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Paxillus involutus
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<h2 id="taxonomy-and-naming">Taxonomy and naming</h2> <p>The brown roll-rim was described by French mycologist <a href="/facts/Jean_Baptiste_Fran%25C3%25A7ois_Pierre_Bulliard/lTITdKN8">Pierre Bulliard</a> in 1785 as <i>Agaricus contiguus</i>,<a class="footnote-ref" id="fnref:1" href="#fn:1"><sup>1</sup></a> although the 1786 combination <i>Agaricus involutus</i> of <a href="/facts/August_Batsch/xPsRRmkD">August Batsch</a><a class="footnote-ref" id="fnref:2" href="#fn:2"><sup>2</sup></a> is taken as the first <a href="/facts/Valid_name_(botany)/fCJj0rfN">valid description</a>.<a class="footnote-ref" id="fnref:3" href="#fn:3"><sup>3</sup></a> <a href="/facts/James_Bolton/z575zVCz">James Bolton</a> published a description of what he called <i>Agaricus adscendibus</i> in 1788;<a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a> the <a href="/facts/Taxonomy_(biology)/Qkg8wuyw">taxonomical</a> authority <a href="/facts/Index_Fungorum/rqBKYxlD">Index Fungorum</a> considers this to be <a href="/facts/Synonym_(biology)/BHLdox7E">synonymous</a> with <i>P. involutus</i>.<a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a> Additional synonyms include <i>Omphalia involuta</i> described by <a href="/facts/Samuel_Frederick_Gray/5brmJ0UH">Samuel Frederick Gray</a> in 1821,<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> and <i>Rhymovis involuta</i>, published by <a href="/facts/Gottlob_Ludwig_Rabenhorst/SXTMMNwF">Gottlob Ludwig Rabenhorst</a> in 1844.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> The species gained its current <a href="/facts/Binomial_nomenclature/jgBqAhPv">binomial name</a> in 1838 when the 'father of mycology', Swedish naturalist <a href="/facts/Elias_Magnus_Fries/IspFbsTk">Elias Magnus Fries</a> erected the genus <i><a href="/facts/Paxillus/1aFh2Uxd">Paxillus</a></i>, and set it as the <a href="/facts/Type_species/kL03T6V4">type species</a>.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> The starting date of fungal <a href="/facts/Taxonomy_(biology)/Qkg8wuyw">taxonomy</a> had been set as January 1, 1821, to coincide with the date of Fries' works, which meant that names coined earlier than this date required <a href="/facts/Sanctioned_name/5QHuxsAV">sanction</a> by Fries (indicated in the name by a colon) to be considered <a href="/facts/Correct_name_(botany)/qT06Fuem">valid</a>. It was thus written <i>Paxillus involutus</i> (Batsch:Fr.) Fr. A 1987 revision of the <a href="/facts/International_Code_of_Botanical_Nomenclature/EbaUF9P5">International Code of Botanical Nomenclature</a> set the starting date at May 1, 1753, the date of publication of <a href="/facts/Carl_Linnaeus/gy5HWkHS">Linnaeus</a>' seminal work, the <i><a href="/facts/Species_Plantarum/SKwpspkt">Species Plantarum</a></i>.<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> Hence the name no longer requires the ratification of Fries' authority. </p><p>The genus was later placed in a new family, <a href="/facts/Paxillaceae/Sz2NxP1P">Paxillaceae</a>, by French mycologist <a href="/facts/Ren%25C3%25A9_Maire/omNoekpb">René Maire</a> who held it to be related to both <a href="/facts/Agaric/pQqJmCeK">agarics</a> and boletes.<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> Although it has gills rather than pores, it has long been recognised as belonging to the pored mushrooms of the order <a href="/facts/Boletales/4gNcsGES">Boletales</a> rather than the traditional agarics.<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> The generic name is derived from the <a href="/facts/Latin/pM3Kge85">Latin</a> for 'peg' or 'plug', and the <a href="/facts/Specific_name_(botany)/31ckrsXv">specific epithet</a> <i>involutus</i>, 'inrolled', refers to the cap margin.<a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> <a href="/facts/Common_name/3KNNezEU">Common names</a> include the naked brimcap,<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a> poison paxillus,<a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a> inrolled pax,<a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a> poison pax, common roll-rim, brown roll-rim,<a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a> and brown chanterelle.<a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a> Gray called it the "involved navel-stool" in his 1821 compendium of British flora.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> </p><p>Studies of the ecology and genetics of <i>Paxillus involutus</i> indicate that it may form a <a href="/facts/Species_complex/u6qEMdRQ">complex</a> of multiple similar-looking species.