Suillus luteus

{{Short description|Species of edible fungus in the family Suillaceae native to Eurasia}} {{Featured article}} {{Use dmy dates|date=January 2020}} {{Speciesbox | image = Suillus luteus 475376.jpg | status = LC | status_system = IUCN3.1 | status_ref = <ref name=IUCN>{{cite iucn |last1=Dahlberg |first1=A. |year=2019 |title=''Suillus luteus'' |volume=2019 |errata=2022 |article-number=e.T122090818A223016348 |doi=10.2305/IUCN.UK.2019-3.RLTS.T122090818A223016348.en |access-date=19 October 2024}}</ref> | genus = Suillus | species = luteus | authority = (L.) Roussel (1796) | synonyms_ref = <ref name="urlFungorum synoynymy: Suillus luteus"/> | synonyms = *''Boletus luteus'' L. (1753) * ''Boletus volvatus'' Batsch (1783) * ''Cricunopus luteus'' (L.) P.Karst. (1881) * ''Viscipellis luteus'' (L.) Quél. (1886) * ''Ixocomus luteus'' (L.) Quél. (1888) * ''Boletopsis lutea'' (L.) Henn. (1898) }} {{mycomorphbox | name = ''Suillus luteus'' | hymeniumType = pores | capShape = convex | whichGills = adnate | whichGills2 = subdecurrent | stipeCharacter = ring | sporePrintColor = brown | ecologicalType = mycorrhizal | howEdible = edible }}

'''''Suillus luteus''''' is a species of bolete fungus commonly referred to as '''slippery jack''' or '''sticky bun'''. The fungus was originally assigned to ''Boletus'' by Carl Linnaeus in 1753, before being made the type species of ''Suillus'' in 1796. The slimy cap is typically up to {{convert|10|cm|0|abbr=on}} wide and conical before flattening. A membranous partial veil connects the cap to the stem when young. Yellow tubes extend downward from the underside of the cap, which spores escape through. The pale stem is up to 10 cm (4 in) tall and {{convert|3|cm|1|abbr=on|frac=4}} thick, bearing small dots near the top. Unlike most other boletes, it bears a distinctive membranous ring, tinged brown to violet on the underside.

A common fungus native all across Eurasia from Ireland to Korea, it has been introduced widely elsewhere, including North and South America, southern Africa, Australia and New Zealand. The fungus grows in coniferous forests in its native range, and pine plantations in countries where it has become naturalized. It forms symbiotic ectomycorrhizal associations with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. The fruit bodies are often produced in large numbers above ground in summer and autumn.

The mushroom is edible, though not as highly regarded as other boletes. It is commonly prepared and eaten in soups, stews or fried dishes. The slime coating, however, may cause indigestion if not removed before eating. It is often sold as a dried mushroom.

==Taxonomy== The species was one of the many first described in 1753 by the "father of taxonomy" Carl Linnaeus, who, in the second volume of his ''Species Plantarum'', gave it the name ''Boletus luteus''.<ref name="Linnaeus 1177"/> The specific epithet is the Latin adjective ''lūtěus'', meaning "yellow".<ref name="Simpson 1979"/> The fungus was reclassified as (and became the type species of) the genus ''Suillus'' by French naturalist Henri François Anne de Roussel in 1796.<ref name="Roussel 1796"/> ''Suillus'' is an ancient term for fungi, and is derived from ''swine''.<ref name="Arora 1986"/> In addition to the British Mycological Society approved name "slippery jack",<ref name="BMS Common names 2014"/> other common names for this bolete include "pine boletus" and "sticky bun"—the latter referring to its resemblance to the pastry.<ref name="Jordan 2015"/>

[[File:Boletus luteus — Flora Batava — Volume v13.jpg|thumb|"''Boletus luteus''", illustrated by Christiaan Sepp]]

