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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A phylogenetic study of <italic>Boletus</italic> section <italic>Boletus</italic> in Europe</title><author><name sortKey="Beugelsdijk, D C M" sort="Beugelsdijk, D C M" uniqKey="Beugelsdijk D" first="D. C. M." last="Beugelsdijk">D. C. M. Beugelsdijk</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author><author><name sortKey="Van Der Linde, S" sort="Van Der Linde, S" uniqKey="Van Der Linde S" first="S." last="Van Der Linde">S. Van Der Linde</name><affiliation><nlm:aff id="A2"> The Macaulay Institute, Craigiebuckler, Aberdeen, AB15 8QH, UK.</nlm:aff></affiliation></author><author><name sortKey="Zuccarello, G C" sort="Zuccarello, G C" uniqKey="Zuccarello G" first="G. C." last="Zuccarello">G. C. Zuccarello</name><affiliation><nlm:aff id="A3"> School of Biological Sciences, Victoria University of Wellington, P.O. Box 600, Wellington, 6140 New Zealand.</nlm:aff></affiliation></author><author><name sortKey="Den Bakker, H C" sort="Den Bakker, H C" uniqKey="Den Bakker H" first="H. C." last="Den Bakker">H. C. Den Bakker</name><affiliation><nlm:aff id="A4"> Department of Plant Pathology, Cornell University, 334 Plant Science, Ithaca, NY 14853, USA.</nlm:aff></affiliation></author><author><name sortKey="Draisma, S G A" sort="Draisma, S G A" uniqKey="Draisma S" first="S. G. A." last="Draisma">S. G. A. Draisma</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author><author><name sortKey="Noordeloos, M E" sort="Noordeloos, M E" uniqKey="Noordeloos M" first="M. E." last="Noordeloos">M. E. Noordeloos</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">20467482</idno><idno type="pmc">2865352</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2865352</idno><idno type="RBID">PMC:2865352</idno><idno type="doi">10.3767/003158508X283692</idno><date when="2008">2008</date><idno type="wicri:Area/Pmc/Corpus">000241</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000241</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A phylogenetic study of <italic>Boletus</italic> section <italic>Boletus</italic> in Europe</title><author><name sortKey="Beugelsdijk, D C M" sort="Beugelsdijk, D C M" uniqKey="Beugelsdijk D" first="D. C. M." last="Beugelsdijk">D. C. M. Beugelsdijk</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author><author><name sortKey="Van Der Linde, S" sort="Van Der Linde, S" uniqKey="Van Der Linde S" first="S." last="Van Der Linde">S. Van Der Linde</name><affiliation><nlm:aff id="A2"> The Macaulay Institute, Craigiebuckler, Aberdeen, AB15 8QH, UK.</nlm:aff></affiliation></author><author><name sortKey="Zuccarello, G C" sort="Zuccarello, G C" uniqKey="Zuccarello G" first="G. C." last="Zuccarello">G. C. Zuccarello</name><affiliation><nlm:aff id="A3"> School of Biological Sciences, Victoria University of Wellington, P.O. Box 600, Wellington, 6140 New Zealand.</nlm:aff></affiliation></author><author><name sortKey="Den Bakker, H C" sort="Den Bakker, H C" uniqKey="Den Bakker H" first="H. C." last="Den Bakker">H. C. Den Bakker</name><affiliation><nlm:aff id="A4"> Department of Plant Pathology, Cornell University, 334 Plant Science, Ithaca, NY 14853, USA.</nlm:aff></affiliation></author><author><name sortKey="Draisma, S G A" sort="Draisma, S G A" uniqKey="Draisma S" first="S. G. A." last="Draisma">S. G. A. Draisma</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author><author><name sortKey="Noordeloos, M E" sort="Noordeloos, M E" uniqKey="Noordeloos M" first="M. E." last="Noordeloos">M. E. Noordeloos</name><affiliation><nlm:aff id="A1"> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</nlm:aff></affiliation></author></analytic><series><title level="j">Persoonia : Molecular Phylogeny and Evolution of Fungi</title><idno type="ISSN">0031-5850</idno><idno type="eISSN">1878-9080</idno><imprint><date when="2008">2008</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>A phylogenetic study of the species in <italic>Boletus</italic> sect. <italic>Boletus</italic> was undertaken using the molecular markers ITS1-5.8S-ITS2 and GAPDH. Four well-supported lineages, one comprising <italic>Boletus edulis</italic> s.l., the others referring to <italic>B. aereus</italic>, <italic>B. reticulatus</italic> and <italic>B. pinophilus</italic> have been distinguished. The ML and MP trees of ITS showed remarkably low resolution within the <italic>B. edulis</italic> clade, and confirmed earlier published results, despite the use of samples from a wider geographical area and different hosts. The results of GAPDH demonstrate clearly that this low resolution must be ascribed to a low genetic variability with the <italic>B. edulis</italic> clade, and make clear that morphological and ecological characters have been overestimated within this species complex. <italic>Boletus edulis</italic> is therefore defined as a variable species with a wide morphological, ecological and geographic range, and includes several specific and subspecific taxa described in the literature (e.g. <italic>B. betulicola</italic>, <italic>B. persoonii</italic>, <italic>B. quercicola</italic> and <italic>B. venturii</italic>). Three other European species (<italic>B. aereus</italic>, <italic>B. pinophilus</italic> and <italic>B. reticulatus</italic>) are well delimited species based on morphology and our genetic data.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Binder, M" uniqKey="Binder M">M Binder</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Bruns, Td" uniqKey="Bruns T">TD Bruns</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Den Bakker, Hc" uniqKey="Den Bakker H">HC Den Bakker</name></author><author><name sortKey="Gravendeel, B" uniqKey="Gravendeel B">B Gravendeel</name></author><author><name sortKey="Kuyper, Tw" uniqKey="Kuyper T">TW Kuyper</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Den Bakker, Hc" uniqKey="Den Bakker H">HC Den Bakker</name></author><author><name sortKey="Noordeloos, Me" uniqKey="Noordeloos M">ME Noordeloos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Den Bakker, Hc" uniqKey="Den Bakker H">HC Den Bakker</name></author><author><name sortKey="Zuccarello, Gc" uniqKey="Zuccarello G">GC Zuccarello</name></author><author><name sortKey="Kuyper, Twm" uniqKey="Kuyper T">TWM Kuyper</name></author><author><name sortKey="Noordeloos, Me" uniqKey="Noordeloos M">ME Noordeloos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Felsenstein, J" uniqKey="Felsenstein J">J Felsenstein</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hasegawa, M" uniqKey="Hasegawa M">M Hasegawa</name></author><author><name sortKey="Kishino, K" uniqKey="Kishino K">K Kishino</name></author><author><name sortKey="Yano, T" uniqKey="Yano T">T Yano</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Huelsenbeck, Jp" uniqKey="Huelsenbeck J">JP Huelsenbeck</name></author><author><name sortKey="Ronquist, F" uniqKey="Ronquist F">F Ronquist</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hughes, Kw" uniqKey="Hughes K">KW Hughes</name></author><author><name