<a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a><a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> In a field study near <a href="/facts/Uppsala/X3HzSeYN">Uppsala</a>, Sweden, conducted from 1981 to 1983, mycologist Nils Fries found that there were three populations of <i>P. involutus</i> unable to breed with each other. One was found under conifers and mixed woodlands, while the other two were found in parklands, associated with nearby birch trees. He found that the first group tended to produce single isolated fruit bodies which had a thinner stipe and cap which was less inrolled at the margins, while the fruit bodies of the other two populations tended to appear in groups, and have thicker stipes, and caps with more inrolled and sometimes undulating margins. There were only general tendencies and he was unable to detect any consistent macroscopic or microscopic features that firmly differentiate them.<a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> A <a href="/facts/Molecular_phylogenetics/FysplHP5">molecular</a> study comparing the <a href="/facts/DNA_sequence/hsPsNCQW">DNA sequences</a> of specimens of <i>Paxillus involutus</i> collected from various habitats in <a href="/facts/Bavaria/CLbr6NIG">Bavaria</a> found that those collected from parks and gardens showed a close relationship with the North American species <i><a href="/facts/Paxillus_vernalis/NKMj8v7n">P. vernalis</a></i>, while those from forests were allied with <i><a href="/facts/Paxillus_filamentosus/DcNxDHAJ">P. filamentosus</a></i>. The authors suggested the park populations may have been introduced from North America.<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a> A multi-gene analysis of European isolates showed that <i>P. involutus</i> <i><a href="/facts/Sensu_lato/ho2gtX2C">sensu lato</a></i> (in the loose sense) could be separated into four distinct, genetically isolated lineages corresponding to <i><a href="/facts/Paxillus_obscurosporus/Y8x0mAQs">P. obscurosporus</a></i>, <i>P. involutus</i> <i>sensu stricto</i> (in the strict sense), <i>P. validus</i>, and a fourth species that has not yet been identified.<a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a> Changes in host range have occurred frequently and independently among strains within this species complex.<a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a> </p> <h2 id="description">Description</h2> <p>Resembling a brown wooden <a href="/facts/Spinning_top/u50YUBP6">top</a>, the <a href="/facts/Epigeous/K7oCphYp">epigeous</a> (aboveground) <a href="/facts/Basidiocarp/xgaQV94l">fruit body</a> may be up to 6 cm (2+3⁄8 in) high.<a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a> The <a href="/facts/Pileus_(mycology)/6d91unJ1">cap</a>, initially convex then more funnel-shaped (infundibuliform) with a depressed centre and rolled rim (hence the common name), may be reddish-, yellowish- or olive-brown in colour and typically 4–12 cm (1+5⁄8–4+3⁄4 in) wide;<a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a> the cap diameter does not get larger than 15 cm (5+7⁄8 in).<a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a> The cap surface is initially downy and later smooth, becoming sticky when wet. The cap and cap margin initially serve to protect the <a href="/facts/Lamella_(mycology)/V18VY7tJ">gills</a> of young fruit bodies: this is termed pilangiocarpic development.<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a> The narrow brownish yellow gills are decurrent and forked, and can be peeled easily from the flesh (as is the case with the pores of boletes). Gills further down toward the stipe become more irregular and <a href="/facts/Anastomosis/fScK17o8">anastomose</a>, and can even resemble the pores of bolete-type fungi. The fungus darkens when bruised and older specimens may have darkish patches. The juicy yellowish <a href="/facts/Trama_(mycology)/C9QvGj7h">flesh</a> has a mild to faintly sour or sharp odor and taste, and has been described as well-flavored upon cooking.<a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a><a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a> Of similar colour to the cap, the short <a href="/facts/Stipe_(mycology)/032SVJIw">stipe</a> measures some 3–6 cm tall and 1–3 wide,<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a> can be crooked, and tapers toward the base.<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a> </p> <p>The <a href="/facts/Spore_print/WFAfnvuK">spore print</a> is brown, and the dimensions of the ellipsoid (oval-shaped) spores are 7.5–9 by 5–6 <a href="/facts/Micrometre/7eDFxD2L">μm</a>. The <a href="/facts/Hymenium/ScjqHmWY">hymenium</a> has <a href="/facts/Cystidia/qReFlXTn">cystidia</a> both on the gill edge and face (cheilo- and pleurocystidia respectively), which are slender and filament-like, typically measuring 40–65 by 8–10.5 μm.<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> </p> <h3>Similar species</h3> <p>The brownish colour and funnel-like shape of <i>P. involutus</i> can lead to its confusion with several species of <i><a href="/facts/Lactarius_(fungus)/C9S5gGqU">Lactarius</a></i>, many of which have some degree of toxicity themselves.