of the basidiomycete ''Lactarius deliciosus''
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}}


==External links==
==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.)
* [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).
* [http://www.mtsn.tn.it/russulales-news Russulales News]
* [http://www2.muse.it/russulales-news/ Russulales News]
* [http://www.svims.ca/council/Lactar.htm Key to species of ''Lactarius'' in the Pacific Northwest]
* [http://www.mushroomexpert.com/lactarius.html The genus ''Lactarius'' on MushroomExpert.com], by Michael Kuo
* [https://www.ugent.be/we/biology/en/research/mycology/projects University of Ghent, mycology group: Research projects]
* [http://www.landcareresearch.co.nz/resources/identification/fungi/russula-lactarius Landcare Research: Keys to ''Russula'' and ''Lactarius'' species in New Zealand]
* [https://www.youtube.com/watch?v=LLFhPlbHU9A Video - Milky Mushroom: How to identify a Lactarius mushroom]

==Literature==

*Courtecuisse R. & Duhem B. 2013: ''Guide des champignons de France et d'Europe'' (Delachaux & Niestlé) – ISBN 978-2-603-02038-8
*Eyssartier G. & Roux P. 2011: Le guide des champignons: France et Europe (Editions Belin) – ISBN 978-2-7011-5428-2
*Heilmann-Clausen J., Verbeken A. & Vesterholt J. 1998: ''Fungi of Northern Europe, Volume 2: The Genus Lactarius'' (2nd ed., Svampetrik, 287 pp.) – ISBN 8798358146
*Kibby G. 2014: ''British milkcaps: Lactarius & Lactifluus'' (priv. published, 114 pp.)
*Moser M. 1983, translated by Simon Plant: ''Keys to Agarics and Boleti'' (Mad River Publishers)
*Roger Phillips 1981: ''Mushrooms and other fungi of Great Britain and Europe'' (Pan Books Ltd.)


[[Category:Russulales]]
[[Category:Russulales]]

Revision as of 14:25, 28 September 2014

For the single-species genus of fish, see False trevally.

Lactarius
Lactarius vietus
Scientific classification
Kingdom:
Fungi
Subkingdom:
Dikarya
Phylum:
Basidiomycota
Subphylum:
Agaricomycotina
Class:
Agaricomycetes
Order:
Russulales
Family:
Russulaceae
Genus:
Lactarius

Pers. 1797
Diversity
c. 350 species
Synonyms[1]
  • 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)

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 roughly 350 known species, mainly distributed in the Northern hemisphere. Recently, the genus Lactifluus has been separated from Lactarius based on molecular phylogenetic evidence.

Nomenclature

The genus Lactarius was described by C. H. Persoon in 1797[2] with L. piperatus as the original type species. In 2010, L. torminosus was accepted as the new type of the genus after the splitting-off of Lactifluus as separate genus (see Systematics section).

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

Description

Macromorphology

Lactarius quietus exuding cream-colored latex from gills upon cut.
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 pink L. chrysorrheus. Spore print color is white to ocre or, in some cases, pinkish. Some species have angiocarp, i.e., closed fruitbodies.[3]

Micromorphology

Spores of Lactarius alnicola showing a reticulate (net-like) ornament with an amyloid stain reaction.

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 Russulaceae family. 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

Separating 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:[4] zonate and viscose to glutinose caps are only found in Lactarius, as well as closed (angiocarpous) and sequestrate fruitbodies. All known annulate adn pleurotoid (i.e., laterally stiped) milk-caps, on the contrary, belong to Lactifluus.