sortKey="Mcghee, Ll" uniqKey="Mcghee L">LL McGhee</name></author><author><name sortKey="Mehven, As" uniqKey="Mehven A">AS Mehven</name></author><author><name sortKey="Johnson, Je" uniqKey="Johnson J">JE Johnson</name></author><author><name sortKey="Petersen, Rh" uniqKey="Petersen R">RH Petersen</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kimura, M" uniqKey="Kimura M">M Kimura</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kretzer, A" uniqKey="Kretzer A">A Kretzer</name></author><author><name sortKey="Li, Y" uniqKey="Li Y">Y Li</name></author><author><name sortKey="Szaro, Tm" uniqKey="Szaro T">TM Szaro</name></author><author><name sortKey="Bruns, Td" uniqKey="Bruns T">TD Bruns</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Leonardi, M" uniqKey="Leonardi M">M Leonardi</name></author><author><name sortKey="Paolocci, F" uniqKey="Paolocci F">F Paolocci</name></author><author><name sortKey="Rubini, A" uniqKey="Rubini A">A Rubini</name></author><author><name sortKey="Simonini, G" uniqKey="Simonini G">G Simonini</name></author><author><name sortKey="Pacioni, G" uniqKey="Pacioni G">G Pacioni</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Maddison, W" uniqKey="Maddison W">W Maddison</name></author><author><name sortKey="Maddison, D" uniqKey="Maddison D">D Maddison</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Moncalvo, Jm" uniqKey="Moncalvo J">JM Moncalvo</name></author><author><name sortKey="Vilgalys, R" uniqKey="Vilgalys R">R Vilgalys</name></author><author><name sortKey="Redhead, Sa" uniqKey="Redhead S">SA Redhead</name></author><author><name sortKey="Johnson, Je" uniqKey="Johnson J">JE Johnson</name></author><author><name sortKey="James, Ty" uniqKey="James T">TY James</name></author><author><name sortKey="Aime, Mc" uniqKey="Aime M">MC Aime</name></author><author><name sortKey="Hofstetter, V" uniqKey="Hofstetter V">V Hofstetter</name></author><author><name sortKey="Verduin, Sjw" uniqKey="Verduin S">SJW Verduin</name></author><author><name sortKey="Larsson, E" uniqKey="Larsson E">E Larsson</name></author><author><name sortKey="Baroni, Tj" uniqKey="Baroni T">TJ Baroni</name></author><author><name sortKey="Thorn, Rg" uniqKey="Thorn R">RG Thorn</name></author><author><name sortKey="Jacobsson, S" uniqKey="Jacobsson S">S Jacobsson</name></author><author><name sortKey="Clemencon, H" uniqKey="Clemencon H">H Clémençon</name></author><author><name sortKey="Miller, Ok" uniqKey="Miller O">OK Miller</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Posada, D" uniqKey="Posada D">D Posada</name></author><author><name sortKey="Crandall, Ka" uniqKey="Crandall K">KA Crandall</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Singer, R" uniqKey="Singer R">R Singer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Swofford, Dl" uniqKey="Swofford D">DL Swofford</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Thompson, Jd" uniqKey="Thompson J">JD Thompson</name></author><author><name sortKey="Gibson, Tj" uniqKey="Gibson T">TJ Gibson</name></author><author><name sortKey="Plewniak, F" uniqKey="Plewniak F">F Plewniak</name></author><author><name sortKey="Jeanmougin, F" uniqKey="Jeanmougin F">F Jeanmougin</name></author><author><name sortKey="Higgins, Dg" uniqKey="Higgins D">DG Higgins</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Van Der Linde, S" uniqKey="Van Der Linde S">S Van der Linde</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Van Der Linde, S" uniqKey="Van Der Linde S">S Van der Linde</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="White, Tj" uniqKey="White T">TJ White</name></author><author><name sortKey="Bruns, Td" uniqKey="Bruns T">TD Bruns</name></author><author><name sortKey="Lee, S" uniqKey="Lee S">S Lee</name></author><author><name sortKey="Taylor, J" uniqKey="Taylor J">J Taylor</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Persoonia</journal-id><journal-id journal-id-type="publisher-id">Persoonia</journal-id><journal-title-group><journal-title>Persoonia : Molecular Phylogeny and Evolution of Fungi</journal-title></journal-title-group><issn pub-type="ppub">0031-5850</issn><issn pub-type="epub">1878-9080</issn><publisher><publisher-name>Nationaal Herbarium Nederland & Centraallbureau voor Schimmelcultures</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">20467482</article-id><article-id pub-id-type="pmc">2865352</article-id><article-id pub-id-type="doi">10.3767/003158508X283692</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>A phylogenetic study of <italic>Boletus</italic> section <italic>Boletus</italic> in Europe</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Beugelsdijk</surname><given-names>D.C.M.</given-names></name><xref ref-type="aff" rid="A1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>van der Linde</surname><given-names>S.</given-names></name><xref ref-type="aff" rid="A2"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>Zuccarello</surname><given-names>G.C.</given-names></name><xref ref-type="aff" rid="A3"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>den Bakker</surname><given-names>H.C.</given-names></name><xref ref-type="aff" rid="A4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Draisma</surname><given-names>S.G.A.</given-names></name><xref ref-type="aff" rid="A1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Noordeloos</surname><given-names>M.E.</given-names></name><xref ref-type="aff" rid="A1"><sup>1</sup></xref><xref ref-type="corresp" rid="COR1"></xref></contrib></contrib-group><aff id="A1"><label>1</label> National Herbarium of the Netherlands, Leiden University branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands;</aff><aff id="A2"><label>2</label> The Macaulay Institute, Craigiebuckler, Aberdeen, AB15 8QH, UK.</aff><aff id="A3"><label>3</label> School of Biological Sciences, Victoria University of Wellington, P.O. Box 600, Wellington, 6140 New Zealand.</aff><aff id="A4"><label>4</label> Department of Plant Pathology, Cornell University, 334 Plant Science, Ithaca, NY 14853, USA.</aff><author-notes><corresp id="COR1">corresponding author e-mail: <email>noordeloos@nhn.leidenuniv.nl</email>.