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> The lack of a milky exudate distinguishes it from any milk cap.<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a> One of the more similar is <i><a href="/facts/Lactarius_turpis/SsrAMAjL">L. turpis</a></i>, which presents a darker olive colouration.<a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> The related North American <i>Paxillus vernalis</i> has a darker spore print, thicker stipe and is found under aspen,<a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a> whereas the closer relative <i>P. filamentosus</i> is more similar in appearance to <i>P. involutus</i>. A rare species that grows only in association with <a href="/facts/Alder/KbD9YG8w">alder</a>, <i>P. filamentosus</i> can be distinguished from it by the pressed-down scales on the cap surface that point towards the cap margin, a light yellow flesh that bruises only slightly brown, and deep yellow-ochre gills that do not change colour upon injury<a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> </p><p>The most similar species are two once thought to be part of <i>P. involutus</i> in Europe. <i><a href="/facts/Paxillus_obscurisporus/Y8x0mAQs">Paxillus obscurisporus</a></i> (originally <i>obscurosporus</i>) has larger fruit bodies than <i>P. involutus</i>, with caps up to 40 cm (16 in) wide whose margins tend to unroll and flatten with age, and a layer of cream-coloured mycelia covering the base of its tapered stipe. <i>P. validus</i>, also known only from Europe, has caps up to 20 cm (7+7⁄8 in) wide with a stipe that is more or less equal in width throughout its length. Found under broadleaved trees in parks, it can be reliably distinguished from <i>P. involutus</i> (and other <i>Paxillus</i> species) by the presence of crystals up to 2.5 μm long in the <a href="/facts/Mycelial_cord/wyRdw1TT">rhizomorphs</a>, as the crystals found in rhizomorphs of other <i>Paxillus</i> species do not exceed 0.5 μm long.<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> </p><p>Other similar species include <i><a href="/facts/Phylloporus_arenicola/apBokXu4">Phylloporus arenicola</a></i>, <i><a href="/facts/Tapinella_atrotomentosa/TQcm2JWH">Tapinella atrotomentosa</a></i>, and <i><a href="/facts/Tapinella_panuoides/8bLymzbk">Tapinella panuoides</a></i>.<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> </p> <h2 id="ecology-distribution-and-habitat">Ecology, distribution and habitat</h2> <p><i>Paxillus involutus</i> forms <a href="/facts/Ectomycorrhiza/HAgj8brf">ectomycorrhizal</a> relationships with a number of <a href="/facts/Pinophyta/vDUgqxCx">coniferous</a> and <a href="/facts/Deciduous/7jQHVtdd">deciduous</a> tree species. Because the fungus has somewhat unspecialized nutrient requirements and a relatively broad <a href="/facts/Host_(biology)/L6mop0kh">host</a> specificity, it has been frequently used in research and seedling inoculation programs.<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a> There is evidence of the benefit to trees of this arrangement: in one experiment where <i>P. involutus</i> was cultivated on the root exudate of red pine (<i><a href="/facts/Pinus_resinosa/uF7q3TT9">Pinus resinosa</a></i>), the root showed markedly increased resistance to pathogenic strains of the ubiquitous soil fungus <i><a href="/facts/Fusarium_oxysporum/meRut0kE">Fusarium oxysporum</a></i>.<a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a> Seedlings inoculated with <i>P. involutus</i> also showed increased resistance to <i>Fusarium</i>.<a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> Thus <i>P. involutus</i> may be producing antifungal compounds which protect the host plants from root rot.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> <i>Paxillus involutus</i> also decreases the uptake of certain toxic elements, acting as a buffer against <a href="/facts/Heavy_metal_(chemistry)/Cblmy8qI">heavy metal</a> toxicity in the host plant. For example, the fungus decreased the toxicity of <a href="/facts/Cadmium/wXkV2eIX">cadmium</a> and <a href="/facts/Zinc/80b1qgk3">zinc</a> to Scots pine (<i><a href="/facts/Pinus_sylvestris/Jd9DJoju">Pinus sylvestris</a></i>) seedlings: even though cadmium itself inhibits ectomycorrhiza formation in seedlings, colonization with <i>P. involutus</i> decreases cadmium and zinc transport to the plant shoots and alters the ratio of zinc transported to the roots and shoots, causing more cadmium to be retained in the roots of the seedlings rather than distributed through its entire metabolism.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> Evidence suggests that the mechanism for this detoxification involves the cadmium binding to the fungal <a href="/facts/Cell_wall/DYXJVrnw">cell walls</a>, as well as accumulating in the <a href="/facts/Vacuole/9QjBVb7j">vacuolar compartments</a>.