German naturalist August Batsch described ''Boletus volvatus'' (the specific epithet derived from the Latin ''volva'', meaning "sheath", "covering" or "womb")<ref name="Simpson 1979"/> alongside ''B. luteus'' in his 1783 work ''Elenchus Fungorum''. Batsch placed both of these species, along with ''B. bovinus'' and the now obsolete names ''Boletus mutabilis'' and ''B. canus'', in a grouping of similar boletes he called "subordo Suilli".<ref name="Batsch 1783"/> ''Boletus volvatus'' is now considered a synonym of ''Suillus luteus''.<ref name="urlFungorum synoynymy: Suillus luteus"/> Several authors have placed the slippery jack in other genera: Finnish mycologist Petter Karsten classified it as ''Cricunopus luteus'' in 1881—the genus ''Cricinopus'' defined by yellow adnate tubes;<ref name="Karsten 1881"/> Lucien Quélet classified it as ''Viscipellis luteus'' in 1886,<ref name="Quélet 1886"/> and ''Ixocomus luteus'' in 1888; and Paul Christoph Hennings placed it in the section ''Cricinopus'' of the genus ''Boletopsis'' in 1900.<ref name="Hennings 1900"/>

In works published before 1987, the slippery jack was written fully as {{nowrap|''Suillus luteus'' (L.:Fr.) Gray}}, as the description by Linnaeus had been name sanctioned in 1821 by the "father of mycology", Swedish naturalist Elias Magnus Fries. The starting date for all the mycota had been set by general agreement as 1 January 1821, the date of Fries's work. Furthermore, as Roussel's description of ''Suillus'' predated this as well, the authority for the genus was assigned to British botanist Samuel Frederick Gray in the first volume of his 1821 work ''A Natural Arrangement of British Plants''.<ref name="Gray 1821"/> The 1987 edition of the International Code of Botanical Nomenclature changed the rules on the starting date and primary work for names of fungi, and names can now be considered valid as far back as 1 May 1753, the date of publication of Linnaeus's work.<ref name="Hawksworth 2001"/> In 1986, a collection of fruit bodies from Sweden was designated as the neotype of ''Suillus luteus''.<ref name="Palm 1986"/>

In their 1964 monograph on North American ''Suillus'' species, Alexander H. Smith and Harry Delbert Thiers classified ''S. luteus'' in the series ''Suilli'' of the section ''Suillus'' in genus ''Suillus''. This group is characterized by the presence of either a ring on the stipe (or stem), a partial veil adhering to the cap margin, or a "false veil" not attached to the stipe but initially covering the tube cavity.<ref name="Smith 1964"/> Species closely related to ''Suillus luteus'' include ''S. pseudobrevipes'' (a sister species), ''S. brevipes'' and ''S. weaverae'' (formerly ''Fuscoboletinus weaverae'').<ref name="Kretzer 1996"/> A genetic study of nucleotide DNA reinforced the species' monophyly and low genetic divergence, with material of ''S. luteus'' from the United Kingdom, Austria, Germany and North America forming a clade, in contrast with some other species, such as ''S. granulatus'', which were shown to be polyphyletic.<ref name="Manian 2001"/>

Chemical analysis of pigments and chromogens showed that ''Suillus'' was more closely related to ''Gomphidius'' and ''Rhizopogon'' than to other boletes, and hence ''Suillus luteus'' and its allies were transferred from the Boletaceae to the newly circumscribed family Suillaceae in 1997.<ref name="Besl 1997"/> Molecular studies have reinforced how distantly related these fungi are from ''Boletus edulis'' and its allies.<ref name="Binder 2006"/>

==Description== {{multiple image|caption_align=center|header_align=center | align = right | total_width = 320 | image1 = Suillus luteus 269479.jpg | width1 = 908 | height1 = 1280 | alt1 = | caption1 = Young fruit bodies have a partial veil that encloses the pores. | image2 = Suillus luteus 113280.jpg | width2 = 2592 | height2 = 1944 | alt2 = | caption2 = In maturity, the partial veil rips away from the cap, remaining on the upper stipe as a well-developed, membranous ring. }}