</corresp></author-notes><pub-date pub-type="epub"><day>1</day><month>2</month><year>2008</year></pub-date><pub-date pub-type="ppub"><month>6</month><year>2008</year></pub-date><volume>20</volume><fpage>1</fpage><lpage>7</lpage><history><date date-type="received"><day>1</day><month>10</month><year>2007</year></date><date date-type="accepted"><day>16</day><month>11</month><year>2007</year></date></history><permissions><copyright-statement>© 2008 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures</copyright-statement><copyright-year>2008</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode"><license-p>© 2008 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures</license-p><license-p>You are free to share - to copy, distribute and transmit the work, under the following conditions:</license-p><license-p>Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).</license-p><license-p>Non-commercial: You may not use this work for commercial purposes.</license-p><license-p>No derivative works: You may not alter, transform, or build upon this work.</license-p><license-p><pmc-comment>CREATIVE COMMONS</pmc-comment>For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode">http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode.</ext-link> Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.</license-p></license></permissions><abstract abstract-type="executive-summary"><p>A phylogenetic study of the species in <italic>Boletus</italic> sect. <italic>Boletus</italic> was undertaken using the molecular markers ITS1-5.8S-ITS2 and GAPDH. Four well-supported lineages, one comprising <italic>Boletus edulis</italic> s.l., the others referring to <italic>B. aereus</italic>, <italic>B. reticulatus</italic> and <italic>B. pinophilus</italic> have been distinguished. The ML and MP trees of ITS showed remarkably low resolution within the <italic>B. edulis</italic> clade, and confirmed earlier published results, despite the use of samples from a wider geographical area and different hosts. The results of GAPDH demonstrate clearly that this low resolution must be ascribed to a low genetic variability with the <italic>B. edulis</italic> clade, and make clear that morphological and ecological characters have been overestimated within this species complex. <italic>Boletus edulis</italic> is therefore defined as a variable species with a wide morphological, ecological and geographic range, and includes several specific and subspecific taxa described in the literature (e.g. <italic>B. betulicola</italic>, <italic>B. persoonii</italic>, <italic>B. quercicola</italic> and <italic>B. venturii</italic>). Three other European species (<italic>B. aereus</italic>, <italic>B. pinophilus</italic> and <italic>B. reticulatus</italic>) are well delimited species based on morphology and our genetic data.</p></abstract><kwd-group><kwd><italic>Boletus</italic></kwd><kwd><italic>Boletus edulis</italic></kwd><kwd>Europe</kwd><kwd>phylogenetic study</kwd></kwd-group></article-meta></front><body><sec id="s1"><title>INTRODUCTION</title><p><italic>Boletus</italic> sect. <italic>Boletus</italic> is well characterised morphologically by the white, unchanging context, white pores becoming yellow to olive with age and a reticulate surface of the stipe (<xref ref-type="bibr" rid="R16">Singer 1986</xref>). Molecular data suggest that sect. <italic>Boletus</italic> is well delimited from the rest of the genus (<xref ref-type="bibr" rid="R1">Binder 1999</xref>).</p><p>Within sect. <italic>Boletus</italic> the species concept and species delimitation has however been a matter of dispute, resulting in a fairly large number of specific and infraspecific names and a complex nomenclatural history. Because most species of sect. <italic>Boletus</italic> are much sought for because of their culinary value, they have received attention not only from taxonomists, but also gastronomists. As a result many taxa have been described, often based on single ecological characters, such as mycorrhizal partner, or on highly variable morphological characters, such as the colour of the pileus. Particularly with regard to taxa similar to the type species of the section, <italic>B. edulis</italic>, a number of species and infraspecific taxa have been described, for example in relation to the mycorrhizal partner or on account of deviating colours of the basidiocarp (<xref ref-type="table" rid="T1">Table 1</xref>).</p><p><xref ref-type="bibr" rid="R19">Van der Linde (2002)</xref> made an extensive literature study followed by a morphological revision (<xref ref-type="bibr" rid="R20">Van der Linde 2004</xref>) of sect. <italic>Boletus</italic> in northwestern Europe. Morphological and morphometric studies on material from various geographic regions revealed that <italic>B. edulis</italic> has a wide ecological range, being associated with numerous different partners, both deciduous and coniferous trees (e.g. <italic>Betula</italic>, <italic>Fagus</italic>, <italic>Picea</italic>, <italic>Pinus</italic>, <italic>Quercus</italic> and <italic>Tilia</italic>). Therefore it includes several taxa that are supposed to be exclusively associated to certain hosts (<italic>B. betulicola</italic> and <italic>B. quercicola</italic>). Morphological characters, such as the colour of the pileus, development of a reticulum on the stipe, size and shape of the spores and the terminal elements of the pileipellis, varied a great deal, and could not be used to distinguish discrete morphological entities (<xref ref-type="bibr" rid="R20">Van der Linde 2004</xref>). As a result it was proposed to reduce the number of taxa to four (<xref ref-type="table" rid="T2">Table 2</xref>). Beside <italic>B. edulis</italic> and a white variety <italic>B. edulis</italic> var. <italic>albus</italic> (= <italic>B. persoonii</italic>)<italic>,</italic> three other species could be distinguished in Europe on morphological criteria, <italic>B. aereus, B. pinophilus</italic> and <italic>B. reticulatus</italic> (= <italic>B. aestivalis</italic>). <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref> distinguished furthermore three morphotaxa within the wide concept of <italic>B. edulis</italic>, one associated with <italic>Fagus</italic>, with a very dark pileus and strongly developed net on the stipe (morphotaxon A in the present paper), and one associated with <italic>Tilia</italic>, with a reddish pileus, strongly developed net and rather wide terminal elements of the pileipellis (= morphotaxon B). Morphotaxon C initially was thought to be <italic>B. pinophilus</italic> on account of its association with <italic>Pinus</italic> on nutrient-poor sandy soil in the Netherlands, but its colour and microscopical characters revealed that it should be considered a form of <italic>B. edulis</italic>.</p><p>Molecular data have been very useful in understanding phylogenetic relationships in Basidiomycota at all taxonomic levels (<xref ref-type="bibr" rid="R1">Binder 1999</xref>, <xref ref-type="bibr" rid="R14">Moncalvo et al. 2002</xref>, <xref ref-type="bibr" rid="R3">Den Bakker et al. 2004a</xref>, <xref ref-type="bibr" rid="R5">b</xref>). Using ITS2 and the glyceraldehyde 3-phosphate dehydrogenase gene (GAPDH) sequences, <xref ref-type="bibr" rid="R5">Den Bakker et al. (2004b)</xref> confirmed that host specificity can be an important character in distinguishing species in the genus <italic>Leccinum</italic>. <xref ref-type="bibr" rid="R4">Den Bakker & Noordeloos (2005)</xref> compared other morphological and ecological data with molecular data to revise the species of the genus <italic>Leccinum</italic> in Europe. <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref> studied the <italic>B. edulis</italic>-complex with molecular data obtained from the internal transcribed spacer 1-5.8S rRNA-internal transcribed spacer 2 (= ITS) using a total of 39 samples from Italy and France. Their results support roughly those of our morphological studies. <italic>Bolletus aereus</italic>, <italic>B. aestivalis</italic>, <italic>B. edulis</italic> and <italic>B. pinophilus</italic> formed four distinct groups. <italic>Bolletus edulis</italic> var. <italic>pusteriensis</italic>, <italic>B. persoonii</italic> and <italic>B. venturii</italic> could molecularly not be distinguished from <italic>B. edulis</italic>.