<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> Further, ectomycorrhizal hyphae exposed to <a href="/facts/Copper/I8PEE8oX">copper</a><a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> or cadmium drastically increase production of a <a href="/facts/Metallothionein/4ntygR3A">metallothionein</a>—a low <a href="/facts/Molecular_weight/FWueGMed">molecular weight</a> protein that binds metals.<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a><a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> </p><p>The presence of <i>Paxillus involutus</i> is related to much reduced numbers of bacteria associated with the roots of <i>Pinus sylvestris</i>. Instead bacteria are found on the external mycelium.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> The types of bacteria change as well; a Finnish study published in 1997 found that bacterial communities under <i>P. sylvestris</i> without mycorrhizae metabolised <a href="/facts/Organic_acid/2Pfb26Lp">organic</a> and <a href="/facts/Amino_acid/WDxYwBny">amino acids</a>, while communities among <i>P. involutus</i> metabolised the sugar <a href="/facts/Fructose/pEqWJC2N">fructose</a>.<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a> <i>Paxillus involutus</i> benefits from the presence of some species of bacteria in the soil it grows in. As the fungus grows it excretes <a href="/facts/Polyphenol/H6Q7QIQH">polyphenols</a>, waste products that are toxic to itself and impede its growth, but these compounds are metabolised by some bacteria, resulting in increased fungal growth. Bacteria also produce certain compounds such as <a href="/facts/Citric_acid/EgKGQBdl">citric</a> and <a href="/facts/Malic_acid/mSbExe77">malic acid</a>, which stimulate <i>P. involutus</i>.<a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> </p> <p>Highly abundant,<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> the brown roll-rim is found across the Northern Hemisphere, Europe and Asia, with records from India,<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> China,<a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a> Japan, Iran,<a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> and Turkey's eastern <a href="/facts/Anatolia/vMkrLrPr">Anatolia</a>.<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> It is equally widely distributed across northern North America,<a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a> extending north to <a href="/facts/Alaska/LlDNr2uR">Alaska</a>, where it has been collected from <a href="/facts/Tundra/BqC0AQ7u">tundra</a> near <a href="/facts/Coldfoot%2c_Alaska/Na5e9wXG">Coldfoot</a> in the interior of the state.<a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></a> In southwestern Greenland, <i>P. involutus</i> has been recorded under the birch species <i><a href="/facts/Betula_nana/DUk7PQ8o">Betula nana</a></i>, <i><a href="/facts/Betula_pubescens/HBLtt76Q">B. pubescens</a></i> and <i><a href="/facts/Betula_glandulosa/o5NaD6hU">B. glandulosa</a></i>.<a class="footnote-ref" id="fnref:60" href="#fn:60"><sup>60</sup></a> The mushroom is more common in coniferous woods in Europe, but is also closely associated with birch (<i><a href="/facts/Betula_pendula/xEth6vrV">Betula pendula</a></i>). Within woodland, it prefers wet places or boggy ground, and avoids <a href="/facts/Calcareous/sGpRISvY">calcareous</a> (chalky) soils. It has been noted to grow alongside <i><a href="/facts/Boletus_badius/xCQEovX7">Boletus badius</a></i> in Europe,<a class="footnote-ref" id="fnref:61" href="#fn:61"><sup>61</sup></a> and <i><a href="/facts/Leccinum_scabrum/dWoVLxay">Leccinum scabrum</a></i> and <i><a href="/facts/Lactarius_plumbeus/SsrAMAjL">Lactarius plumbeus</a></i> in the <a href="/facts/Pacific_Northwest/EjJhETXc">Pacific Northwest</a> region of North America.<a class="footnote-ref" id="fnref:62" href="#fn:62"><sup>62</sup></a> There it is found in both deciduous and coniferous woodland, commonly under plantings of white birch (<i><a href="/facts/Betula_papyrifera/MUAPFrr6">Betula papyrifera</a></i>) in urban areas.<a class="footnote-ref" id="fnref:63" href="#fn:63"><sup>63</sup></a> It is one of a small number of fungal species which thrive in <i><a href="/facts/Pinus_radiata/RaQ5nXhy">Pinus radiata</a></i> plantations planted outside their natural range.<a class="footnote-ref" id="fnref:64" href="#fn:64"><sup>64</sup></a> A study of polluted Scots pine forest around <a href="/facts/Oulu/6d8fo60J">Oulu</a> in northern Finland found that <i>P. involutus</i> became more abundant in more polluted areas while other species declined. Emissions from pulp mills, fertiliser, heating and traffic were responsible for the pollution, which was measured by sulfur levels in the pine needles.<a class="footnote-ref" id="fnref:65" href="#fn:65"><sup>65</sup></a> </p><p><i>Paxillus involutus</i> can be found growing on <a href="/facts/Lawn/8j5Yf84C">lawns</a> and old <a href="/facts/Meadow/gXJr3U3P">meadows</a> throughout its distribution. Fruit bodies are generally terrestrial, though they may be found on woody material around tree stumps.