The cap is chestnut, rusty, olive brown, or dark brown in color with a diameter of {{Convert|4–10|cm|frac=4}}, rarely up to 20 cm.<ref name="Zeitlmayr l976"/> The cap has a distinctive conical shape, later flattening out. It is slimy to the touch, bare, smooth, and glossy even when dry, and the cuticle is easily peeled off. The tiny, circular pores of the tubes are initially yellow but turn olive to dark yellow with maturity. Like the skin of the cap, they can be readily peeled away from the flesh.<ref name="Haas 1969"/>

Tubes comprising the hymenophore on the underside of the cap are {{convert|3–7|mm|1|abbr=on|frac=8}} deep, with an attachment to the stipe ranging from adnate to somewhat decurrent. The pores are tiny, numbering 3 per mm in young specimens and 1–2 per mm in maturity.<ref name="Smith 1964"/> The stipe is {{convert|5–10|cm|abbr=on|frac=4}} tall and {{convert|2–3|cm|1|abbr=on|frac=4}} wide.<ref name="Phillips 2006"/> It is pale yellow and more or less cylindrical but may bear a swollen base. A membranous partial veil initially links the stipe with the edge of the cap. When it ruptures, it forms a membranous, hanging ring.<ref name="Haas 1969"/> The top side of the ring is whitish, while the underside is characteristically dark brown to violet. This species is one of the few members of the genus ''Suillus'' that have such a ring.<ref name="Haas 1969"/> Above the ring, the stipe features glandular dots—minute clumps of pigmented cells. Below the ring, the stipe is dingy white, sometimes streaked with brownish slime.<ref name="Ammirati 1987"/> In humid conditions, the ring has a gelatinous texture.<ref name="Smith 1964"/> The white flesh of the entire fungus does not discolour when damaged, and is soft—particularly in mature specimens.<ref name="Haas 1969"/> It has a "pleasant" taste and lacks any distinctive odour.<ref name="Smith 1964"/>

The spore print is ochre or clay coloured, the elongated elliptical spores measuring 7–10 by 3–3.5 μm.<ref name="Phillips 2006"/> Basidia (spore-producing cells) are four spored, with dimensions of 14–18 by 4–5 μm. Cystidia are present on both the tube faces (pleurocystidia) and edges (cheilocystidia), either scattered or, more rarely, as bundles. They measure 20–35 by 5–7 μm and have a narrow club shape. Clamp connections are not present in the hyphae of ''S. luteus''.<ref name="Smith 1964"/>

===Similar species=== [[File:Körnchen-Röhrling Suillus granulatus 1.jpg|thumb|''Suillus granulatus'', a frequent lookalike]]

Good field characteristics for the species include the slimy brown cap, glandular dots on the upper stipe, and prominent purplish ring. A frequent lookalike is ''S. granulatus'', which is another common, widely distributed and edible species occurring in the same habitat; it is yellow fleshed and exudes latex droplets when young, but most conspicuously bears neither a partial veil nor a ring.<ref name="Min 2014"/> Other than that, ''S. luteus'' is unlikely to be confused with other mushrooms, especially if its preferred habitat under pine trees and the whitish partial veil are considered. In Europe, the related ''S. grevillei'' is found under larch and has a yellow cap, while immature fruit bodies of ''Gomphidius glutinosus'' may look comparable from above but have gills rather than pores underneath.<ref name="Haas 1969"/> In North America, ''Suillus borealis'' and ''S. pseudobrevipes'' also have partial veils, but lack the distinctive ring of ''S. luteus''.<ref name="Arora 1986"/> ''S. cothurnatus'' forms a band-like ring on the stipe that tends to be brownish rather than purplish.<ref name="Kuo 2007"/>

