Phylogeny of Penicillium and the segregation of Trichocomaceae into three families
Identifieur interne : 000246 ( Pmc/Corpus ); précédent : 000245; suivant : 000247Phylogeny of Penicillium and the segregation of Trichocomaceae into three families
Auteurs : J. Houbraken ; R. A. SamsonSource :
- Studies in Mycology [ 0166-0616 ] ; 2011.
Abstract
Species of
Url:
DOI: 10.3114/sim.2011.70.01
PubMed: 22308045
PubMed Central: 3233907
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Phylogeny of <italic>Penicillium</italic> and the segregation of <italic>Trichocomaceae</italic> into three families</title><author><name sortKey="Houbraken, J" sort="Houbraken, J" uniqKey="Houbraken J" first="J." last="Houbraken">J. Houbraken</name><affiliation><nlm:aff id="A1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="A2"><italic>Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.</italic></nlm:aff></affiliation></author><author><name sortKey="Samson, R A" sort="Samson, R A" uniqKey="Samson R" first="R. A." last="Samson">R. A. Samson</name><affiliation><nlm:aff id="A1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">22308045</idno><idno type="pmc">3233907</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3233907</idno><idno type="RBID">PMC:3233907</idno><idno type="doi">10.3114/sim.2011.70.01</idno><date when="2011">2011</date><idno type="wicri:Area/Pmc/Corpus">000246</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000246</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Phylogeny of <italic>Penicillium</italic> and the segregation of <italic>Trichocomaceae</italic> into three families</title><author><name sortKey="Houbraken, J" sort="Houbraken, J" uniqKey="Houbraken J" first="J." last="Houbraken">J. Houbraken</name><affiliation><nlm:aff id="A1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation><affiliation><nlm:aff id="A2"><italic>Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.</italic></nlm:aff></affiliation></author><author><name sortKey="Samson, R A" sort="Samson, R A" uniqKey="Samson R" first="R. A." last="Samson">R. A. Samson</name><affiliation><nlm:aff id="A1"><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></nlm:aff></affiliation></author></analytic><series><title level="j">Studies in Mycology</title><idno type="ISSN">0166-0616</idno><idno type="eISSN">1872-9797</idno><imprint><date when="2011">2011</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p id="P6">Species of <italic>Trichocomaceae</italic> occur commonly and are important to both industry and medicine. They are associated with food spoilage and mycotoxin production and can occur in the indoor environment, causing health hazards by the formation of β-glucans, mycotoxins and surface proteins. Some species are opportunistic pathogens, while others are exploited in biotechnology for the production of enzymes, antibiotics and other products. <italic>Penicillium</italic> belongs phylogenetically to <italic>Trichocomaceae</italic> and more than 250 species are currently accepted in this genus. In this study, we investigated the relationship of <italic>Penicillium</italic> to other genera of <italic>Trichocomaceae</italic> and studied in detail the phylogeny of the genus itself. In order to study these relationships, partial <italic>RPB1</italic>, <italic>RPB2</italic> (RNA polymerase II genes), <italic>Tsr1</italic> (putative ribosome biogenesis protein) and <italic>Cct8</italic> (putative chaperonin complex component TCP-1) gene sequences were obtained. The <italic>Trichocomaceae</italic> are divided in three separate families: <italic>Aspergillaceae</italic>, <italic>Thermoascaceae</italic> and <italic>Trichocomaceae</italic>. The <italic>Aspergillaceae</italic> are characterised by the formation flask-shaped or cylindrical phialides, asci produced inside cleistothecia or surrounded by Hülle cells and mainly ascospores with a furrow or slit, while the <italic>Trichocomaceae</italic> are defined by the formation of lanceolate phialides, asci borne within a tuft or layer of loose hyphae and ascospores lacking a slit. <italic>Thermoascus</italic> and <italic>Paecilomyces</italic>, both members of <italic>Thermoascaceae</italic>, also form ascospores lacking a furrow or slit, but are differentiated from <italic>Trichocomaceae</italic> by the production of asci from croziers and their thermotolerant or thermophilic nature. Phylogenetic analysis shows that <italic>Penicillium</italic> is polyphyletic. The genus is re-defined and a monophyletic genus for both anamorphs and teleomorphs is created (<italic>Penicillium sensu stricto</italic>). The genera <italic>Thysanophora</italic>, <italic>Eupenicillium</italic>, <italic>Chromocleista</italic>, <italic>Hemicarpenteles</italic> and <italic>Torulomyces</italic> belong in <italic>Penicillium</italic> <italic>s</italic>. <italic>str</italic>. and new combinations for the species belonging to these genera are proposed. Analysis of <italic>Penicillium</italic> below genus rank revealed the presence of 25 clades. A new classification system including both anamorph and teleomorph species is proposed and these 25 clades are treated here as sections. An overview of species belonging to each section is presented.</p><sec id="S1"><title>Taxonomic novelties:</title><p id="P7"><bold>New sections, all in <italic>Penicillium:</italic></bold> sect. Sclerotiora Houbraken & Samson, sect. <italic>Charlesia</italic> Houbraken & Samson, sect. <italic>Thysanophora</italic> Houbraken & Samson,sect. <italic>Ochrosalmonea</italic> Houbraken & Samson, sect. <italic>Cinnamopurpurea</italic> Houbraken & Samson, <italic>Fracta</italic> Houbraken & Samson, sect. <italic>Stolkia</italic> Houbraken & Samson, sect. <italic>Gracilenta</italic> Houbraken & Samson, sect. <italic>Citrina</italic> Houbraken & Samson, sect. <italic>Turbata</italic> Houbraken & Samson, sect. <italic>Paradoxa</italic> Houbraken & Samson, sect. <italic>Canescentia</italic> Houbraken & Samson.</p><p id="P8"><bold>New combinations:</bold> <italic>Penicillium asymmetricum</italic> (Subramanian & Sudha) Houbraken & Samson, <italic>P. bovifimosum</italic> (Tuthill & Frisvad) Houbraken & Samson, <italic>P. glaucoalbidum</italic> (Desmazières) Houbraken & Samson, <italic>P. laeve</italic> (K. Ando & Manoch) Houbraken & Samson, <italic>P. longisporum</italic> (Kendrick) Houbraken & Samson, <italic>P. malachiteum</italic> (Yaguchi & Udagawa) Houbraken & Samson, <italic>P. ovatum</italic> (K. Ando & Nawawi) Houbraken & Samson, <italic>P. parviverrucosum</italic> (K. Ando & Pitt) Houbraken & Samson, <italic>P. saturniforme</italic> (Wang & Zhuang) Houbraken & Samson, <italic>P. taiwanense</italic> (Matsushima) Houbraken & Samson.</p><p id="P9"><bold>New names:</bold> <italic>Penicillium coniferophilum</italic> Houbraken & Samson, <italic>P. hennebertii</italic> Houbraken & Samson, <italic>P. melanostipe</italic> Houbraken & Samson, <italic>P. porphyreum</italic> Houbraken & Samson.</p></sec></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Abe, S" uniqKey="Abe S">S Abe</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Aguileta, G" uniqKey="Aguileta G">G Aguileta</name></author><author><name sortKey="Marthey, S" uniqKey="Marthey S">S Marthey</name></author><author><name sortKey="Chiapello, H" uniqKey="Chiapello H">H Chiapello</name></author><author><name sortKey="Lebrun, Mh" uniqKey="Lebrun M">MH Lebrun</name></author><author><name sortKey="Rodolphe, F" 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Stud Mycol</journal-id><journal-id journal-id-type="hwp">simycol</journal-id><journal-title-group><journal-title>Studies in Mycology</journal-title></journal-title-group><issn pub-type="ppub">0166-0616</issn><issn pub-type="epub">1872-9797</issn><publisher><publisher-name>CBS Fungal Biodiversity Centre</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">22308045</article-id><article-id pub-id-type="pmc">3233907</article-id><article-id pub-id-type="doi">10.3114/sim.2011.70.01</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>Phylogeny of <italic>Penicillium</italic> and the segregation of <italic>Trichocomaceae</italic> into three families</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Houbraken</surname><given-names>J.</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Samson</surname><given-names>R.A.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib></contrib-group><aff id="A1"><label>1</label><italic>CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands</italic></aff><aff id="A2"><label>2</label><italic>Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.</italic></aff><author-notes><corresp>*<italic>Correspondence</italic>: Jos Houbraken, <email>j.houbraken@cbs.knaw.nl</email></corresp></author-notes><pub-date pub-type="ppub"><day>15</day><month>11</month><year>2011</year></pub-date><volume>70</volume><issue>1</issue><issue-title>Phylogenetic and taxonomic studies on the genera Penicillium and Talaromyces</issue-title><fpage>1</fpage><lpage>51</lpage><permissions><copyright-statement>Copyright 2011 CBS-KNAW Fungal Biodiversity Centre</copyright-statement><copyright-year>2011</copyright-year><license license-type="creativecommons"><license-p>You are free to share - to copy, distribute and transmit the work, under the following conditions:</license-p><license-p><bold>Attribution:</bold>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><bold>Non-commercial:</bold> You may not use this work for commercial purposes.</license-p><license-p><bold>No derivative works:</bold> You may not alter, transform, or build upon this work.</license-p><license-p>For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at <uri xlink:type="simple" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode">http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode</uri>. 