Characters important for identification of milk-caps (Lactarius and Lactifluus) are:[5][6][7] 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 curticle (pileipellis). The habitat and especially the type of host tree can also be critical. While there are some easily recognizable species, others can be quite hard to determine without microscopical examination.[7]

Distribution

Lactarius is one of the most prominent genera of mushroom-forming fungi in the Northern hemisphere. It also occcurs natively in Northern Africa,[5] tropical Africa,[8] tropical Asia,[9][10] and Central America.[11] Its possible native distribution in South America and Oceania is unclear, as many species in those regions, poorly known, might in fact belong to Lactifluus, which has a more tropical distribution than Lactarius.[12] Several species have also been introduced with their host trees outside their native range,[13] e.g. in South America,[14] Southern Africa,[8] Australia,[15] and New Zealand.[16]

Ecology

Lactarius belongs to a lineage of obligate ectomycorrhizal symbionts.[17] As such, they are dependent on the occurrence of possible host plants. Confirmed habitats apart from temperate forests include arctic tundra and boreal forest,[18] mediterranean maquis,[19][20][5] tropical African shrubland,[8] tropical Asian rainforest,[9][10] mesoamerican tropical oak forests,[11] and Australian Eucalyptus forests.[21]

Lactarius pyrogalus mainly associates with common hazel.

While most species display a preference towards either broadleaf or coniferous hosts,[5][6] 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.[22] Other examples of specialized associations of Lactarius are with Cistus shrubs (L. cistophilus and L. tesquorum),[19][20] 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 seem to be rather generalist in tropical Africa.[8]

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.[23] However, species symbiotic with early colonizing trees, such as L. pubescens with birch, will rather occur in early successional stages.[24] Several species have preferences regarding soil pH and humidity,[5][6] which will determine the habitats in which they occur.

Systematics and taxonomy

Placement within Russulaceae

Lactarius

Multifurca

Russula

Lactifluus

Phylogenetic relationships of Lactarius, Lactifluus, Multifurca, and Russula according to Buyck et al. 2010.[25]

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.[25][26] 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.[26]

Relationships within Lactarius

Arcangiella crassa is one of the milk-caps with closed fruitbodies that are phylogenetically nested 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.[3] The angiocarpous genera Arcangeliella and Zelleromyces are phylogenetically part of Lactarius.[3][10]

Systematics within Lactarius is a subject of ongoing research. Three subgenera are currently accepted and supported by molecular phylogenetics:[4]

  • 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 more species, all tropical, do not seem to fall into these subgenera and occupy more basal positions within Lactarius.[10] This includes for example L. chromospermus from tropical Africa with an odd brown spore color.[27][10]

Currently, around 600 Lactarius species are described,[28] but roughly one fourth or 150 of these are believed to belong to Lactifluus,[12] 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.[4]

Edibility

Lactarius deliciosus for sale on a market in Barcelona, Spain.
Lactarius fruitbodies, including the blue L. indigo, on a market in Guatemala.

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 others, such as L. indigo or L. deterrimus. Several species are reported to be regularly collected for food in Tanzania and Hunan, China.[29] Some Lactarius are considered toxic, for example Lactarius turpis, which contains a mutagenic compound,[30] 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.[31] 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.[32][33]

Chemistry

Different bioactive compounds have been isolated from Lactarius species, such as sesquiterpenoids,[34] aromatic volatiles[35][36], and mutagenic substances.[30] Pigments have been isolated from colored Lactarius species, such as L. deliciosus[37] or L. indigo.[38]