<a class="footnote-ref" id="fnref:66" href="#fn:66"><sup>66</sup></a> They generally appear in autumn and late summer.<a class="footnote-ref" id="fnref:67" href="#fn:67"><sup>67</sup></a> In California, <a href="/facts/David_Arora/Iwu9RXPl">David Arora</a> discerned a larger form associated with oak and pine which appears in late autumn and winter, as well as the typical form that is associated with birch plantings and appears in autumn.<a class="footnote-ref" id="fnref:68" href="#fn:68"><sup>68</sup></a> Several species of flies and beetles have been recorded using the fruit bodies to rear their young.<a class="footnote-ref" id="fnref:69" href="#fn:69"><sup>69</sup></a> The mushroom can be infected by <i><a href="/facts/Hypomyces_chrysospermus/MeBt16bY">Hypomyces chrysospermus</a></i>, or bolete eater, a <a href="/facts/Mold_(fungus)/7nHqptM7">mould</a> species that parasitises <a href="/facts/Boletales/4gNcsGES">Boletales</a> members.<a class="footnote-ref" id="fnref:70" href="#fn:70"><sup>70</sup></a> Infection results in the appearance of a whitish powder that first manifests on the pores, then spreads over the surface of the mushroom, becoming golden yellow to reddish-brown in maturity.<a class="footnote-ref" id="fnref:71" href="#fn:71"><sup>71</sup></a> </p><p>Australian mycologist <a href="/facts/John_Burton_Cleland/5fNDhfhM">John Burton Cleland</a> noted it occurring under <a href="/facts/Larch/BDy77MTP">larch</a> (<i>Larix</i>), oak, pine, birch and other <a href="/facts/Introduced_species/s0aHefyp">introduced</a> trees in <a href="/facts/South_Australia/R7UdxV97">South Australia</a> in 1934,<a class="footnote-ref" id="fnref:72" href="#fn:72"><sup>72</sup></a> and it has subsequently been recorded in <a href="/facts/New_South_Wales/aW7eJXSS">New South Wales</a>, <a href="/facts/Victoria_(Australia)/76P2nqtz">Victoria</a><a class="footnote-ref" id="fnref:73" href="#fn:73"><sup>73</sup></a> (where it was found near <i>Betula</i> and <i>Populus</i>)<a class="footnote-ref" id="fnref:74" href="#fn:74"><sup>74</sup></a> and <a href="/facts/Western_Australia/1ouCbwNh">Western Australia</a>. It has been recorded under introduced <a href="/facts/Birch/XG0bl0GF">birch</a> (<i>Betula</i>) and <a href="/facts/Hazel/K0gposzR">hazel</a> (<i>Corylus</i>) in New Zealand.<a class="footnote-ref" id="fnref:75" href="#fn:75"><sup>75</sup></a> Mycologist <a href="/facts/Rolf_Singer/pKcuccv4">Rolf Singer</a> reported a similar situation in South America, with the species recorded under introduced trees in <a href="/facts/Chile/uEyTfEcx">Chile</a>. It is likely to have been transported to those countries in the soil of imported European trees.<a class="footnote-ref" id="fnref:76" href="#fn:76"><sup>76</sup></a> </p> <h2 id="toxicity">Toxicity</h2> <p><i>Paxillus involutus</i> was widely eaten in <a href="/facts/Central_Europe/VdxzSbJz">Central</a> and <a href="/facts/Eastern_Europe/f5CDGzNA">Eastern Europe</a> until World War II, although English guidebooks did not recommend it.<a class="footnote-ref" id="fnref:77" href="#fn:77"><sup>77</sup></a><a class="footnote-ref" id="fnref:78" href="#fn:78"><sup>78</sup></a> In Poland, the mushroom was often eaten after pickling or salting.<a class="footnote-ref" id="fnref:79" href="#fn:79"><sup>79</sup></a> It was known to be a gastrointestinal irritant when ingested raw but had been presumed edible after cooking.<a class="footnote-ref" id="fnref:80" href="#fn:80"><sup>80</sup></a> Questions were first raised about its toxicity after German mycologist <a href="/facts/Julius_Sch%25C3%25A4ffer/QhDpBACQ">Julius Schäffer</a> died after eating it in October 1944. About an hour after he and his wife ate a meal prepared with the mushrooms, Schäffer developed vomiting, diarrhea, and fever. His condition worsened to the point where he was admitted to hospital the following day and developed <a href="/facts/Kidney_failure/gsqb5nos">kidney failure</a>, perishing after 17 days.<a class="footnote-ref" id="fnref:81" href="#fn:81"><sup>81</sup></a><a class="footnote-ref" id="fnref:82" href="#fn:82"><sup>82</sup></a> </p><p>In the mid-1980s, Swiss physician René Flammer discovered an <a href="/facts/Antigen/9JlAernG">antigen</a> within the mushroom that stimulates an <a href="/facts/Autoimmune/5cASmcfI">autoimmune</a> reaction causing the body's immune cells to consider its own <a href="/facts/Red_blood_cells/Kggpq8Jh">red blood cells</a> as foreign and attack them. Despite this, it was not until 1990 that guidebooks firmly warned against eating <i>P. involutus</i>, and one Italian guidebook recommended it as late as 1998.<a class="footnote-ref" id="fnref:83" href="#fn:83"><sup>83</sup></a> The relatively rare immunohemolytic syndrome occurs following the repeated ingestion of <i>Paxillus</i> mushrooms.<a class="footnote-ref" id="fnref:84" href="#fn:84"><sup>84</sup></a> Most commonly it arises when the person has ingested the mushroom for a long period of time, sometimes for many years, and has shown mild <a href="/facts/Gastrointestinal_symptom/S4UzQRFA">gastrointestinal symptoms</a> on previous occasions.