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><self-uri xlink:title="pdf" xlink:type="simple" xlink:href="1.pdf"></self-uri><abstract><p id="P6">Species of <italic>Trichocomaceae</italic> occur commonly and are important to both industry and medicine. They are associated with food spoilage and mycotoxin production and can occur in the indoor environment, causing health hazards by the formation of β-glucans, mycotoxins and surface proteins. Some species are opportunistic pathogens, while others are exploited in biotechnology for the production of enzymes, antibiotics and other products. <italic>Penicillium</italic> belongs phylogenetically to <italic>Trichocomaceae</italic> and more than 250 species are currently accepted in this genus. In this study, we investigated the relationship of <italic>Penicillium</italic> to other genera of <italic>Trichocomaceae</italic> and studied in detail the phylogeny of the genus itself. In order to study these relationships, partial <italic>RPB1</italic>, <italic>RPB2</italic> (RNA polymerase II genes), <italic>Tsr1</italic> (putative ribosome biogenesis protein) and <italic>Cct8</italic> (putative chaperonin complex component TCP-1) gene sequences were obtained. The <italic>Trichocomaceae</italic> are divided in three separate families: <italic>Aspergillaceae</italic>, <italic>Thermoascaceae</italic> and <italic>Trichocomaceae</italic>. The <italic>Aspergillaceae</italic> are characterised by the formation flask-shaped or cylindrical phialides, asci produced inside cleistothecia or surrounded by Hülle cells and mainly ascospores with a furrow or slit, while the <italic>Trichocomaceae</italic> are defined by the formation of lanceolate phialides, asci borne within a tuft or layer of loose hyphae and ascospores lacking a slit. <italic>Thermoascus</italic> and <italic>Paecilomyces</italic>, both members of <italic>Thermoascaceae</italic>, also form ascospores lacking a furrow or slit, but are differentiated from <italic>Trichocomaceae</italic> by the production of asci from croziers and their thermotolerant or thermophilic nature. Phylogenetic analysis shows that <italic>Penicillium</italic> is polyphyletic. The genus is re-defined and a monophyletic genus for both anamorphs and teleomorphs is created (<italic>Penicillium sensu stricto</italic>). The genera <italic>Thysanophora</italic>, <italic>Eupenicillium</italic>, <italic>Chromocleista</italic>, <italic>Hemicarpenteles</italic> and <italic>Torulomyces</italic> belong in <italic>Penicillium</italic> <italic>s</italic>. <italic>str</italic>. and new combinations for the species belonging to these genera are proposed. Analysis of <italic>Penicillium</italic> below genus rank revealed the presence of 25 clades. A new classification system including both anamorph and teleomorph species is proposed and these 25 clades are treated here as sections. An overview of species belonging to each section is presented.</p><sec id="S1"><title>Taxonomic novelties:</title><p id="P7"><bold>New sections, all in <italic>Penicillium:</italic></bold> sect. Sclerotiora Houbraken & Samson, sect. <italic>Charlesia</italic> Houbraken & Samson, sect. <italic>Thysanophora</italic> Houbraken & Samson,sect. <italic>Ochrosalmonea</italic> Houbraken & Samson, sect. <italic>Cinnamopurpurea</italic> Houbraken & Samson, <italic>Fracta</italic> Houbraken & Samson, sect. <italic>Stolkia</italic> Houbraken & Samson, sect. <italic>Gracilenta</italic> Houbraken & Samson, sect. <italic>Citrina</italic> Houbraken & Samson, sect. <italic>Turbata</italic> Houbraken & Samson, sect. <italic>Paradoxa</italic> Houbraken & Samson, sect. <italic>Canescentia</italic> Houbraken & Samson.</p><p id="P8"><bold>New combinations:</bold> <italic>Penicillium asymmetricum</italic> (Subramanian & Sudha) Houbraken & Samson, <italic>P. bovifimosum</italic> (Tuthill & Frisvad) Houbraken & Samson, <italic>P. glaucoalbidum</italic> (Desmazières) Houbraken & Samson, <italic>P. laeve</italic> (K. Ando & Manoch) Houbraken & Samson, <italic>P. longisporum</italic> (Kendrick) Houbraken & Samson, <italic>P. malachiteum</italic> (Yaguchi & Udagawa) Houbraken & Samson, <italic>P. ovatum</italic> (K. Ando & Nawawi) Houbraken & Samson, <italic>P. parviverrucosum</italic> (K. Ando & Pitt) Houbraken & Samson, <italic>P. saturniforme</italic> (Wang & Zhuang) Houbraken & Samson, <italic>P. taiwanense</italic> (Matsushima) Houbraken & Samson.</p><p id="P9"><bold>New names:</bold> <italic>Penicillium coniferophilum</italic> Houbraken & Samson, <italic>P. hennebertii</italic> Houbraken & Samson, <italic>P. melanostipe</italic> Houbraken & Samson, <italic>P. porphyreum</italic> Houbraken & Samson.</p></sec></abstract><kwd-group><title>Keywords:</title><kwd><italic>Aspergillus</italic></kwd><kwd><italic>Eupenicillium</italic></kwd><kwd>nomenclature</kwd><kwd><italic>Penicillium</italic></kwd><kwd><italic>Talaromyces</italic></kwd><kwd>taxonomy.</kwd></kwd-group></article-meta></front><body><sec id="S2"><title>INTRODUCTION</title><p id="P10">The <italic>Trichocomaceae</italic> comprise a relatively large family of fungi well-known for their impact, both positive and negative, on human activities. The most well-known species of this family belong to the genera <italic>Aspergillus, Penicillium</italic> and <italic>Paecilomyces</italic>. Species belonging to <italic>Trichocomaceae</italic> are predominantly saprobic and represent some of the most catabolically and anabolically diverse microorganisms known. Some species are capable of growing at extremely low water activities (<italic>i.e.</italic> xerotolerant and/or osmotolerant), low temperatures (psychrotolerant) and high temperatures (thermotolerant). Members of <italic>Trichocomaceae</italic> secrete secondary metabolites (extrolites) that are known as mycotoxins (<italic>e.g.</italic> aflatoxins, ochratoxins, patulin), while other extrolites are used as pharmaceuticals, including antibiotics such as penicillin and the cholesterol-lowering agent lovastatin. Furthermore, members of <italic>Trichocomaceae</italic> are also known for their production of organic acids and diverse enzymes that degrade a wide variety of complex biomolecules (<xref ref-type="bibr" rid="R45">Geiser <italic>et al</italic>. 2006</xref>, <xref ref-type="bibr" rid="R128">Pitt & Hocking 2009</xref>, <xref ref-type="bibr" rid="R144">Samson <italic>et al</italic>. 2010</xref>).</p><p id="P11">The taxon <italic>Trichocomaceae</italic> was introduced by Fischer (<xref ref-type="bibr" rid="R30">1897</xref>) and the classification of this family was studied extensively using phenotypic characters (<xref ref-type="bibr" rid="R92">Malloch & Cain 1972</xref>, <xref ref-type="bibr" rid="R169">Subramanian 1972</xref>, <xref ref-type="bibr" rid="R90">Malloch 1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>, von Arx1986). These studies include only teleomorph genera because <italic>Trichocomaceae</italic> is based on <italic>Trichocoma</italic>, a teleomorph genus, and thus not applicable for anamorph genera (<xref ref-type="bibr" rid="R91">Malloch 1985b</xref>). However, it is noted that anamorph genera with phialidic structures are linked to <italic>Trichocomaceae</italic> (<xref ref-type="bibr" rid="R92">Malloch & Cain 1972</xref>). Currently, only the phylogenetic relationships within certain genera of <italic>Trichocomaceae, e.g. Aspergillus, Penicillium</italic> and <italic>Paecilomyces</italic>, are elucidated (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, <xref ref-type="bibr" rid="R113">b</xref>, <xref ref-type="bibr" rid="R147">Samson <italic>et al</italic>. 2004</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>, <xref ref-type="bibr" rid="R145">Samson <italic>et al</italic>. 2009</xref>), but the relationships among the genera are still poorly studied.</p><p id="P12"><italic>Penicillium</italic> is an anamorph genus and belongs phylogenetically to <italic>Trichocomaceae</italic> (<xref ref-type="bibr" rid="R15">Berbee 1995</xref>, <xref ref-type="bibr" rid="R112">Peterson 2000a</xref>). The name <italic>Penicillium</italic> is derived from <italic>penicillus</italic>, which means “little brush” and was introduced by <xref ref-type="bibr" rid="R80">Link in 1809</xref>. Many new species were described in the 19<sup>th</sup> century, and Dierckx (<xref ref-type="bibr" rid="R25">1901</xref>) was the first researcher who introduced a subgeneric classification system for the genus.</p><p id="P13">He proposed the subgenera <italic>Aspergilloides, Biverticillium</italic> and <italic>Eupenicillium</italic> and Biourge (<xref ref-type="bibr" rid="R18">1923</xref>) followed Dierckx's classification system and expanded it with two sections, four series and six subsections. Thom (<xref ref-type="bibr" rid="R178">1930</xref>: 155–159) did not accept Dierckx's and Biourge's subgeneric classification system and introduced a new system with four divisions (subgenera), 12 sections and 18 subsections (series). His system was mainly based on colony characteristics and conidiophore branching and the monographs of Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>) and Ramírez (<xref ref-type="bibr" rid="R132">1982</xref>) are in line with that of Thom (<xref ref-type="bibr" rid="R178">1930</xref>). Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) did not follow Thom's concept and, based on conidiophore characters, phialide shapes and growth characteristics, divided <italic>Penicillium</italic> into four subgenera, 10 sections and 21 series. In addition, he treated <italic>Eupenicillium</italic> separately from <italic>Penicillium</italic> and subdivided the former genus into eight series. In 1985, Stolk & Samson proposed another taxonomic scheme for <italic>Penicillium</italic> anamorphs and this classification was primary based on phialide shape and conidiophore branching. They divided <italic>Penicillium</italic> in 10 sections and 18 series and this taxonomic scheme treated strict anamorphs, as well as anamorphs of sexual <italic>Penicillium</italic> species. More recently, Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) studied subgenus <italic>Penicillium</italic> and five sections and 17 series were recognised.