</p><p>We wish to extend this analysis using two nuclear encoded regions: one the ITS regions as used by <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref>, and the GAPDH region which has been shown to give good interspecific resolution in Basidiomycota (<xref ref-type="bibr" rid="R5">Den Bakker et al. 2004b</xref>).</p><p>We will also use the specimens studied morphologically by <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref> from western and northern Europe to address the following questions:</p><list list-type="simple"><list-item><p>– Are taxa (species, subspecies, morphotypes) within the <italic>B. edulis</italic> complex genetically distinct?</p></list-item><list-item><p>– What further insights can be gained into the evolutionary history of <italic>Boletus</italic> with the addition of a second genetic marker (GAPDH)?</p></list-item><list-item><p>– How does our ITS data compare to previous results by <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref>?</p></list-item></list></sec><sec id="s2" sec-type="methods"><title>MATERIALS AND METHODS</title><sec id="s2a"><title>Selection of specimens</title><p>A selection was made from the material used by <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref> in his morphological studies. These were collected mainly in the Netherlands in 2001 and 2002 and during collecting trips to Borgjö (Sweden) in August 2001, Vuokatti (Finland) in September 2001 and central Austria in 2002. A list of collections used is given in <xref ref-type="table" rid="T3">Table 3</xref>.</p></sec><sec id="s2b"><title>DNA extraction</title><p>DNA was extracted from herbarium specimens using the method described in <xref ref-type="bibr" rid="R3">Den Bakker et al. (2004a)</xref>. The presence of DNA was checked on a 1 % agarose gel in 0.5× TBE, stained with ethidium bromide.</p></sec><sec id="s2c"><title>Amplification</title><p>For amplification of ITS, the forward primer ITS5 and the reverse ITS4 (<xref ref-type="bibr" rid="R21">White et al. 1990</xref>) were used. Total reaction volume was 25 μl containing 2 μl of diluted template DNA. The PCR reaction was performed using the following protocol: an initial denaturation step of 94 °C for 4 min, followed by 37 cycles of: 1 min denaturation at 94 °C, annealing at 52 °C for 1 min and extension at 72 °C for 1 min. This was followed by a final extension of 5 min at 72 °C. The amplified DNA samples were checked on a 1 % agarose gel stained with ethidium bromide. PCR products were cleaned using the QIAquick PCR Purification (Qiagen, Hilden, Germany). In cases where multiple bands were found, the PCR products with the correct length (± 850 bp) were cut out and cleaned following the QIAquick Gel Extraction Kit Protocol (Qiagen, Hilden, Germany).</p><p>From the single copy nuclear gene GAPDH only the last half of the gene was amplified. Forward primer GPD0623F (5′-TTGCCAAGGTCGTCAACG-3′) and reverse primer GPD reverse (5′-GAGTAWCCSCATTCGTTATCGTACC-3′) were used from <xref ref-type="bibr" rid="R5">Den Bakker et al. (2004b)</xref>. The PCR reactions for amplification of GAPDH followed <xref ref-type="bibr" rid="R5">Den Bakker et al. (2004b)</xref>. The products were checked and cleaned as for ITS products.</p></sec><sec id="s2d"><title>Sequencing</title><p>The cleaned PCR products of ITS were sequenced using the ITS2, ITS3, ITS4 and ITS5 primers (<xref ref-type="bibr" rid="R21">White et al. 1990</xref>). The primers GPD0623F and GPD reverse, plus the PCR primers, were used for GAPDH sequencing. The samples were sequenced on an ABI 377 automated sequencer (Applied Biosystems, Foster City, CA, USA) using standard dye-terminator chemistry following the manufacturer’s protocols.</p></sec><sec id="s2e"><title>Phylogenetic analysis</title><p>The raw data were processed using Sequencher v. 4.0 (Gene Codes Corporation Ann Arbor, Michigan, USA). ITS sequences (with and without the samples from <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref>) were aligned in ClustalX (<xref ref-type="bibr" rid="R18">Thompson et al. 1997</xref>) and refined by eye. The ClustalX parameters were gap opening penalty of 5 and a gap extension penalty of 3. Alignment of the GAPDH sequences was done by eye in Se-Al (v. 2.0-11) (A. Rambaut, University of Oxford, available from <ext-link ext-link-type="uri" xlink:href="http://evolve.zoo.ox.ac.uk/software.html">http://evolve.zoo.ox.ac.uk/software.html</ext-link>).</p><p>To increase phylogenetic resolution the ITS and GAPDH data sets were combined in MacClade 4.0.5 (<xref ref-type="bibr" rid="R13">Maddison & Maddison 2002</xref>) for 31 taxa in which both genes were sequenced. To test a priori whether the ITS and GAPDH datasets contained a congruent phylogenetic signal a partition homogeneity test was performed using 100 replicates, 10 random sequence additions, maxtrees set to 100, TBR branch swapping, unordered and unweighted characters and gaps treated as missing. Maximum parsimony (MP) and maximum likelihood (ML) analyses were performed in PAUP v. 4.0 (<xref ref-type="bibr" rid="R17">Swofford 2002</xref>) for the combined ITS-GAPDH sequences and all the ITS sequences (this study and <xref ref-type="bibr" rid="R12">Leonardi et al. 2005</xref>).</p><p>For the MP analysis the heuristic search option, 100 random sequence additions, tree bisection reconnection (TBR) branch swapping, gaps treated as missing data and unordered and unweighted characters were used. For measuring relative support of the clades, a bootstrap analysis (<xref ref-type="bibr" rid="R6">Felsenstein 1985</xref>) was performed using 1 000 bootstrap replicates, 10 random sequence additions and TBR branch swapping. To limit the length of the bootstrap analysis no more than 10 trees per (random sequence addition) replicate were saved.</p><p>Modeltest v. 3.06 (<xref ref-type="bibr" rid="R15">Posada & Crandall 1998</xref>) was used to find the model of sequence evolution that best fitted the data set using the hierarchical likelihood ratio test (hLRT). These parameters of the model were used in a ML analysis and a distance bootstrap in PAUP.</p><p>For the combined ITS-GAPDH sequences the HKY85+G substitution model (with Ti/Tv ratio = 2.1349, gamma shape parameter = 0.2526, and base frequencies set to: A = 0.2164, C = 0.2682, G = 0.2423 and T = 0.2731) was used (<xref ref-type="bibr" rid="R7">Hasegawa et al. 1985</xref>).</p><p>The ML analysis was performed with 10 random sequence additions and TBR branch swapping. No more than 100 trees were saved for each bootstrap replicate. For the combined ITS-GAPDH dataset, the single ML tree had a log-likelihood score of -3326.71559. A distance bootstrap analysis was conducted with 1 000 bootstrap replicates, neighbour-joining, TBR branch swapping and not more than 5 trees were saved per bootstrap replicate.