In some specimens of ''S. luteus'', the partial veil separates from the stipe (rather than the cap margin), leaving cottony patches of veil hanging from the cap margin. In this state, fruit bodies can be confused with those of ''S. albidipes''. Unlike ''S. luteus'', however, ''S. albidipes'' does not have glandular dots on its stipe.<ref name="Smith 1964"/>

==Distribution and habitat== thumb|left|Growing under host tree, Finland

''Suillus luteus'' can be found all over the Northern Hemisphere. Native to Eurasia, it is widespread across the British Isles.<ref name="Nilper 1977"/> To the east it has been recorded from Pakistan, where it was found along canals in Dashkin in the district of Astore,<ref name="Sarwar 2007"/> and as far east as South Korea.<ref name="Min 2014"/> It has also been widely introduced elsewhere by way of pine plantations around the globe. It is very commonly found in Monterey pine (''Pinus radiata'') plantations, despite the tree being native to California and hence not in the fungus' native range.<ref name="Vellinga 2009"/> In North America it is found in the northeastern, northwestern, and southwestern United States.<ref name="Arora 1986"/> According to Ernst Both,<ref name="Both 1998"/> it was Charles Horton Peck who first suggested in 1887 that the fungus was introduced to the state of New York on ''Pinus sylvestris''.<ref name="Peck 1887"/> DNA studies show that the North American populations differ little genetically from European populations, supporting the idea that the fungus arrived to North America relatively recently as a result of human activity.<ref name="Kuo 2007"/> ''Suillus luteus'' is found in coastal and mountainous pine forests and exhibits a tolerance of the northern latitudes. Southern Hemisphere locales where the slippery jack grow with plantation pines include South America, Africa, Australia, and New Zealand.<ref name="Roberts 2011"/> In southwestern Australia, the bolete is limited to areas of greater than 1000 mm (40 in) annual rainfall.<ref name="Dunstan 1998"/> It has been recorded as far north as the Darling Downs and southern Queensland,<ref name="QMS"/> and occasionally in Tasmania.<ref name="Ratkowsky 2005"/> The fungus fruits in spring, summer and fairly prolifically in autumn, following periods of wet weather. Mushrooms can appear in large troops or fairy rings.<ref name="Haas 1969"/>

In Ecuador, ''Pinus radiata'' plantations were planted extensively around Cotopaxi National Park, and ''Suillus luteus'' boletes appear in abundance year-round. A 1985 field study estimated production to be 3000–6000 mushrooms per hectare—up to {{convert|1000|kg}} (dry weight) of mushrooms hectare per year. This continuous production contrasts with the bolete's seasonal appearance elsewhere. The fungus is not found in adjacent areas of native vegetation.<ref name="Hedger 1986"/> The fruiting is so bountiful that the harvest of slippery jacks has become the main reason that pine plantations are established or maintained in parts of Ecuador.<ref name="Farley 2004"/>

In southern Brazil, it has been recorded in plantations of slash pine (''P. elliottii'') in the municipalities of Pelotas, Nova Petrópolis and Canela in Rio Grande do Sul, and Colombo in Paraná.<ref name="Sulzbacher 2013"/> It is particularly common in plantations in Patagonia.<ref name="Barroetavena 2005"/> ''Suillus luteus'' is the commonest bolete encountered in the Falkland Islands, where it is found in windbreaks and gardens.<ref name="Watling 2012"/>

In South Africa, ''Suillus luteus'' has been occasionally recorded under pines in Bloemfontein, Johannesburg and Royal Natal National Park.<ref name="Van der Westhuizen"/>