</p><p id="P14">The first attempt to make a subgeneric classification of <italic>Eupenicillium</italic> was undertaken by Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) and eight series were introduced. This classification was based on a combination of various characters, such as growth rates in standard conditions, colony morphology and microscopical characters of both teleomorphic and anamorphic states. In the monograph of Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>), four sections were introduced for the classification of <italic>Eupenicillium</italic>, and Pitt's concept of using series of species was abandoned.</p><p id="P15">To date, only a limited number of studies have investigated the phylogenetic relationship of <italic>Penicillium</italic> at genus level. Berbee (<xref ref-type="bibr" rid="R15">1995</xref>), based of 18S rDNA sequences, demonstrated that <italic>Penicillium</italic> is polyphyletic. The genus splits up in two clades: one clade includes <italic>Talaromyces</italic> species and members of the subgenus <italic>Biverticillium</italic> and the other clade includes <italic>Eupenicillium</italic> species and <italic>Penicillium</italic> species accommodated in the subgenera <italic>Penicillium, Furcatum</italic> and <italic>Aspergilloides</italic> (<xref ref-type="bibr" rid="R83">LoBuglio & Taylor 1993</xref>, <xref ref-type="bibr" rid="R84">LoBuglio <italic>et al</italic>. 1993</xref>, <xref ref-type="bibr" rid="R15">Berbee <italic>et al</italic>. 1995</xref>, <xref ref-type="bibr" rid="R106">Ogawa <italic>et al</italic>. 1997</xref>, <xref ref-type="bibr" rid="R192">Wang & Zhuang 2007</xref>). Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>) studied the phylogeny of <italic>Eupenicillium</italic> and members of the subgenera <italic>Penicillium, Furcatum</italic> and <italic>Aspergilloides</italic> in more detail. He subsequently divided the studied species in six groups and showed that many subgeneric taxa in <italic>Penicillium</italic> are polyphyletic. Furthermore, his data indicated that the current classification systems based on conidiophore branching is not congruent with the phylogeny and a new subgeneric classification system is needed.</p><p id="P16">Pleomorphism in fungi was first demonstrated by Tulasne (<xref ref-type="bibr" rid="R179">1851</xref>). Together with his discovery, he was already aware of the problem raised by the nomenclature of composite species and he stated that the imperfect forms must someday be submerged in the Ascomycota. He thus established a first principle of pleomorphic nomenclature and suggested the precedence of the perfect state name over imperfect names (<xref ref-type="bibr" rid="R53">Hennebert 1971</xref>). In 1910, “dual nomenclature” was introduced and this was established in the International Code of Botanical Nomenclature (ICBN). The problem of naming fungi that exhibit pleomorphic life cycles was addressed in previous versions of article 59 of the ICBN and implied that more than one name for a single taxon can be used (<xref ref-type="bibr" rid="R22">Cline 2005</xref>). Recently, the proposal to revise article 59 was accepted at the 2011 IBC Nomenclature Section at Melbourne and the principle of “one fungus: one name” was established (<xref ref-type="bibr" rid="R104">Norvell <italic>et al</italic>. 2011</xref>).</p><p id="P17">In the present study, the phylogenetic relationships between <italic>Penicillium</italic> and other members of the family <italic>Trichocomaceae</italic> are studied using a combined analysis of four loci (<italic>RPB1, RPB2, Tsr1</italic> and <italic>Cct8</italic>). In this study, the principle “one fungus- one name” is applied and priority is given to the oldest family, genus and section names using the single-name nomenclature (<xref ref-type="bibr" rid="R50">Hawksworth <italic>et al</italic>. 2011</xref>, <xref ref-type="bibr" rid="R104">Norvell 2011</xref>). <italic>Penicillium</italic> is delimited, various genera are placed in synonymy, and new combinations in <italic>Penicillium</italic> are made for the species belonging to the genera <italic>Thysanophora, Eupenicillium, Chromocleista, Hemicarpenteles</italic> and <italic>Torulomyces</italic>. Subsequently, the phylogeny of <italic>Penicillium</italic> is studied and a new sectional classification system is proposed. In addition, an overview of species in each section is presented.</p></sec><sec sec-type="materials|methods" id="S3"><title>MATERIAL AND METHODS</title><sec id="S4"><title>Strains</title><p id="P18">The first part of this study treats the phylogenetic relationships of the <italic>Penicillium</italic> species among <italic>Trichocomaceae</italic>. A selection of strains is made in order to study these relationships and in most cases the types of the genera were selected. The second part deals with the phylogeny of <italic>Penicillium.</italic> For this study, the type species of the various subgenera and sections in <italic>Penicillium</italic> and <italic>Eupenicillium</italic> were selected, and this selection is supplemented with other related species. An overview of strains used in the study of the phylogeny of <italic>Trichocomaceae</italic> and <italic>Penicillium</italic> presented in <xref ref-type="table" rid="T1">Table 1</xref>. In the third part of this study, a new sectional classification system for <italic>Penicillium</italic> is proposed and lists of species in each section are compiled. For the preparation of these lists, mostly type strains were selected of accepted <italic>Penicillium</italic> and <italic>Eupenicillium</italic> species. This selection is based on the overview of “accepted species and their synonyms in the <italic>Trichocomaceae</italic>” by Pitt <italic>et al</italic>. (<xref ref-type="bibr" rid="R130">2000</xref>) and supplemented with species described after 2000. An overview of these strains is shown in Table S1 (Supplementary Information- online only) and partly in <xref ref-type="table" rid="T1">Table 1</xref> (species names indicated with two asterisks). All strains are maintained in the CBS-KNAW culture collection and additional strains were obtained from IBT (culture collection of Center for Microbial Biotechnology (CMB) at Department of Systems Biology, Technical University of Denmark), NRRL (ARS Culture Collection, U.S. Department of Agriculture, Peoria, Illinois, USA), ATCC (American Type Culture Collection, Manassas, VA, USA) and IMI (CABI Genetic Resources Collection, Surrey, UK).</p><table-wrap id="T1" position="float"><label>Table 1</label><caption><p>Strains used in phylogenetic analysis of <italic>Trichocomaceae</italic> and other families.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top" rowspan="1" colspan="1">CBS no.</th><th align="left" valign="top" rowspan="1" colspan="1">Name</th><th align="left" valign="top" rowspan="1" colspan="1">Other collections</th><th align="left" valign="top" rowspan="1" colspan="1">Origin</th><th colspan="4" align="center" valign="top" rowspan="1">GenBank accession or reference<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></th></tr><tr><th align="left" valign="top" rowspan="1" colspan="1"></th><th align="left" valign="top" rowspan="1" colspan="1"></th><th align="left" valign="top" rowspan="1" colspan="1"></th><th align="left" valign="top" rowspan="1" colspan="1"></th><th align="left" valign="top" rowspan="1" colspan="1"><italic>RPB1</italic></th><th align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></th><th align="left" valign="top" rowspan="1" colspan="1"><italic>Tsr1</italic></th><th align="left" valign="top" rowspan="1" colspan="1"><italic>Cct8</italic></th></tr></thead><tbody><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=267.72&link_type=cbs">CBS 267.72</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aphanoascus cinnabarinus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 26215</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121625</td><td align="left" valign="top" rowspan="1" colspan="1">JN121477</td><td align="left" valign="top" rowspan="1" colspan="1">JN121783</td><td align="left" valign="top" rowspan="1" colspan="1">JN121903</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=172.66&link_type=cbs">CBS 172.66</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus aculeatus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16872 = IMI 211388</td><td align="left" valign="top" rowspan="1" colspan="1">Tropical soil</td><td align="left" valign="top" rowspan="1" colspan="1">JN121590</td><td align="left" valign="top" rowspan="1" colspan="1">JN121448</td><td align="left" valign="top" rowspan="1" colspan="1">JN121755</td><td align="left" valign="top" rowspan="1" colspan="1">JN121895</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=600.67&link_type=cbs">CBS 600.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus amylovorus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18351 = IMI 129961 = MUCL 15648</td><td align="left" valign="top" rowspan="1" colspan="1">Wheat starch, Kharkiv, Ukraine</td><td align="left" valign="top" rowspan="1" colspan="1">JN121705</td><td align="left" valign="top" rowspan="1" colspan="1">JN121538</td><td align="left" valign="top" rowspan="1" colspan="1">JN121844</td><td align="left" valign="top" rowspan="1" colspan="1">JN121931</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=463.65&link_type=cbs">CBS 463.