</p><p>The ML analysis of all the ITS sequences (<xref ref-type="bibr" rid="R12">Leonardi et al. 2005</xref>, and this study) was performed using the substitution model HKY+G (with Ti/Tv ratio = 1.7679, gamma shape parameter = 0.3354, and base frequencies set to: A = 0.2154, C = 0.2542, G = 0.2416 and T = 0.2888) (<xref ref-type="bibr" rid="R7">Hasegawa et al. 1985</xref>). The other parameters were 10 random sequence additions, TBR branch swapping and maxtrees set to 100. The ML tree had a log likelihood score of 2492.05247.</p><p>For both the ITS-GAPDH and the ITS dataset (sequences of this study and <xref ref-type="bibr" rid="R12">Leonardi et al. 2005</xref>), a Bayesian analysis was performed in MrBayes v. 3.0b4 (<xref ref-type="bibr" rid="R8">Huelsenbeck & Ronquist 2001</xref>). The combined ITS-GAPDH dataset was partitioned to apply a different model of sequence evolution to each gene. These models were determined using MrModeltest (J.J.A. Nylander, available from: <ext-link ext-link-type="uri" xlink:href="http://www.ebc.uu.se/systzoo/staff/nylander.html">http://www.ebc.uu.se/systzoo/staff/nylander.html</ext-link>). The K80+G substition model with a Ti/Tv of 1.9919, equal base frequencies and gamma shape parameter of 0.2569 was applied to the ITS part of the alignment (<xref ref-type="bibr" rid="R10">Kimura 1980</xref>). For the GAPDH partition a HKY85+G subsitution model was used (with Ti/Tv = 2.0506, gamma shape parameter = 0.2956 and base frequencies set to: A = 0.2092, C = 0.3047, G = 0.2320 and T = 0.2540) (<xref ref-type="bibr" rid="R7">Hasegawa et al. 1985</xref>). The number of generations was set to 3 million and one tree was saved per 100 generations. A 50 % majority rule consensus tree was made in PAUP from the outcomes of the Bayesian analysis. A ‘burn-in’ of 1 million generations was used, which was well after stationarity was reached.</p></sec></sec><sec id="s3"><title>RESULTS</title><p>The ITS and GAPDH alignment consisted of 860 and 496 characters respectively of which 57 and 39 were parsimony informative. The partition homogeneity test indicated that the two molecular data sets (ITS and GAPDH) could be combined (<italic>P</italic> = 0.560). This alignment consisted of 1356 basepairs (bp) and contained 96 parsimony-informative sites. The MP analysis produced 6 600 most parsimonious trees of 305 steps. The strict consensus tree of the MP trees is congruent to the ML tree (<xref ref-type="fig" rid="F1">Fig. 1</xref>).</p><p>In <xref ref-type="fig" rid="F1">Fig. 1</xref> four well-supported clades are seen. Well-supported is used here for a MP and ML bootstrap (BP) above 70 % and/or posterior probability (PP) above 0.95. The first, <italic>B. edulis</italic> clade, includes the species <italic>B. betulicola</italic>, <italic>B. edulis</italic>, <italic>B. persoonii</italic>, <italic>B. quercicola</italic> and the three unnamed morphotaxa. There was no phylogenetic resolution within this group and very little genetic variation. The second clade, containing the <italic>B. pinophilus</italic> from Sweden (samples 22 and 23), is a sistergroup of the first clade. The third clade, containing <italic>B. aereus</italic> (samples 1 and 2) and <italic>B. reticulatus</italic> (sample 28) formed a sister group to the <italic>B. edulis</italic>/ <italic>B. pinophilus</italic> clades. This group was further divided with the two <italic>B. aereus</italic> samples (group 3) and their sister species <italic>B. reticulatus</italic> (sample 28, group 4).</p><p>The total ITS alignment consisted of 964 base pairs of which 157 bases were parsimony informative. The MP analysis produced > 10 000 most parsimonious trees of 214 steps. A strict consensus topology is shown in <xref ref-type="fig" rid="F2">Fig. 2</xref>. The ML tree and MP tree did not differ significantly in topology. Most ingroup relationships were not resolved, only four clades were supported with good to moderate BP or PP support. Moderate support is defined here as high support for BP (> 70 %) and low support for PP (< 0.95). The first, moderately supported clade is <italic>B. edulis,</italic> which includes the species <italic>B. betulicola</italic>, <italic>B. edulis</italic>, <italic>B. edulis</italic> var. <italic>pusteriensis</italic>, <italic>B. persoonii</italic>, <italic>B. quercicola</italic>, <italic>B. venturii</italic> and the three morphotaxa from various hosts and locations in Europe. There was no supported resolution within any of these clades. A second clade contains <italic>B. pinophilus</italic>. Finally, a <italic>B. aereus</italic> clade (group 3) and a <italic>B. reticulatus</italic> clade (group 4) (<italic>B. reticulatus</italic> = <italic>B. aestivalis</italic>), which also grouped into a well-supported clade.</p></sec><sec id="s4"><title>DISCUSSION</title><p>Phylogenetic analysis of the ITS and GAPDH and the ITS sequences including the samples of <xref ref-type="bibr" rid="R12">Leonaridi et al. (2005)</xref> consistently produced the same four clades. These clades are:</p><list list-type="order"><list-item><p>A clade containing a large number of samples identified as being <italic>B. betulicola</italic>, <italic>B. edulis</italic>, <italic>B. edulis</italic> var. <italic>albus, B. edulis</italic> var. <italic>pusteriensis, B. quercicola, B. venturii</italic> and morphotaxon A, B, C;</p></list-item><list-item><p><italic>B. pinophilus</italic>;</p></list-item><list-item><p><italic>B. aereus</italic>; and</p></list-item><list-item><p><italic>B. reticulatus</italic>.</p></list-item></list><p>There is no resolution within the <italic>B. edulis</italic> group. All the clades are well to moderately supported using bootstrap and posterior probabilities.</p><p>Within the <italic>B. edulis</italic> clade, almost no genetic variation was observed, though many distinguishable morphotypes were included plus a wide geographic sampling. Within each group no differences has been found between Northern Europe and Southern Europe <italic>B. edulis</italic>, <italic>B. reticulatus</italic> and <italic>B. pinophilus</italic>.</p><p>Our molecular results and those of <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref> indicate that within the <italic>B. edulis</italic> clade there is little genetic variation, even for a marker often used in species level distinction in the fungi. The ITS is commonly used for many organisms and is useful for separating species in Basidiomycetes. For example, <xref ref-type="bibr" rid="R9">Hughes et al. (1999)</xref> used ITS sequences to distinguish species of the genus <italic>Flammulina</italic> and <xref ref-type="bibr" rid="R11">Kretzer et al. (1996)</xref> used ITS for recognizing different species in the genus <italic>Suillus</italic> s.l. But the ITS region also has limitations. According to <xref ref-type="bibr" rid="R2">Bruns (2001)</xref>, there is often little genetic variation among very closely related species. To corroborate the results of the ITS data, we used an additional section of DNA (GAPDH) as this region was useful in distinguishing species of the genus <italic>Leccinum</italic> (<xref ref-type="bibr" rid="R5">Den Bakker 2004b</xref>). Very little genetic differences were also found using the GAPDH dataset. The assignment of different putative species, for specimens that show slight morphological variation or different host plants, cannot be supported by our genetic data suggesting that they are all one species.