==Ecology== thumb|Depicted on a 1964 Soviet Union postage stamp

''Suillus luteus'' is a pioneer species that typically establishes itself in the early stages of forest succession.<ref name="Muller 2004"/> The fungus forms mycorrhizal associations with various species of pine, including Scots pine (''P. sylvestris''), black pine (''P. nigra''),<ref name="Loizides 2011"/> and Macedonian pine (''P. peuce'') in Europe,<ref name="Karadelev 2007"/><ref name="Yagiz 2006"/> and red pine (''P. resinosa'') and white pine (''P. strobus'') in North America.<ref name="mushroomexpert"/> An ''in vitro'' experiment demonstrated that the species could form an ectomycorrhizal association with Aleppo pine (''P. halepensis''), a key species used in reforestation in the Mediterranean.<ref name="Torres 1994"/> A study of the ectomycorrhizal fungi associated with a lodgepole pine (''P. contorta'') invasion front near Coyhaique, Chile, showed that many invasive trees were supported by ''S. luteus'' as the sole mycorrhizal partner.<ref name="Hayward 2015"/>

The ectomycorrhizae formed between the fungus and host plant can be influenced by soil microorganisms present in the mycorrhizosphere. For example, soil bacteria from the genera ''Paenibacillus'' and ''Burkholderia'' alter the branching structure of the root, whereas ''Bacillus'' species increase root growth and mycorrhizal colonization.<ref name="Churchland 2014"/> The fungus does not require a specific soil but seems to prefer acidic and nutrient-deficient soil.<ref name="Haas 1969"/> ''Suillus luteus'' produces hydroxamic acid-based siderophores, which are compounds that can chelate iron and extract it from the soil in nutrient-poor conditions.<ref name="Haselwandter 2011"/> Ignacio Chapela and colleagues analysed the carbon uptake of ''S. luteus'' in Ecuador, concluding pine plantations accompanied by ''S. luteus'' deplete carbon stored in the soil and raising concerns that these might not be a remedy for rising carbon dioxide levels in the atmosphere.<ref name="Chapela 2001"/>

The fungus has been shown to provide a protective effect against heavy metal toxicity when associated with the host ''Pinus sylvestris'', preventing copper accumulation in the needles, and protecting seedlings against cadmium toxicity.<ref name="Mackova 2006"/><ref name="Krznaric 2009"/> Owing to its frequent rate of sexual reproduction and the resulting extensive gene flow within populations, the fungus can rapidly evolve a trait to tolerate otherwise toxic levels of heavy metals in the environment.<ref name="Muller 2007"/> The genetic basis of this adaptation—intriguing to researchers investigating the bioremediation potential of metal-adapted plants and their fungal associates<ref name="JGI"/>—are contained in the genome sequence of ''S. luteus'', published in 2015.<ref name="Kohler 2015"/>

''Suillus luteus'' fruit bodies are sometimes infested with larvae, though not nearly as often as ''S. granulatus'' or ''B. edulis''.<ref name="Haas 1969"/> Damage from maggots is much more common in warmer months, and rare late in the season with cooler weather.<ref name="Nilper 1977"/> In a Finnish study, researchers found that 70–95% of fruit bodies collected from typical forest habitats were infested with larvae; the most common species were the flies ''Mycetophila fungorum'', ''Pegomya deprimata'', and ''Pegohylemyia silvatica''.<ref name="Hackmann 1979"/> In contrast, other studies have shown that fruit bodies collected from pine plantations are relatively free of larvae.<ref name="Zambonelli 2013"/> The fungus produces microscopic crystals of oxalic acid at the surface of its hyphae, a feature that is thought to help deter grazing by the springtail species ''Folsomia candida''.<ref name="Böllmann 2009"/>