<a class="footnote-ref" id="fnref:85" href="#fn:85"><sup>85</sup></a> The <i>Paxillus</i> syndrome is better classed as a hypersensitivity reaction than a toxicological reaction as it is caused not by a genuinely poisonous substance but by the antigen in the mushroom. The antigen is still of unknown structure but it stimulates the formation of <a href="/facts/Immunoglobulin_G/cKzc83tk">IgG</a> <a href="/facts/Antibodies/MX8pdTdP">antibodies</a> in the <a href="/facts/Blood_serum/a22tvug9">blood serum</a>. In the course of subsequent meals, <a href="/facts/Immune_complex/s4kKT3lc">antigen-antibody complexes</a> are formed; these complexes attach to the surface of blood cells and eventually lead to their <a href="/facts/Lysis/th8pXeEn">breakdown</a>.<a class="footnote-ref" id="fnref:86" href="#fn:86"><sup>86</sup></a> </p><p>Poisoning symptoms are rapid in onset, consisting initially of vomiting, diarrhea, abdominal pain, and associated <a href="/facts/Hypovolemia/MmP8TsPw">decreased blood volume</a>.<a class="footnote-ref" id="fnref:87" href="#fn:87"><sup>87</sup></a> Shortly after these initial symptoms appear, <a href="/facts/Hemolysis/1WpVYHkP">hemolysis</a> develops, resulting in <a href="/facts/Oliguria/T7t2onvx">reduced urine output</a>, <a href="/facts/Hemoglobinuria/C2v7r5NJ">hemoglobin in the urine</a> or outright <a href="/facts/Anuria/aYLfTfb7">absence of urine formation</a>, and <a href="/facts/Anemia/AgEPPmg8">anemia</a>. Medical laboratory tests consist of testing for the presence of increasing <a href="/facts/Bilirubin/HCHyCjkt">bilirubin</a> and free hemoglobin, and falling <a href="/facts/Haptoglobin/FvX19ch9">haptoglobins</a>. Hemolysis may lead to numerous complications including acute kidney injury, <a href="/facts/Shock_(circulatory)/aIKGDS0e">shock</a>, acute <a href="/facts/Respiratory_failure/R3GcdkJ1">respiratory failure</a>, and <a href="/facts/Disseminated_intravascular_coagulation/CuWaSKQW">disseminated intravascular coagulation</a>.<a class="footnote-ref" id="fnref:88" href="#fn:88"><sup>88</sup></a><a class="footnote-ref" id="fnref:89" href="#fn:89"><sup>89</sup></a><a class="footnote-ref" id="fnref:90" href="#fn:90"><sup>90</sup></a> These complications can cause significant <a href="/facts/Disease/55eeQlGL">morbidity</a> with fatalities having been reported.<a class="footnote-ref" id="fnref:91" href="#fn:91"><sup>91</sup></a> </p><p>There is no antidote for poisoning, only <a href="/facts/Supportive_treatment/zQlnUC9x">supportive treatment</a> consisting of monitoring <a href="/facts/Complete_blood_count/NF9lNJL3">complete blood count</a>, renal function, blood pressure, and fluid and electrolyte balance<a class="footnote-ref" id="fnref:92" href="#fn:92"><sup>92</sup></a> and correcting abnormalities. The use of <a href="/facts/Corticosteroid/jeL2QeIL">corticosteroids</a> may be a useful adjunct in treatment, as they protect blood cells against hemolysis, thereby reducing complications.<a class="footnote-ref" id="fnref:93" href="#fn:93"><sup>93</sup></a> <a href="/facts/Plasmapheresis/pt2AfdrU">Plasmapheresis</a> reduces the circulating immune complexes in the blood which cause the hemolysis, and may be beneficial in improving the outcome.<a class="footnote-ref" id="fnref:94" href="#fn:94"><sup>94</sup></a><a class="footnote-ref" id="fnref:95" href="#fn:95"><sup>95</sup></a> Additionally, <a href="/facts/Hemodialysis/oB7fIKP7">hemodialysis</a> can be used for patients with compromised kidney function or kidney failure.<a class="footnote-ref" id="fnref:96" href="#fn:96"><sup>96</sup></a> </p><p><i>Paxillus involutus</i> also contains agents which appear to damage <a href="/facts/Chromosome/FRGf11FC">chromosomes</a>; it is unclear whether these have <a href="/facts/Carcinogen/soH8Y9CE">carcinogenic</a> or <a href="/facts/Mutagen/xI5lKvmp">mutagenic</a> potential.<a class="footnote-ref" id="fnref:97" href="#fn:97"><sup>97</sup></a> Two compounds that have been identified are the <a href="/facts/Natural_phenol/sMCro6sW">phenols</a> involutone<a class="footnote-ref" id="fnref:98" href="#fn:98"><sup>98</sup></a> and <a href="/facts/Involutin/Tuq0W11v">involutin</a>; the latter is responsible for the brownish discolouration upon bruising.<a class="footnote-ref" id="fnref:99" href="#fn:99"><sup>99</sup></a> </p><p>Despite the poisonings, <i>Paxillus involutus</i> is still consumed in parts of <a href="/facts/Poland/eXJ7fd0D">Poland</a>, <a href="/facts/Russia/jfkCwyxn">Russia</a>, and <a href="/facts/Ukraine/Qof0uqAj">Ukraine</a>, where people die from it every year.