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus arenarius</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16830 = IMI 055632 = IMI 055632ii</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Mysore, Karnataka, India</td><td align="left" valign="top" rowspan="1" colspan="1">JN121684</td><td align="left" valign="top" rowspan="1" colspan="1">JN121520</td><td align="left" valign="top" rowspan="1" colspan="1">JN121825</td><td align="left" valign="top" rowspan="1" colspan="1">JN121917</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=653.74&link_type=cbs">CBS 653.74</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus aureofulgens</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Natural truffle soil, Provence, France</td><td align="left" valign="top" rowspan="1" colspan="1">JN121712</td><td align="left" valign="top" rowspan="1" colspan="1">JN121545</td><td align="left" valign="top" rowspan="1" colspan="1">JN121851</td><td align="left" valign="top" rowspan="1" colspan="1">JN121936</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.46&link_type=cbs">CBS 109.46</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus avenaceus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16861 = IMI 016140 = NRRL 517</td><td align="left" valign="top" rowspan="1" colspan="1">Seed of <italic>Pisum sativum</italic> (pea), England, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121565</td><td align="left" valign="top" rowspan="1" colspan="1">JN121424</td><td align="left" valign="top" rowspan="1" colspan="1">JN121731</td><td align="left" valign="top" rowspan="1" colspan="1">JN121878</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=468.65&link_type=cbs">CBS 468.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus biplanus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16858 = IMI 235602</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Tilaran, Costa Rica</td><td align="left" valign="top" rowspan="1" colspan="1">JN121685</td><td align="left" valign="top" rowspan="1" colspan="1">JN121520</td><td align="left" valign="top" rowspan="1" colspan="1">JN121826</td><td align="left" valign="top" rowspan="1" colspan="1">JN121917</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=707.71&link_type=cbs">CBS 707.71</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus bisporus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22527 = NRRL 3693</td><td align="left" valign="top" rowspan="1" colspan="1">Soil injected into mouse, Clarksburg, Maryland, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121715</td><td align="left" valign="top" rowspan="1" colspan="1">JN121548</td><td align="left" valign="top" rowspan="1" colspan="1">JN121854</td><td align="left" valign="top" rowspan="1" colspan="1">JN121939</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127.61&link_type=cbs">CBS 127.61</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus brunneouniseriatus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16916 = IMI 227677</td><td align="left" valign="top" rowspan="1" colspan="1">Soil under Dalbergia sissoo, India</td><td align="left" valign="top" rowspan="1" colspan="1">JN121583</td><td align="left" valign="top" rowspan="1" colspan="1">JN121442</td><td align="left" valign="top" rowspan="1" colspan="1">JN121749</td><td align="left" valign="top" rowspan="1" colspan="1">JN121889</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=121611&link_type=cbs">CBS 121611</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus calidoustus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Patient (case 4), man with allogeneic HSCT, probably lung infection, man, Washington, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121579</td><td align="left" valign="top" rowspan="1" colspan="1">JN121438</td><td align="left" valign="top" rowspan="1" colspan="1">JN121745</td><td align="left" valign="top" rowspan="1" colspan="1">JN121887</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=566.65&link_type=cbs">CBS 566.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus candidus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1002 = IMI 091889 = NRRL 303</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">JN121702</td><td align="left" valign="top" rowspan="1" colspan="1">JN121535</td><td align="left" valign="top" rowspan="1" colspan="1">JN121841</td><td align="left" valign="top" rowspan="1" colspan="1">JN121929</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=196.64&link_type=cbs">CBS 196.64</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus cervinus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 15508 = IMI 107684</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, West Malaysia, Malaysia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121595</td><td align="left" valign="top" rowspan="1" colspan="1">JN121452</td><td align="left" valign="top" rowspan="1" colspan="1">JN121759</td><td align="left" valign="top" rowspan="1" colspan="1">JN121896</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=473.65&link_type=cbs">CBS 473.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus clavatoflavus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16866 = IMI 124937</td><td align="left" valign="top" rowspan="1" colspan="1">Rain forest soil,Tulley, Queensland, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121686</td><td align="left" valign="top" rowspan="1" colspan="1">JN121521</td><td align="left" valign="top" rowspan="1" colspan="1">JN121827</td><td align="left" valign="top" rowspan="1" colspan="1">JN121918</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus clavatus<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NRRL 1 (= ATCC 1007 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=15949)&link_type=cbs">CBS 513.65</ext-link> = IMI 15949)</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">Fedorova <italic>et al.</italic> (<xref ref-type="bibr" rid="R28">2008</xref>)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=476.65&link_type=cbs">CBS 476.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus conjunctus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16796 = IMI 135421</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil, Palmar, Province of Puntarenas, Costa Rica</td><td align="left" valign="top" rowspan="1" colspan="1">JN121688</td><td align="left" valign="top" rowspan="1" colspan="1">JN121523</td><td align="left" valign="top" rowspan="1" colspan="1">JN121829</td><td align="left" valign="top" rowspan="1" colspan="1">JN121920</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=553.77&link_type=cbs">CBS 553.77</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus coremiiformis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 38576 = 223069</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Ivory Coast</td><td align="left" valign="top" rowspan="1" colspan="1">JN121700</td><td align="left" valign="top" rowspan="1" colspan="1">JN12153</td><td align="left" valign="top" rowspan="1" colspan="1">JN121839</td><td align="left" valign="top" rowspan="1" colspan="1">JN121926</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=656.73&link_type=cbs">CBS 656.73</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus egyptiacus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 141415</td><td align="left" valign="top" rowspan="1" colspan="1">Sandy soil, under <italic>Olea europaea</italic> (olive tree), Mediterranean Coast, Ras-el-Hikma, Egypt</td><td align="left" valign="top" rowspan="1" colspan="1">JN121713</td><td align="left" valign="top" rowspan="1" colspan="1">JN121546</td><td align="left" valign="top" rowspan="1" colspan="1">JN121852</td><td align="left" valign="top" rowspan="1" colspan="1">JN121937</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=128202&link_type=cbs">CBS 128202</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus flavus<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NRRL 3357 (= ATCC 200026)</td><td align="left" valign="top" rowspan="1" colspan="1">Peanut cotyledons, USA</td><td align="left" valign="top" rowspan="1" colspan="1">Unpublished</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus fumigatus<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">Af293</td><td align="left" valign="top" rowspan="1" colspan="1">Patient with invasive aspergillosis</td><td align="left" valign="top" rowspan="1" colspan="1">Nierman <italic>et al.</italic> (<xref ref-type="bibr" rid="R102">2005</xref>)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116.56&link_type=cbs">CBS 116.56N</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus funiculosus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16846 = IMI 054397 = IMI 054397ii</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Ibadan, Nigeria</td><td align="left" valign="top" rowspan="1" colspan="1">JN121572</td><td align="left" valign="top" rowspan="1" colspan="1">JN121431</td><td align="left" valign="top" rowspan="1" colspan="1">JN121738</td><td align="left" valign="top" rowspan="1" colspan="1">JN121883</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=118.45&link_type=cbs">CBS 118.45</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus janus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16835 = IMI 016065 = IMI 016065ii = MUCL 31307 = NRRL 1787</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Panama</td><td align="left" valign="top" rowspan="1" colspan="1">JN121576</td><td align="left" valign="top" rowspan="1" colspan="1">JN121435</td><td align="left" valign="top" rowspan="1" colspan="1">JN121742</td><td align="left" valign="top" rowspan="1" colspan="1">JN121885</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=538.65&link_type=cbs">CBS 538.