</p><p>A collection with a purely white fruit body, often referred to as <italic>B. persoonii</italic> in the literature, appeared to be similar to <italic>B. edulis</italic> in morphology, and must therefore be considered a mere white form of that species, for which the name <italic>B. edulis</italic> var. <italic>albus</italic> is available. Three morphotaxa, which could be distinguished from <italic>B. edulis</italic> in the morphological study of <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref>, did not get support from the molecular data and must be considered to fall within the genetic variability of <italic>B. edulis</italic> (see <xref ref-type="table" rid="T2">Table 2</xref>).</p><p>The lack of resolution in the <italic>B. edulis</italic> clade due to low variability of ITS and GAPDH, suggests that there is an overestimation of the significance of morphological and ecological characters, as was already made apparent by the study of <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref>. We therefore consider all taxa in the <italic>B. edulis</italic> clade as belonging to one morphologically variable species.</p><p>GAPDH data was able to improve the support of the recognised clades from the ITS data but not resolve relationships, especially within the <italic>B. edulis</italic> clade.</p><p>In conclusion, host plant specificity has a very restricted value for the recognition of species in sect. <italic>Boletus</italic>. <italic>Boletus edulis</italic> appears to be associated with a large number of host trees, both deciduous and coniferous. <italic>Boletus pinophilus</italic> seems to be the only species within the section which has a rather strict association with <italic>Pinus</italic>, although it sometimes also is found in pure <italic>Picea</italic> or <italic>Abies</italic> stands. The latter is sometimes referred to as <italic>B. pinophilus</italic> var. <italic>fuscoruber</italic>. <italic>Boletus reticulatus</italic> and <italic>B. aereus</italic> are restricted to a number of <italic>Quercus</italic> species.</p><p>We therefore recognise four species throughout Europe: <italic>B. aereus, B. edulis,</italic><italic>B. pinophilus</italic> and <italic>B. reticulatus</italic>.</p></sec></body><back><ack><p>Marcel Eurlings and Rene Glas are greatly thanked for their assistance in the molecular laboratory; Dr H.C. Mauri Korhonen, University of Helsinki, contributed much to our understanding of the morphological concepts in sect. <italic>Boletus</italic> during numerous discussions while collecting in Sweden and Finland, for which we are very grateful. Prof. dr Th.W. 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</mixed-citation></ref></ref-list></back><floats-group><table-wrap id="T1" orientation="portrait" position="float"><label>Table 1</label><caption><p>Species and infraspecific taxa described in section <italic>Boletus</italic> in Europe.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Name and authors</th><th align="left" rowspan="1" colspan="1">Host</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>B. aereus</italic> Bull.</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic> (Vasilkov) Pilat & Dermek</td><td align="left" rowspan="1" colspan="1"><italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. carpinaceus</italic> Velen.</td><td align="left" rowspan="1" colspan="1"><italic>Carpinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> Bull. var. <italic>edulis</italic></td><td align="left" rowspan="1" colspan="1">Various hosts (coniferous and deciduous)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>albus</italic> Persoon = <italic>B. persoonii</italic> M.Bon</td><td align="left" rowspan="1" colspan="1"><italic>Pinus sylvestris</italic> and deciduous trees</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>arcticus</italic> Vasilkov</td><td align="left" rowspan="1" colspan="1"><italic>Betula nana</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>arenarius</italic> Engel & al.</td><td align="left" rowspan="1" colspan="1"><italic>Pinus sylvestris</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> forma <italic>aurantioruber</italic> (Dick & Snell) Vassilkov</td><td align="left" rowspan="1" colspan="1"><italic>Abies</italic>, <italic>Picea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>citrinus</italic> Pelt.</td><td align="left" rowspan="1" colspan="1">Unknown</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> subsp. <italic>clavipes</italic> (Peck) Singer</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic>, <italic>Abies</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>pusturiensis</italic> Ferrarese & Simonini</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic> and <italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> forma <italic>roseipes</italic> Vassilkov</td><td align="left" rowspan="1" colspan="1">Unknown</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>subhepaticus</italic> Fayod</td><td align="left" rowspan="1" colspan="1"><italic>Fagus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> forma <italic>viridicaerulescens</italic> Estades & Lannoy</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic>, <italic>Abies</italic>, and <italic>Pinus uncinata</italic>, <italic>P. cembra</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> subsp. <italic>trisporus</italic> Watling</td><td align="left" rowspan="1" colspan="1">Unknown</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. fulvomaculatus</italic> Estades & Lannoy</td><td align="left" rowspan="1" colspan="1">Deciduous trees (<italic>Quercus</italic>)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. pinophilus</italic> Pilat & Dermek</td><td align="left" rowspan="1" colspan="1"><italic>Pinus</italic> (<italic>Picea</italic>)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. pinophilus</italic> var. <italic>fuscoruber</italic> (Forq.) Estades & Lannoy</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic>, <italic>Abies</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. quercicola</italic> (Vassilkov) Singer</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. reticulatus</italic> Schaeff. (= <italic>B. aestivalis</italic>) Paulet</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>B. venturii</italic> M. Bon</td><td align="left" rowspan="1" colspan="1">Deciduous and coniferous trees</td></tr></tbody></table></table-wrap><table-wrap id="T2" orientation="portrait" position="float"><label>Table 2</label><caption><p>Species in sect. <italic>Boletus</italic> as accepted by <xref ref-type="bibr" rid="R20">Van der Linde (2004)</xref>.