<a class="footnote-ref" id="fnref:100" href="#fn:100"><sup>100</sup></a><a class="footnote-ref" id="fnref:101" href="#fn:101"><sup>101</sup></a><a class="footnote-ref" id="fnref:102" href="#fn:102"><sup>102</sup></a><a class="footnote-ref" id="fnref:103" href="#fn:103"><sup>103</sup></a> </p> <h2 id="see-also">See also</h2> <ul> <li>Fungi portal</li></ul> <ul><li><a href="/facts/List_of_deadly_fungi/jXuGQQRh">List of deadly fungi</a></li></ul> <h2 id="external-links">External links</h2> <ul><li> Media related to Paxillus involutus at Wikimedia Commons</li> <li> Data related to Paxillus involutus at Wikispecies</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Bulliard, J.B.F. (1785). Herbier de la France (in French). Vol. 5. Paris, France: Chez l'auteur. pp. 192–240. <a href="/wiki/Jean_Baptiste_Fran%C3%A7ois_Pierre_Bulliard" target="_blank">/wiki/Jean_Baptiste_Fran%C3%A7ois_Pierre_Bulliard</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Batsch, August (1786). Elenchus Fungorum (Discussion of Fungi), Continuatio Prima (in Latin). Halle, Magdeburg, Germany: Apud Joannem J. Gebauer. p. 39. <a href="/wiki/August_Batsch" target="_blank">/wiki/August_Batsch</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>"Paxillus involutus (Batsch) Fr. 1838". MycoBank. International Mycological Association. Retrieved 29 June 2011. <a href="http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=191172" target="_blank">http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=191172</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Bolton, James (1788). An History of Fungusses, Growing about Halifax. Vol. 2. Huddersfield, UK: self-published. p. 55. <a href="/wiki/James_Bolton" target="_blank">/wiki/James_Bolton</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>"Agaricus adscendibus Bolton". Index Fungorum. CAB International. Retrieved 15 July 2011. <a href="http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=462407" target="_blank">http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=462407</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Gray, Samuel F. (1821). Natural Arrangement of British Plants. London: Baldwin, Cradock, and Joy. p. 611. <a href="/wiki/Samuel_Frederick_Gray" target="_blank">/wiki/Samuel_Frederick_Gray</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Rabenhorst, Gottlob Ludwig (1844). Deutschlands Kryptogamenflora (in German). Vol. 1 (2 ed.). Leipzig, Germany: E. Kummer. p. 453. <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Fries, Elias Magnus (1838). Epicrisis Systematis Mycologici (in Latin). Uppsala, Sweden: Typographia Academica. p. 317. Archived from the original on 23 September 2015. Retrieved 15 July 2011. <a href="/wiki/Elias_Magnus_Fries" target="_blank">/wiki/Elias_Magnus_Fries</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Esser, K.; Lemke, P.A. (1994). The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research. Heidelberg, Germany: Springer. p. 81. ISBN 3-540-66493-9. <a href="3-540-66493-9" target="_blank">3-540-66493-9</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Maire, René (1902). "Recherches cytologiques et taxonomiques sur les Basidiomycetes". Bulletin trimestriel de la Société Mycologique de France (in French). 18 (supplement): 165. <a href="/wiki/Ren%C3%A9_Maire" target="_blank">/wiki/Ren%C3%A9_Maire</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Nilson, Sven; Persson, Ole (1977). Fungi of Northern Europe 1: Larger Fungi (Excluding Gill-Fungi). Harmondsworth, UK: Penguin. p. 121. ISBN 0-14-063005-8. <a href="0-14-063005-8" target="_blank">0-14-063005-8</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Nilson, Sven; Persson, Ole (1977). Fungi of Northern Europe 1: Larger Fungi (Excluding Gill-Fungi). Harmondsworth, UK: Penguin. p. 121. ISBN 0-14-063005-8. <a href="0-14-063005-8" target="_blank">0-14-063005-8</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Ziobro, G. (2012). Lampel, Keith A.; Al-Khaldi, Sufian; Cahil, Susan Mary (eds.). "Mushroom Toxins" (PDF). Bad Bug Book: Foodborne Pathogenic Microorganisms and Natural Toxins Handbook (2nd ed.). USFDA/Center for Food Safety & Applied Nutrition. p. 204. <a href="https://www.fda.gov/downloads/Food/FoodSafety/FoodborneIllness/FoodborneIllnessFoodbornePathogensNaturalToxins/BadBugBook/UCM297627.pdf" target="_blank">https://www.fda.gov/downloads/Food/FoodSafety/FoodborneIllness/FoodborneIllnessFoodbornePathogensNaturalToxins/BadBugBook/UCM297627.pdf</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Lincoff, Gary H. (2008). National Audubon Society Field Guide to North American Mushrooms. New York: Alfred A. Knopf. p. 671. ISBN 978-0-394-51992-0. <a href="978-0-394-51992-0" target="_blank">978-0-394-51992-0</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>Arora, David (1986). Mushrooms Demystified: a Comprehensive Guide to the Fleshy Fungi. Berkeley, California: Ten Speed Press. pp. 477–78. ISBN 0-89815-169-4. <a href="0-89815-169-4" target="_blank">0-89815-169-4</a> <a href="#fnref:15" class="footnote-back-ref">↩</a></p></li> <li id="fn:16"><p>Benjamin, Denis R. (1995). Mushrooms: Poisons and Panaceas – A Handbook for Naturalists, Mycologists and Physicians. New York: WH Freeman and Company. pp. 382–84. ISBN 0-7167-2600-9. <a