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus kanagawaensis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16143 = IMI 126690</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil under <italic>Pinus banksiana</italic>, Wisconsin, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121698</td><td align="left" valign="top" rowspan="1" colspan="1">JN121531</td><td align="left" valign="top" rowspan="1" colspan="1">JN121837</td><td align="left" valign="top" rowspan="1" colspan="1">JN121925</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=151.66&link_type=cbs">CBS 151.66</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus leporis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16490</td><td align="left" valign="top" rowspan="1" colspan="1">Dung of <italic>Lepus townsendii</italic> (white-tailed Jackrabbit ), near Saratoga, Wyoming, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121589</td><td align="left" valign="top" rowspan="1" colspan="1">JN121446</td><td align="left" valign="top" rowspan="1" colspan="1">JN121753</td><td align="left" valign="top" rowspan="1" colspan="1">JN121893</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=513.88&link_type=cbs">CBS 513.88</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus niger<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Derived from NRRL 3122 and currently used as enzyme production strain.</td><td align="left" valign="top" rowspan="1" colspan="1">Pel <italic>et al.</italic> (<xref ref-type="bibr" rid="R108">2007</xref>)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101887&link_type=cbs">CBS 101887</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus ochraceoroseus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 42001 = IBT 14580</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Tai National Forest, Ivory Coast</td><td align="left" valign="top" rowspan="1" colspan="1">JN121557</td><td align="left" valign="top" rowspan="1" colspan="1">JN121416</td><td align="left" valign="top" rowspan="1" colspan="1">JN121723</td><td align="left" valign="top" rowspan="1" colspan="1">JN121871</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=108.08&link_type=cbs">CBS 108.08</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus ochraceus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1008 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=547.65&link_type=cbs">CBS 547.65</ext-link> = IMI 016247 = IMI 016247iii = IMI 016247iv = NRRL 1642 = NRRL 398</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">JN121562</td><td align="left" valign="top" rowspan="1" colspan="1">JN121421</td><td align="left" valign="top" rowspan="1" colspan="1">JN121728</td><td align="left" valign="top" rowspan="1" colspan="1">JN121875</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=622.67&link_type=cbs">CBS 622.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus penicilliformis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18328 = IMI 129968= IMI 132431</td><td align="left" valign="top" rowspan="1" colspan="1">Soil under <italic>Nicotiana tabacum</italic>, Moldavia, Romania</td><td align="left" valign="top" rowspan="1" colspan="1">JN121708</td><td align="left" valign="top" rowspan="1" colspan="1">JN121542</td><td align="left" valign="top" rowspan="1" colspan="1">JN121848</td><td align="left" valign="top" rowspan="1" colspan="1">JN121934</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130294&link_type=cbs">CBS 130294</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus penicillioides</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">DTO 11C3</td><td align="left" valign="top" rowspan="1" colspan="1">Indoor environment, Germany</td><td align="left" valign="top" rowspan="1" colspan="1">JN121578</td><td align="left" valign="top" rowspan="1" colspan="1">JN121437</td><td align="left" valign="top" rowspan="1" colspan="1">JN121744</td><td align="left" valign="top" rowspan="1" colspan="1">JN121886</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=578.65&link_type=cbs">CBS 578.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus pulvinus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16842 = IMI 139628</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil, Liberia, Province of Guanacaste, Costa Rica</td><td align="left" valign="top" rowspan="1" colspan="1">JN121703</td><td align="left" valign="top" rowspan="1" colspan="1">JN121536</td><td align="left" valign="top" rowspan="1" colspan="1">JN121842</td><td align="left" valign="top" rowspan="1" colspan="1">JN121930</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=117.33&link_type=cbs">CBS 117.33</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus restrictus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16912 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=541.65&link_type=cbs">CBS 541.65</ext-link> = IMI 016267 = MUCL 31313 = NRRL 154 = NRRL 4155</td><td align="left" valign="top" rowspan="1" colspan="1">Cloth, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121574</td><td align="left" valign="top" rowspan="1" colspan="1">JN121432</td><td align="left" valign="top" rowspan="1" colspan="1">JN121740</td><td align="left" valign="top" rowspan="1" colspan="1">JN121884</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=649.93&link_type=cbs">CBS 649.93</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus robustus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=428.77&link_type=cbs">CBS 428.77</ext-link> = IBT 14305</td><td align="left" valign="top" rowspan="1" colspan="1">Surface soil from thorn-forest, near Mombasa, Kenya</td><td align="left" valign="top" rowspan="1" colspan="1">JN121711</td><td align="left" valign="top" rowspan="1" colspan="1">JN121544</td><td align="left" valign="top" rowspan="1" colspan="1">JN121850</td><td align="left" valign="top" rowspan="1" colspan="1">JN121935</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=139.61&link_type=cbs">CBS 139.61</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus sparsus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16851 = IMI 019394 = IMI 019394ii = MUCL 31314 = NRRL 1933</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Costa Rica</td><td align="left" valign="top" rowspan="1" colspan="1">JN121586</td><td align="left" valign="top" rowspan="1" colspan="1">JN121444</td><td align="left" valign="top" rowspan="1" colspan="1">JN121751</td><td align="left" valign="top" rowspan="1" colspan="1">JN121891</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112812&link_type=cbs">CBS 112812</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus steynii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 23096</td><td align="left" valign="top" rowspan="1" colspan="1">Dried arabica green coffee bean, on parchment, internal infection, Chamumdeshuran Estata, Karnataka, district Giris, India</td><td align="left" valign="top" rowspan="1" colspan="1">JN121569</td><td align="left" valign="top" rowspan="1" colspan="1">JN121428</td><td align="left" valign="top" rowspan="1" colspan="1">JN121735</td><td align="left" valign="top" rowspan="1" colspan="1">JN121880</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=264.81&link_type=cbs">CBS 264.81</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus sydowii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Grains and milling fractions, <italic>Triticum aestivum</italic>, India</td><td align="left" valign="top" rowspan="1" colspan="1">JN121624</td><td align="left" valign="top" rowspan="1" colspan="1">JN121476</td><td align="left" valign="top" rowspan="1" colspan="1">JN121782</td><td align="left" valign="top" rowspan="1" colspan="1">JN121902</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus terreus<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NIH 2624</td><td align="left" valign="top" rowspan="1" colspan="1">Clinical isolate</td><td align="left" valign="top" rowspan="1" colspan="1">Unpublished</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=272.89&link_type=cbs">CBS 272.89</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus togoensis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NRRL 13550</td><td align="left" valign="top" rowspan="1" colspan="1">Seed, near La Maboké, Central African Republic</td><td align="left" valign="top" rowspan="1" colspan="1">JN121627</td><td align="left" valign="top" rowspan="1" colspan="1">JN121480</td><td align="left" valign="top" rowspan="1" colspan="1">JN121785</td><td align="left" valign="top" rowspan="1" colspan="1">JN121904</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=245.65&link_type=cbs">CBS 245.65</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus versicolor</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 11730 = ATCC 16020= IMI 045554 = IMI 045554ii = IMI 045554iii = IMI 045554iv = MUCL 19008</td><td align="left" valign="top" rowspan="1" colspan="1">Cellophane, Indiana, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121614</td><td align="left" valign="top" rowspan="1" colspan="1">JN121468</td><td align="left" valign="top" rowspan="1" colspan="1">JN121775</td><td align="left" valign="top" rowspan="1" colspan="1">JN121899</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=104.07&link_type=cbs">CBS 104.07</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus wentii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1023 = IMI 017295 = IMI 017295ii = NRRL 1269 = NRRL 375</td><td align="left" valign="top" rowspan="1" colspan="1">Soybeans, Java, Indonesia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121559</td><td align="left" valign="top" rowspan="1" colspan="1">JN121418</td><td align="left" valign="top" rowspan="1" colspan="1">JN121725</td><td align="left" valign="top" rowspan="1" colspan="1">JN121873</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=506.65&link_type=cbs">CBS 506.