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Species</th><th align="left" rowspan="1" colspan="1">Morphological characteristics</th><th align="left" rowspan="1" colspan="1">Host and ecology</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>Boletus aereus</italic> Bull.</td><td align="left" rowspan="1" colspan="1">Pileus dry, finely tomentose, often also finely pruinose-pulverulent in marginal zone; very dark brown; stipe reticulate over whole length; dark brown; stipe reticulate over whole length; pileipellis elements up to 11 μm wide with fine intracellular grains</td><td align="left" rowspan="1" colspan="1">With <italic>Quercus</italic>, on heavy loamy, often somewhat calcareous soil; in road-sides and parks; thermophilic</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Boletus edulis</italic> Bull. var. <italic>edulis</italic></td><td align="left" rowspan="1" colspan="1">Pileus surface glabrous, greasy to touch, pale to dark brown, yellow brown, reddish brown (white in var. <italic>albus</italic>); stipe with fine reticulation in upper half, rarely over entire length. Pileipellis with terminal elements 4.0–19 μm wide in intracellular pigment</td><td align="left" rowspan="1" colspan="1">With a broad spectre of both deciduous and coniferous trees (<italic>Picea</italic>, <italic>Quercus</italic>, <italic>Betula</italic>, <italic>Fagus</italic>, <italic>Tilia</italic>, rarely <italic>Pinus</italic>). On various soil types, ranging from moist to fairly dry</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Boletus edulis</italic> var. <italic>albus</italic> Persoon</td><td align="left" rowspan="1" colspan="1">As in the type-variety, but fruitbodies purely white</td><td align="left" rowspan="1" colspan="1">With <italic>Pinus</italic> or <italic>Quercus</italic>, on humose soil</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Boletus pinophilus</italic> Pilát & Dermek</td><td align="left" rowspan="1" colspan="1">Pileus with thick, gelatinous pellicle, dark red brown, glabrous; stipe with coarse reticulation over the top, rarely over entire length; pileipellis with terminal elements up to 27 μm wide with reddish brown parietal and incrusting pigment, which dissolves in KOH</td><td align="left" rowspan="1" colspan="1">Almost exclusively with <italic>Pinus</italic>, rarely with <italic>Picea</italic>, on poor, acid, sandy soil</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Boletus reticulatus</italic> Schaeff.</td><td align="left" rowspan="1" colspan="1">Pileus moderately dark to dark yellow-brown or reddish brown, dry, tomentose, with age often somewhat fissurate or craqued; stipe with rather pronounced reticulation all over; pileipellis elements up to 20 μm wide with fine intracellular grains</td><td align="left" rowspan="1" colspan="1">With <italic>Quercus</italic>, on clay or sand mixed loam, on fairly dry places, such as road-sides and parks</td></tr><tr><td align="left" rowspan="1" colspan="1">Morphotype A ‘Fagus’.</td><td align="left" rowspan="1" colspan="1">Pileus very dark brown, dry; pruinose-tomentose; stipe strongly swollen and reticulate all over; pores long covered with a white layer</td><td align="left" rowspan="1" colspan="1">With <italic>Fagus</italic> on rich, humose soil</td></tr><tr><td align="left" rowspan="1" colspan="1">Morphotaxon B ‘Tilia’</td><td align="left" rowspan="1" colspan="1">Pileus surface glabrous, greasy to touch, vivid red-brown; stipe with a strong white net all over on reddish brown background; terminal elements of pileipellis tapering towards apex; 12–50 × 4.5–15 μm</td><td align="left" rowspan="1" colspan="1">With <italic>Tilia</italic> in road-sides on peaty soil</td></tr><tr><td align="left" rowspan="1" colspan="1">Morphotaxon C ‘Pinus’</td><td align="left" rowspan="1" colspan="1">Pileus grey-brown to reddish brown; glabrous, but not viscid; terminal elements of pileipellis slightly swollen, up to 15 μm wide</td><td align="left" rowspan="1" colspan="1">With <italic>Pinus</italic> on poor, acid, sandy soil</td></tr></tbody></table></table-wrap><table-wrap id="T3" orientation="portrait" position="float"><label>Table 3</label><caption><p>Collections used in this research project, showing collection number, GenBank accession number (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov">http://www.ncbi.nlm.nih.gov</ext-link>), country of origin and presumed host. Country names are abbreviated; A = Austria, B = Belgium, FIN = Finland, NL = The Netherlands, S = Sweden.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Sample No.</th><th align="left" rowspan="1" colspan="1">Collection No.</th><th align="left" rowspan="1" colspan="1">Species</th><th align="left" rowspan="1" colspan="1">GenBank accession No. (ITS/GAPDH)</th><th align="left" rowspan="1" colspan="1">Country</th><th align="left" rowspan="1" colspan="1">Host genus</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">SvdL49</td><td align="left" rowspan="1" colspan="1"><italic>B. aereus</italic></td><td align="left" rowspan="1" colspan="1">EU417844/EU417876</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">SvdL94</td><td align="left" rowspan="1" colspan="1"><italic>B. aereus</italic></td><td align="left" rowspan="1" colspan="1">EU417845/EU417877</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">SvdL24</td><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic></td><td align="left" rowspan="1" colspan="1">EU417846/EU417896</td><td align="left" rowspan="1" colspan="1">FIN</td><td align="left" rowspan="1" colspan="1"><italic>Betula</italic> or <italic>Picea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">SvdL58</td><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic></td><td align="left" rowspan="1" colspan="1">EU417847/EU417895</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">SvdL74</td><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic></td><td align="left" rowspan="1" colspan="1">EU417848/EU417894</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic> or <italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">SvdL36</td><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic></td><td align="left" rowspan="1" colspan="1">EU417849/EU417893</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">SvdL41</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> A</td><td align="left" rowspan="1" colspan="1">EU417850/EU417879</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Fagus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">SvdL32</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> A</td><td align="left" rowspan="1" colspan="1">EU417851/EU417880</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Fagus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">SvdL21</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417852/EU417881</td><td