href="0-7167-2600-9" target="_blank">0-7167-2600-9</a> <a href="#fnref:16" class="footnote-back-ref">↩</a></p></li> <li id="fn:17"><p>Barceloux, Donald G. (2008). Medical Toxicology of Natural Substances: Foods, Fungi, Medicinal Herbs, Plants, and Venomous Animals. Hoboken, New Jersey: John Wiley and Sons. pp. 312–13. ISBN 978-0-471-72761-3. <a href="978-0-471-72761-3" target="_blank">978-0-471-72761-3</a> <a href="#fnref:17" class="footnote-back-ref">↩</a></p></li> <li id="fn:18"><p>Gray, Samuel F. (1821). Natural Arrangement of British Plants. London: Baldwin, Cradock, and Joy. p. 611. <a href="/wiki/Samuel_Frederick_Gray" target="_blank">/wiki/Samuel_Frederick_Gray</a> <a href="#fnref:18" class="footnote-back-ref">↩</a></p></li> <li id="fn:19"><p>Hahn, C.; Agerer, R. (1999). "Studien zum Paxillus involutus Formenkreis" [Studies on the Paxillus involutus complex]. Nova Hedwigia (in German). 69 (1–2): 241–310. doi:10.1127/nova.hedwigia/69/1999/241. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:19" class="footnote-back-ref">↩</a></p></li> <li id="fn:20"><p>Hedh, Jenny; Samson, Peter; Erland, Susanne; Tunlid, Anders (2008). "Multiple gene genealogies and species recognition in the ectomycorrhizal fungus Paxillus involutus". Mycological Research. 112 (8): 965–75. doi:10.1016/j.mycres.2008.01.026. PMID 18554888. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:20" class="footnote-back-ref">↩</a></p></li> <li id="fn:21"><p>Fries, Nils (1985). "Intersterility groups in Paxillus involutus". Mycotaxon. 24: 403–10. Archived from the original on 23 September 2015. Retrieved 30 June 2011. <a href="https://web.archive.org/web/20150923232648/http://www.cybertruffle.org.uk/cyberliber/59575/0024/0403.htm" target="_blank">https://web.archive.org/web/20150923232648/http://www.cybertruffle.org.uk/cyberliber/59575/0024/0403.htm</a> <a href="#fnref:21" class="footnote-back-ref">↩</a></p></li> <li id="fn:22"><p>Jarosch, Margit; Bresinsky, A. (1999). "Speciation and phylogenetic distances within Paxillus s. str. (Basidiomycetes, Boletales)". Plant Biology. 1 (6): 701–05. Bibcode:1999PlBio...1..701J. doi:10.1111/j.1438-8677.1999.tb00283.x. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:22" class="footnote-back-ref">↩</a></p></li> <li id="fn:23"><p>Hedh, Jenny; Samson, Peter; Erland, Susanne; Tunlid, Anders (2008). "Multiple gene genealogies and species recognition in the ectomycorrhizal fungus Paxillus involutus". Mycological Research. 112 (8): 965–75. doi:10.1016/j.mycres.2008.01.026. PMID 18554888. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:23" class="footnote-back-ref">↩</a></p></li> <li id="fn:24"><p>Hehd, Jenny; Johannsson, Tomas; Tunlid, Anders (2009). "Variation in host specificity and gene content in strains from genetically isolated lineages of the ectomycorrhizal fungus Paxillus involutus s. lat.". Mycorrhiza. 19 (8): 549–58. Bibcode:2009Mycor..19..549H. doi:10.1007/s00572-009-0252-3. PMID 19452174. S2CID 21611651. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:24" class="footnote-back-ref">↩</a></p></li> <li id="fn:25"><p>Nilson, Sven; Persson, Ole (1977). Fungi of Northern Europe 1: Larger Fungi (Excluding Gill-Fungi). Harmondsworth, UK: Penguin. p. 121. ISBN 0-14-063005-8. <a href="0-14-063005-8" target="_blank">0-14-063005-8</a> <a href="#fnref:25" class="footnote-back-ref">↩</a></p></li> <li id="fn:26"><p>Haas, Hans (1969). The Young Specialist looks at Fungi. London: Burke. p. 54. ISBN 0-222-79409-7. <a href="0-222-79409-7" target="_blank">0-222-79409-7</a> <a href="#fnref:26" class="footnote-back-ref">↩</a></p></li> <li id="fn:27"><p>Henrici, Alick (2004). "A key to Paxillus s.l. in Europe". Field Mycology. 5 (3): 87–88. doi:10.1016/S1468-1641(10)60560-9. <a href="https://doi.org/10.1016%2FS1468-1641%2810%2960560-9" target="_blank">https://doi.org/10.1016%2FS1468-1641%2810%2960560-9</a> <a href="#fnref:27" class="footnote-back-ref">↩</a></p></li> <li id="fn:28"><p>Chiu, Siu-Wai; Moore, David (1996). Patterns in Fungal Development. Cambridge, UK: Cambridge University Press. p. 207. ISBN 0-521-56047-0. <a href="0-521-56047-0" target="_blank">0-521-56047-0</a> <a href="#fnref:28" class="footnote-back-ref">↩</a></p></li> <li id="fn:29"><p>Haas, Hans (1969). The Young Specialist looks at Fungi. London: Burke. p. 54. ISBN 0-222-79409-7. <a href="0-222-79409-7" target="_blank">0-222-79409-7</a> <a href="#fnref:29" class="footnote-back-ref">↩</a></p></li> <li id="fn:30"><p>Zeitlmayr, Linus (1976). Wild Mushrooms: An Illustrated Handbook. Hertfordshire, UK: Garden City Press. p. 75. ISBN 0-584-10324-7. <a href="0-584-10324-7" target="_blank">0-584-10324-7</a> <a href="#fnref:30" class="footnote-back-ref">↩</a></p></li> <li id="fn:31"><p>Davis, R. Michael; Sommer, Robert; Menge, John A. (2012). Field Guide to Mushrooms of Western North America. Berkeley: University of California Press. pp. 232–233. ISBN 978-0-520-95360-4. 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