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus zonatus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16867 = IMI 124936</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil, Province of Linon, Fortuna, Costa Rica</td><td align="left" valign="top" rowspan="1" colspan="1">JN121691</td><td align="left" valign="top" rowspan="1" colspan="1">JN121526</td><td align="left" valign="top" rowspan="1" colspan="1">JN121832</td><td align="left" valign="top" rowspan="1" colspan="1">JN121921</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=380.74&link_type=cbs">CBS 380.74</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Basipetospora halophilica</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IFO 9650</td><td align="left" valign="top" rowspan="1" colspan="1">Undaria pinnatifida (Wakame), Osaka, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121666</td><td align="left" valign="top" rowspan="1" colspan="1">JN121509</td><td align="left" valign="top" rowspan="1" colspan="1">JN121815</td><td align="left" valign="top" rowspan="1" colspan="1">JN121910</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100.11&link_type=cbs">CBS 100.11</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Byssochlamys nivea</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22260</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">JN121511</td><td align="left" valign="top" rowspan="1" colspan="1">JF417414</td><td align="left" valign="top" rowspan="1" colspan="1">JF417381</td><td align="left" valign="top" rowspan="1" colspan="1">JF417514</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101075&link_type=cbs">CBS 101075</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Byssochlamys spectabilis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 90900 = FRR 5219</td><td align="left" valign="top" rowspan="1" colspan="1">Heat processed fruit beverage; Tokyo Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121554</td><td align="left" valign="top" rowspan="1" colspan="1">JF417446</td><td align="left" valign="top" rowspan="1" colspan="1">JF417412</td><td align="left" valign="top" rowspan="1" colspan="1">JF417546</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=605.74&link_type=cbs">CBS 605.74</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Byssochlamys verrucosa</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 34163</td><td align="left" valign="top" rowspan="1" colspan="1">Nesting material of <italic>Leipoa ocellata</italic> (Malleefowl), Pulletop Nature Reserve, New South Wales, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN680311</td><td align="left" valign="top" rowspan="1" colspan="1">JN121540</td><td align="left" valign="top" rowspan="1" colspan="1">JN121746</td><td align="left" valign="top" rowspan="1" colspan="1">JN121932</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=132.31&link_type=cbs">CBS 132.31</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Chrysosporium inops</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 096729 = UAMH 802</td><td align="left" valign="top" rowspan="1" colspan="1">Skin man, Italy</td><td align="left" valign="top" rowspan="1" colspan="1">JN121584</td><td align="left" valign="top" rowspan="1" colspan="1">JN121443</td><td align="left" valign="top" rowspan="1" colspan="1">JN121750</td><td align="left" valign="top" rowspan="1" colspan="1">JN121890</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Coccidioides immitis<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">Strain “RS”</td><td align="left" valign="top" rowspan="1" colspan="1">Vaccine strain - origin unknown</td><td align="left" valign="top" rowspan="1" colspan="1">Sharpton <italic>et al.</italic> (<xref ref-type="bibr" rid="R154">2009</xref>)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=525.83&link_type=cbs">CBS 525.83</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cristaspora arxii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 52744 = FMR 416</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Tarragona, Spain</td><td align="left" valign="top" rowspan="1" colspan="1">JN121695</td><td align="left" valign="top" rowspan="1" colspan="1">JN121529</td><td align="left" valign="top" rowspan="1" colspan="1">JN121835</td><td align="left" valign="top" rowspan="1" colspan="1">JN121924</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=157.66&link_type=cbs">CBS 157.66</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichotomomyces cejpii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Orchard soil, near Tiraspol, Moldova</td><td align="left" valign="top" rowspan="1" colspan="1">JN121589</td><td align="left" valign="top" rowspan="1" colspan="1">JN121447</td><td align="left" valign="top" rowspan="1" colspan="1">JN121754</td><td align="left" valign="top" rowspan="1" colspan="1">JN121894</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Emericella nidulans<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">FGSC A4 (= ATCC 38163 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112.46&link_type=cbs">CBS 112.46</ext-link>)</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">Galagan <italic>et al.</italic> (<xref ref-type="bibr" rid="R41">2005</xref>)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=229.60&link_type=cbs">CBS 229.60</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eupenicillium hirayamae</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18312 = IMI 078255 = IMI 078255ii = NRRL 143</td><td align="left" valign="top" rowspan="1" colspan="1">Milled rice, Thailand</td><td align="left" valign="top" rowspan="1" colspan="1">JN121604</td><td align="left" valign="top" rowspan="1" colspan="1">JN121459</td><td align="left" valign="top" rowspan="1" colspan="1">JN121766</td><td align="left" valign="top" rowspan="1" colspan="1">JN121946</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=518.65&link_type=cbs">CBS 518.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eurotium amstelodami</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16464 = IMI 229971 = NRRL 90</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown substrate</td><td align="left" valign="top" rowspan="1" colspan="1">JN121694</td><td align="left" valign="top" rowspan="1" colspan="1">JN121528</td><td align="left" valign="top" rowspan="1" colspan="1">JN121834</td><td align="left" valign="top" rowspan="1" colspan="1">JN121923</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=516.65&link_type=cbs">CBS 516.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eurotium herbariorum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16469 = IMI 211383 = NRRL 116</td><td align="left" valign="top" rowspan="1" colspan="1">Unpainted board, Washington, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121693</td><td align="left" valign="top" rowspan="1" colspan="1">JN121527</td><td align="left" valign="top" rowspan="1" colspan="1">JN121833</td><td align="left" valign="top" rowspan="1" colspan="1">JN121922</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=260.73&link_type=cbs">CBS 260.73</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fennellia flavipes</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 24484 = IMI 171883 = NRRL 5504</td><td align="left" valign="top" rowspan="1" colspan="1">Cellulose material buried in forest soil, Pak Thong Chai, Thailand</td><td align="left" valign="top" rowspan="1" colspan="1">JN121623</td><td align="left" valign="top" rowspan="1" colspan="1">JN121475</td><td align="left" valign="top" rowspan="1" colspan="1">JN121781</td><td align="left" valign="top" rowspan="1" colspan="1">JN121901</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=252.87&link_type=cbs">CBS 252.87</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Geosmithia viridis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 288716</td><td align="left" valign="top" rowspan="1" colspan="1">Soil; bank of creek flowing into Little River; New South Wales; Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121620</td><td align="left" valign="top" rowspan="1" colspan="1">JF417422</td><td align="left" valign="top" rowspan="1" colspan="1">JF417389</td><td align="left" valign="top" rowspan="1" colspan="1">JF417522</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=295.48&link_type=cbs">CBS 295.48</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hamigera avellanea</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10414 = IMI 040230 = NRRL 1938</td><td align="left" valign="top" rowspan="1" colspan="1">Soil; San Antonio, Texas, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121632</td><td align="left" valign="top" rowspan="1" colspan="1">JF417424</td><td align="left" valign="top" rowspan="1" colspan="1">JF417391</td><td align="left" valign="top" rowspan="1" colspan="1">JF417524</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=377.48&link_type=cbs">CBS 377.