A selection of well-known species

Main article: List of Lactarius species

Some Lactifluus species formerly classified in Lactarius

Main article: List of Lactifluus species

See also

References

  1. ^ "MycoBank: Lactarius". Retrieved 2014-09-28.
  2. ^ Persoon, C.H. (1797). Tentamen dispositionis methodicae Fungorum.
  3. ^ a b c Eberhardt U. & Verbeken A. (2004). "Sequestrate Lactarius species from tropical Africa: L. angiocarpus sp. nov. and L. dolichocaulis comb. nov". Mycological Research. 108: 1042–1052. doi:10.1017/S0953756204000784.
  4. ^ a b c Verbeken A. & Nuytinck J. (2013). "Not every milkcap is a Lactarius" (PDF). Scripta Botanica Belgica. 51: 162–168.
  5. ^ a b c d e Courtecuisse R. & Duhem B. (2013). Champignons de France et d'Europe. Guide Delachaux. Paris: Delachaux & Niestlé. ISBN 978-2-603-02038-8.
  6. ^ a b c Eyssartier G. & Roux P. (2011). Le guide des champignons: France et Europe. Paris: Editions Belin. ISBN 978-2-7011-5428-2.
  7. ^ a b Kuo M. (2011). "MushroomExpert.com: The genus Lactarius". Retrieved 2014-09-28.
  8. ^ a b c d Verbeken, A. & Buyck B. (2002). Diversity and ecology of tropical ectomycorrhizal fungi in Africa, in: Tropical Mycology: Macromycetes (eds. Watling R., Frankland J.C., Ainsworth A.M., Isaac S. & Robinson C.H.) (PDF). pp. 11–21.
  9. ^ a b Le H.T., Stubbe D., Verbeken A., Nuytinck J., Lumyong S. & Desjardin D.E. (2007). "Lactarius in Northern Thailand: 2. Lactarius subgenus Plinthogali" (PDF). Fungal Diversity. 27: 61–94.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ^ a b c d e Verbeken A., Stubbe D., van de Putte K., Eberhardt U. & Nuytinck J. (2014). "Tales of the unexpected: angiocarpous representatives of the Russulaceae in tropical South East Asia". Persoonia. 32: 13–24. doi:10.3767/003158514X679119.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b Halling R.E. & Mueller G.M. (2002). Agarics and boletes of neotropical oakwoods, in: Tropical Mycology: Macromycetes (eds. Watling R., Frankland J.C., Ainsworth A.M., Isaac S. & Robinson C.H.) (PDF). pp. 1–10.
  12. ^ a b "Contrasting evolutionary patterns in two sister genera of macrofungi: Lactarius and Lactifluus". Retrieved 2014-09-27.
  13. ^ Vellinga E.C., Wolfe B.E. & Pringle A. (2009). "Global patterns of ectomycorrhizal introductions". New Phytologist. 181: 960–973. doi:10.1111/j.1469-8137.2008.02728.x.
  14. ^ Sà M.C.A., Baseia I.G. & Wartchow F. (2013). "Checklist of Russulaceae from Brazil" (PDF). Mycotaxon: online 125: 303.
  15. ^ "Australian Fungi - A blog: #15 Lactarius deliciosus". Retrieved 2014-09-28.
  16. ^ McNabb R.F.R. (1971). "The Russulaceae of New Zealand 1. Lactarius DC ex S. F. Gray". New Zealand Journal of Botany. 9: 46–66. doi:10.1080/0028825X.1971.10430170.
  17. ^ Rinaldi A.C., Comandini O. & Kuyper T.W. (2008). "Ectomycorrhizal fungal diversity: separating the wheat from the chaff" (PDF). Fungal Diversity. 33: 1–45.
  18. ^ Geml J., Laursen G.A., Timling I., McFarland J.M., Booth M.G., Lennon N., Nusbaum C. & Tayler D.L. (2009). "Molecular phylogenetic biodiversity assessment of arctic and boreal ectomycorrhizal Lactarius Pers. (Russulales; Basidiomycota) in Alaska, based on soil and sporocarp DNA" (PDF). Molecular Ecology. 18: 2213–2227. doi:10.1111/j.1365-294X.2009.04192.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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Literature

  • Courtecuisse R. & Duhem B. 2013: Guide des champignons de France et d'Europe (Delachaux & Niestlé) – ISBN 978-2-603-02038-8
  • Eyssartier G. & Roux P. 2011: Le guide des champignons: France et Europe (Editions Belin) – ISBN 978-2-7011-5428-2
  • Heilmann-Clausen J., Verbeken A. & Vesterholt J. 1998: Fungi of Northern Europe, Volume 2: The Genus Lactarius (2nd ed., Svampetrik, 287 pp.) – ISBN 8798358146
  • Kibby G. 2014: British milkcaps: Lactarius & Lactifluus (priv. published, 114 pp.)
  • Moser M. 1983, translated by Simon Plant: Keys to Agarics and Boleti (Mad River Publishers)
  • Roger Phillips 1981: Mushrooms and other fungi of Great Britain and Europe (Pan Books Ltd.)
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