==Edibility== thumb|Slippery jack mushrooms collected in Ukraine ''Suillus luteus'' is an edible mushroom, but the slime/pileipellis must be removed.<ref>{{cite book |last=Phillips |first=Roger |title=Mushrooms and Other Fungi of North America |year=2010 |publisher=Firefly Books |location=Buffalo, NY |isbn=978-1-55407-651-2 |page=288}}</ref><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|page=364|author-link=Orson K. Miller Jr.}}</ref> Although some authors regard it as one of low quality,<ref name="Arora 1986" /><ref name="Jordan 1995" /> and generally inferior to co-occurring species such as ''Boletus pinophilus'',<ref name="Lamaison 2005" /> the species is considered a delicacy in Slavic cultures (known as ''maslyata'' in Russian or ''maslyuky'' in Ukrainian or ''maślaki'' in Polish, deriving from words meaning "buttery"). It was highly regarded in Calabria, even more than ''Boletus edulis'', until the 1940s when increased interest in the latter species eclipsed the former.<ref name="Sitta 2008" /> Mushrooms conforming to ''Suillus luteus'' are exported from Chile to Italy,<ref name="Sitta 2008" /> and, since the 1970s, the United States.<ref name="Dahlberg 1999" /> As of 2002, harvesters in Chile were paid on average US$0.5 per kilogram of fruit bodies.<ref name="De Roman 2010" />

In Burundi, ''Suillus luteus'' mushrooms are sold to Europeans as ''cepes'' in Bujumbura but not generally eaten by the Barundi.<ref name="Buyck 1995" /> Based on samples collected from Chile, the boletes contain (as a percentage of dry weight) 20% protein, 57% carbohydrates, 6% fat, and 6% ash.<ref name="Verde 2012" /> ''Pinus radiata'' plantations in southeastern Australia have become tourist attractions as people flock to them in autumn to pick slippery jacks and saffron milk-caps (''Lactarius deliciosus'');<ref name="Oberon Australia" /> Belanglo State Forest in particular has attracted large numbers of Polish foragers.<ref name="Kwiatkowski 2004" />

Slippery jacks do not keep for long after picking,<ref name="Haas 1969"/> Zeitmar considers them unsuitable for drying,<ref name="Zeitlmayr l976"/> as their water content is too high. They are suited for frying,<ref name="Zeitlmayr l976"/> or cooking in stews and soups, either alone or with other mushroom species.<ref name="Carluccio 2003"/> Puréeing the mushroom is not recommended, however: "We once made the mistake of running it through a blender to make a soup. The result was a substance recommending itself for use when hanging wallpaper."<ref name="Kuo 2007"/> ''S. luteus'' and other ''Suillus'' species may cause allergic reactions in some people<ref name="Bruhn 1991"/> or digestive problems that appear to result from consuming the slimy skin.<ref name="Kuo 2007"/> The fungus is better cooked before eating, and some authors recommend discarding the glutinous cuticle and tubes before cooking.<ref name="Ammirati 1987"/><ref name="McKnight 1998"/> Moreover, the skin can spoil other fungi with which slippery jacks are collected.<ref name="Zeitlmayr l976"/>

Inexpensive powdered ''S. luteus'' fruit bodies are sometimes added to the more expensive ''B. edulis'' mushroom soup powder, a fraudulent practice that is difficult to detect by microscope because the tissues are no longer intact.<ref name="Murat 2008"/> This adulteration can be determined chemically, however, by testing for increased levels of the sugar alcohols arabitol and mannitol.<ref name="Sforza 2013"/> The practice can also be determined with a DNA-based method that is sensitive enough to detect the addition of 1–2% of ''S. luteus'' to ''B. edulis'' powder.<ref name="Moor 2002"/>

==See also== {{Portal|Fungi}} * List of North American boletes

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}}

==External links== {{Commons category|Suillus luteus}} {{Wikispecies|Suillus luteus}}

* ''[http://www.mykoweb.com/boletes/species/Suillus_luteus.html Suillus luteus]'' at MykoWeb

luteus Category:Edible fungi Category:Fungi of Europe Category:Fungi described in 1753 Category:Fungi of Africa Category:Fungi of Asia Category:Fungi of Central America Category:Fungi of New Zealand Category:Fungi of the United States Category:Fungi found in fairy rings Category:Fungal taxa named by Carl Linnaeus Category:Fungus species