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hamigera striata</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10501 IMI 039741 = NRRL 717</td><td align="left" valign="top" rowspan="1" colspan="1">Canned blueberries, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121665</td><td align="left" valign="top" rowspan="1" colspan="1">JN121508</td><td align="left" valign="top" rowspan="1" colspan="1">JN121814</td><td align="left" valign="top" rowspan="1" colspan="1">JN121909</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=527.65&link_type=cbs">CBS 527.65</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemicarpenteles paradoxus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16918 = IMI 061446 = NRRL 2162</td><td align="left" valign="top" rowspan="1" colspan="1">Dung of <italic>Opossum</italic>, Wellington, New Zealand</td><td align="left" valign="top" rowspan="1" colspan="1">JN121696</td><td align="left" valign="top" rowspan="1" colspan="1">JN121530</td><td align="left" valign="top" rowspan="1" colspan="1">JN121836</td><td align="left" valign="top" rowspan="1" colspan="1">JN121989</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=607.74&link_type=cbs">CBS 607.74</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Leiothecium ellipsoideum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 32453</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, between rocks, Mystras, Peloponnesos, Greece</td><td align="left" valign="top" rowspan="1" colspan="1">JN121707</td><td align="left" valign="top" rowspan="1" colspan="1">JN121541</td><td align="left" valign="top" rowspan="1" colspan="1">JN121847</td><td align="left" valign="top" rowspan="1" colspan="1">JN121933</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109402&link_type=cbs">CBS 109402</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Monascus argentinensis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">FMR 7393</td><td align="left" valign="top" rowspan="1" colspan="1">Soil sample, El Infiernillo, Tafi del Valle, Tucumán province, Argentina</td><td align="left" valign="top" rowspan="1" colspan="1">JN121564</td><td align="left" valign="top" rowspan="1" colspan="1">JN121423</td><td align="left" valign="top" rowspan="1" colspan="1">JN121730</td><td align="left" valign="top" rowspan="1" colspan="1">JN121877</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=113675&link_type=cbs">CBS 113675</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Monascus lunisporas</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">FMR 6679</td><td align="left" valign="top" rowspan="1" colspan="1">Soil sample, Corcovado Mountain, Tijuca National Park, Rio de Janeiro, Brazil</td><td align="left" valign="top" rowspan="1" colspan="1">JN121570</td><td align="left" valign="top" rowspan="1" colspan="1">JN121429</td><td align="left" valign="top" rowspan="1" colspan="1">JN121736</td><td align="left" valign="top" rowspan="1" colspan="1">JN121881</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109.07&link_type=cbs">CBS 109.07</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Monascus purpureus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16365 = ATCC 16426 = IMI210765 = NRRL 1596</td><td align="left" valign="top" rowspan="1" colspan="1">Fermented rice grain, ‘ang-quac’ (purple coloured rice), Kagok-Tegal, imported from China, Prov. Quouan-toung, Java, Indonesia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121563</td><td align="left" valign="top" rowspan="1" colspan="1">JN121422</td><td align="left" valign="top" rowspan="1" colspan="1">JN121729</td><td align="left" valign="top" rowspan="1" colspan="1">JN121876</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=558.71&link_type=cbs">CBS 558.71</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Neocarpenteles acanthosporum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22931 = IMI 164621</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Bougainville Island, Solomon Islands</td><td align="left" valign="top" rowspan="1" colspan="1">JN121701</td><td align="left" valign="top" rowspan="1" colspan="1">JN121534</td><td align="left" valign="top" rowspan="1" colspan="1">JN121840</td><td align="left" valign="top" rowspan="1" colspan="1">JN121928</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Neosartorya fischeri</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NRRL 181</td><td align="left" valign="top" rowspan="1" colspan="1">Canned fruit</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=350.66&link_type=cbs">CBS 350.66</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Paecilomyces aerugineus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 105412</td><td align="left" valign="top" rowspan="1" colspan="1">Debris of <italic>Glyceria maxima</italic>, Attenborough, Notts., UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121657</td><td align="left" valign="top" rowspan="1" colspan="1">JN121502</td><td align="left" valign="top" rowspan="1" colspan="1">JN121808</td><td align="left" valign="top" rowspan="1" colspan="1">JN121907</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=761.68&link_type=cbs">CBS 761.68</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicilliopsis clavariiformis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">CSIR 1135</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Pretoria, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121716</td><td align="left" valign="top" rowspan="1" colspan="1">JN121549</td><td align="left" valign="top" rowspan="1" colspan="1">JN121855</td><td align="left" valign="top" rowspan="1" colspan="1">JN121940</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=246.67&link_type=cbs">CBS 246.67</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium abidjanum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18385 = FRR 1156 = IMI 136244</td><td align="left" valign="top" rowspan="1" colspan="1">Savannah soil, near Abidjan, Ivory Coast</td><td align="left" valign="top" rowspan="1" colspan="1">JN121615</td><td align="left" valign="top" rowspan="1" colspan="1">JN121469</td><td align="left" valign="top" rowspan="1" colspan="1">JN121777</td><td align="left" valign="top" rowspan="1" colspan="1">JN121954</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=209.28&link_type=cbs">CBS 209.28</ext-link><sup>LT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium adametzii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10407 = IMI 039751 = MUCL 29106 = NRRL 737</td><td align="left" valign="top" rowspan="1" colspan="1">Soil under conifers, Poznan, Poland</td><td align="left" valign="top" rowspan="1" colspan="1">JN121598</td><td align="left" valign="top" rowspan="1" colspan="1">JN121455</td><td align="left" valign="top" rowspan="1" colspan="1">JN121762</td><td align="left" valign="top" rowspan="1" colspan="1">JN121944</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=317.67&link_type=cbs">CBS 317.67</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium alutaceum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18542 = FRR 1158 = IFO 31728 = IMI 136243</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, near Pretoria, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121641</td><td align="left" valign="top" rowspan="1" colspan="1">JN121489</td><td align="left" valign="top" rowspan="1" colspan="1">JN121795</td><td align="left" valign="top" rowspan="1" colspan="1">JN121968</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=220.66&link_type=cbs">CBS 220.66</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium arenicola</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18321 = ATCC 18330 = IMI117658 = NRRL 3392</td><td align="left" valign="top" rowspan="1" colspan="1">Soil from pine forest, Kiev, Ukraine</td><td align="left" valign="top" rowspan="1" colspan="1">JN121601</td><td align="left" valign="top" rowspan="1" colspan="1">JN121457</td><td align="left" valign="top" rowspan="1" colspan="1">JN121764</td><td align="left" valign="top" rowspan="1" colspan="1">JN121897</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=241.56&link_type=cbs">CBS 241.56</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium atrovenetum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 13352 = FRR 2571 = IFO 8138 = IMI 061837</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Sussex Downs, England</td><td align="left" valign="top" rowspan="1" colspan="1">JN121614</td><td align="left" valign="top" rowspan="1" colspan="1">JN121467</td><td align="left" valign="top" rowspan="1" colspan="1">JN121774</td><td align="left" valign="top" rowspan="1" colspan="1">JN121953</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=299.48&link_type=cbs">CBS 299.48</ext-link><sup>AUT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium camemberti</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1105 = ATCC 4845 = FRR 878 = IBT 21508 = IMI 027831 = IMI 092200 = MUCL 29790 = NRRL 877 = NRRL 878</td><td align="left" valign="top" rowspan="1" colspan="1">French Camembert cheese, Connecticut, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121635</td><td align="left" valign="top" rowspan="1" colspan="1">JN121484</td><td align="left" valign="top" rowspan="1" colspan="1">JN121790</td><td align="left" valign="top" rowspan="1" colspan="1">JN121963</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=300.48&link_type=cbs">CBS 300.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium canescens</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10419 = IMI 028260 = MUCL 29169 = NRRL 910</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, England</td><td align="left" valign="top" rowspan="1" colspan="1">JN121636</td><td align="left" valign="top" rowspan="1" colspan="1">JN121485</td><td align="left" valign="top" rowspan="1" colspan="1">JN121791</td><td align="left" valign="top" rowspan="1" colspan="1">JN121964</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=233.81&link_type=cbs">CBS 233.81</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium caperatum</italic></td><td align="left" valign="top" rowspan="1" colspan="1">FRR 71 = IMI 216895</td><td align="left" valign="top" rowspan="1" colspan="1">Neotype of <italic>E. brefeldianum</italic>; soil, Murrumbidgee Irrigation Area, N.S.W., Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121610</td><td align="left" valign="top" rowspan="1" colspan="1">JN121465</td><td align="left" valign="top" rowspan="1" colspan="1">JN121772</td><td align="left" valign="top" rowspan="1" colspan="1">JN121952</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=352.67&link_type=cbs">CBS 352.67</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium catenatum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18543 = IMI 136241</td><td align="left" valign="top" rowspan="1" colspan="1">Desert soil, Upington, Cape Province, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121659</td><td align="left" valign="top" rowspan="1" colspan="1">JN121504</td><td align="left" valign="top" rowspan="1" colspan="1">JN121810</td><td align="left" valign="top" rowspan="1" colspan="1">JN121980</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=304.48&link_type=cbs">CBS 304.