align="left" rowspan="1" colspan="1">FIN</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1">SvdL14</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417853/EU417882</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic> or <italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">11</td><td align="left" rowspan="1" colspan="1">SvdL16</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417854/EU417883</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">SvdL9i</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417855/EU417884</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic> or <italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">13</td><td align="left" rowspan="1" colspan="1">SvdL69</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417856/EU417885</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">14</td><td align="left" rowspan="1" colspan="1">SvdL61</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417857/EU417886</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">15</td><td align="left" rowspan="1" colspan="1">SvdL43</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417858/EU417888</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">16</td><td align="left" rowspan="1" colspan="1">SvdL45</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417859/EU417889</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">17</td><td align="left" rowspan="1" colspan="1">SvdL35</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> B</td><td align="left" rowspan="1" colspan="1">EU417860/EU417890</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Tilia</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">18</td><td align="left" rowspan="1" colspan="1">SvdL67</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> B</td><td align="left" rowspan="1" colspan="1">EU417861/EU417891</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Tilia</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">19</td><td align="left" rowspan="1" colspan="1">O1101601</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic> var. <italic>albus</italic></td><td align="left" rowspan="1" colspan="1">EU417862/EU417892</td><td align="left" rowspan="1" colspan="1">B</td><td align="left" rowspan="1" colspan="1"><italic>Picea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">20</td><td align="left" rowspan="1" colspan="1">SvdL50</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> C</td><td align="left" rowspan="1" colspan="1">EU417863/EU417905</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">21</td><td align="left" rowspan="1" colspan="1">SvdL93</td><td align="left" rowspan="1" colspan="1"><italic>Morphotaxon</italic> C</td><td align="left" rowspan="1" colspan="1">EU417864/EU417904</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">SvdL12</td><td align="left" rowspan="1" colspan="1"><italic>B. pinophilus</italic></td><td align="left" rowspan="1" colspan="1">EU417865/EU417903</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">23</td><td align="left" rowspan="1" colspan="1">SvdL18</td><td align="left" rowspan="1" colspan="1"><italic>B. pinophilus</italic></td><td align="left" rowspan="1" colspan="1">EU417866/EU417902</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Pinus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">24</td><td align="left" rowspan="1" colspan="1">O1097</td><td align="left" rowspan="1" colspan="1"><italic>B. quercicola</italic></td><td align="left" rowspan="1" colspan="1">EU417867/EU417901</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">25</td><td align="left" rowspan="1" colspan="1">O1095</td><td align="left" rowspan="1" colspan="1"><italic>B. quercicola</italic></td><td align="left" rowspan="1" colspan="1">EU417868/EU417900</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">26</td><td align="left" rowspan="1" colspan="1">O1085</td><td align="left" rowspan="1" colspan="1"><italic>B. quercicola</italic></td><td align="left" rowspan="1" colspan="1">EU417869/EU417899</td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic> or <italic>Castanea</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">27</td><td align="left" rowspan="1" colspan="1">SvdL47</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417870/EU417898</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">SvdL79</td><td align="left" rowspan="1" colspan="1"><italic>B. reticulates</italic></td><td align="left" rowspan="1" colspan="1">EU417871/EU417878</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">29</td><td align="left" rowspan="1" colspan="1">MEN 9336</td><td align="left" rowspan="1" colspan="1"><italic>B. fechtneri</italic></td><td align="left" rowspan="1" colspan="1">EU417872/EU417875</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">SvdL15</td><td align="left" rowspan="1" colspan="1"><italic>B. edulis</italic></td><td align="left" rowspan="1" colspan="1">EU417873/EU417887</td><td align="left" rowspan="1" colspan="1">S</td><td align="left" rowspan="1" colspan="1"><italic>Betula</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">31</td><td align="left" rowspan="1" colspan="1">SvdL63</td><td align="left" rowspan="1" colspan="1"><italic>B. betulicola</italic></td><td align="left" rowspan="1" colspan="1">EU417874/EU417897</td><td align="left" rowspan="1" colspan="1">NL</td><td align="left" rowspan="1" colspan="1"><italic>Quercus</italic> or <italic>Betula</italic></td></tr></tbody></table></table-wrap><fig id="F1" orientation="portrait" position="float"><label>Fig. 1</label><caption><p>Maximum likelihood tree from the combined ITS and GAPDH data. <italic>Bolletus fechtneri</italic> is selected as the outgroup. Above the branches bootstrap support values are displayed based on maximum parsimony (percentage), distance (percentage) and posterior probability (frequency) respectively. Behind the name and number of the samples the collection location and host genus are shown. NL = The Netherlands, FIN = Finland, S = Sweden, B = Belgium, A = Austria. ■ = <italic>Picea</italic>, ♦ = <italic>Pinus</italic>, * = <italic>Quercus</italic>, ○ = <italic>Betula</italic>, ▵ = <italic>Fagus</italic>, ▿ = <italic>Tilia</italic>, + = <italic>Castanea</italic>.</p></caption><graphic xlink:href="per-20-1-g001"></graphic></fig><fig id="F2" orientation="portrait" position="float"><label>Fig. 2</label><caption><p>A strict consesus MP topology of over >10 000 trees. <italic>B. fechtneri</italic> and <italic>B. luridus</italic> form the outgroup. Numerals associated with branches show BP support (percentage) followed by posterior probability (frequency). <xref ref-type="bibr" rid="R12">Leonardi et al. (2005)</xref> sequences preceded by their GenBank Accession numbers and country of origin. Samples from this study followed by sample number (<xref ref-type="table" rid="T1">Table 1</xref>) and country of origin. A = Austria, B = Belgium, F = France, FIN = Finland, NL = The Netherlands, I = Italy, S = Sweden.</p></caption><graphic xlink:href="per-20-1-g002"></graphic></fig></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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