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium charlesii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 8730 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=342.51&link_type=cbs">CBS 342.51</ext-link> = IMI 040232 = NRRL 1887 = NRRL 778</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121637</td><td align="left" valign="top" rowspan="1" colspan="1">JN121486</td><td align="left" valign="top" rowspan="1" colspan="1">JN121792</td><td align="left" valign="top" rowspan="1" colspan="1">JN121965</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=306.48&link_type=cbs">CBS 306.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium chrysogenum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10106 = FRR 807 = IBT 5233 = IMI 024314 = IMI 092208 = MUCL 29079 = MUCL 29145 = NRRL 807 = NRRL 810</td><td align="left" valign="top" rowspan="1" colspan="1">Cheese, Storrs, Connecticut</td><td align="left" valign="top" rowspan="1" colspan="1">JN121638</td><td align="left" valign="top" rowspan="1" colspan="1">JN121487</td><td align="left" valign="top" rowspan="1" colspan="1">JN121793</td><td align="left" valign="top" rowspan="1" colspan="1">JN121966</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium chrysogenum<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">Wisconsin 54-1255</td><td align="left" valign="top" rowspan="1" colspan="1">Moldy cantaloupe Peoria, Illinois, USA</td><td align="left" valign="top" rowspan="1" colspan="1">van den Berg <italic>et al</italic>. (2008)</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=490.66&link_type=cbs">CBS 490.66</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium cinnamopurpureum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18337 = IMI 114483</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>E. cinnamopurpureum</italic>; cultivated soil, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121690</td><td align="left" valign="top" rowspan="1" colspan="1">JN121525</td><td align="left" valign="top" rowspan="1" colspan="1">JN121831</td><td align="left" valign="top" rowspan="1" colspan="1">JN121988</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=258.29&link_type=cbs">CBS 258.29</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium citreonigrum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 48736 = 092209 = MUCL 28648 = MUCL 29062 = MUCL 29116 = NRRL 761</td><td align="left" valign="top" rowspan="1" colspan="1">Rotting stem, Belgium</td><td align="left" valign="top" rowspan="1" colspan="1">JN121622</td><td align="left" valign="top" rowspan="1" colspan="1">JN121474</td><td align="left" valign="top" rowspan="1" colspan="1">JN121780</td><td align="left" valign="top" rowspan="1" colspan="1">JN121957</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=139.45&link_type=cbs">CBS 139.45</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium citrinum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1109 = IMI 091961 = MUCL 29781 = NRRL 1841</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown</td><td align="left" valign="top" rowspan="1" colspan="1">JN121585</td><td align="left" valign="top" rowspan="1" colspan="1">JF417416</td><td align="left" valign="top" rowspan="1" colspan="1">JF417383</td><td align="left" valign="top" rowspan="1" colspan="1">JF417516</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=232.38&link_type=cbs">CBS 232.38</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium citrinum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">Thom 4733.73</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>P. implicatum</italic>; unknown source, Belgium</td><td align="left" valign="top" rowspan="1" colspan="1">JN121608</td><td align="left" valign="top" rowspan="1" colspan="1">JN121463</td><td align="left" valign="top" rowspan="1" colspan="1">JN121770</td><td align="left" valign="top" rowspan="1" colspan="1">JN121950</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=119387&link_type=cbs">CBS 119387</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium coffeae</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 27866 = NRRL 35363</td><td align="left" valign="top" rowspan="1" colspan="1">Peduncle, <italic>Coffea arabica</italic>, Oahu, Aiea, Hawaii, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121577</td><td align="left" valign="top" rowspan="1" colspan="1">JN121436</td><td align="left" valign="top" rowspan="1" colspan="1">JN121743</td><td align="left" valign="top" rowspan="1" colspan="1">JN121862</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=231.38&link_type=cbs">CBS 231.38</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium corylophilum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10452 = IFO 7726 = IMI 039817 = NRRL 872</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>P. humuli; Humus lupulus</italic> (hops), Weihenstephan, Germany</td><td align="left" valign="top" rowspan="1" colspan="1">JN121606</td><td align="left" valign="top" rowspan="1" colspan="1">JN121461</td><td align="left" valign="top" rowspan="1" colspan="1">JN121768</td><td align="left" valign="top" rowspan="1" colspan="1">JN121948</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=271.89&link_type=cbs">CBS 271.89</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium cryptum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 60138 = IMI 296794 = NRRL 13460</td><td align="left" valign="top" rowspan="1" colspan="1">Soil from <italic>Quercus-Betula</italic> forest, Hempstead Lake State Park, Long Island, New York</td><td align="left" valign="top" rowspan="1" colspan="1">JN121626</td><td align="left" valign="top" rowspan="1" colspan="1">JN121478</td><td align="left" valign="top" rowspan="1" colspan="1">JN121784</td><td align="left" valign="top" rowspan="1" colspan="1">JN121958</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=660.80&link_type=cbs">CBS 660.80</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium dendriticum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 216897</td><td align="left" valign="top" rowspan="1" colspan="1">Leaf litter of <italic>Eucalyptus pauciflora</italic>, Kosciusko National Park, New South Wales, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121714</td><td align="left" valign="top" rowspan="1" colspan="1">JN121547</td><td align="left" valign="top" rowspan="1" colspan="1">JN121853</td><td align="left" valign="top" rowspan="1" colspan="1">JN121938</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112082&link_type=cbs">CBS 112082</ext-link><sup>epiT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium digitatum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 13068</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Citrus limon</italic>, Italy</td><td align="left" valign="top" rowspan="1" colspan="1">JN121567</td><td align="left" valign="top" rowspan="1" colspan="1">JN121426</td><td align="left" valign="top" rowspan="1" colspan="1">JN121733</td><td align="left" valign="top" rowspan="1" colspan="1">JN121858</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=456.70&link_type=cbs">CBS 456.70</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium dimorphosporum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22783 = ATCC 52501 = FRR 1120 = IMI 149680</td><td align="left" valign="top" rowspan="1" colspan="1">Mangrove swamp soil, below high tide level, Tooraddin, Westernport Bay, Sawtell's Inlet, Victoria, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121683</td><td align="left" valign="top" rowspan="1" colspan="1">JN121517</td><td align="left" valign="top" rowspan="1" colspan="1">JN121823</td><td align="left" valign="top" rowspan="1" colspan="1">JN121985</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=322.48&link_type=cbs">CBS 322.48</ext-link><sup>AUT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium duclauxii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030</td><td align="left" valign="top" rowspan="1" colspan="1">Canvas, France</td><td align="left" valign="top" rowspan="1" colspan="1">JN121643</td><td align="left" valign="top" rowspan="1" colspan="1">JN121491</td><td align="left" valign="top" rowspan="1" colspan="1">JN121797</td><td align="left" valign="top" rowspan="1" colspan="1">JN121905</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=112493&link_type=cbs">CBS 112493</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium ellipsoideosporum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">AS 3.5688</td><td align="left" valign="top" rowspan="1" colspan="1">Banyan seeds, Pingxiang, Guanbxi Province, China (data after Wang <italic>et al</italic>. 2007)</td><td align="left" valign="top" rowspan="1" colspan="1">JN121568</td><td align="left" valign="top" rowspan="1" colspan="1">JN121427</td><td align="left" valign="top" rowspan="1" colspan="1">JN121734</td><td align="left" valign="top" rowspan="1" colspan="1">JN121859</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=318.67&link_type=cbs">CBS 318.67</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium erubescens</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18544 = FRR 814 = IFO 31734 = IMI 136204</td><td align="left" valign="top" rowspan="1" colspan="1">Nursery soil, Pretoria, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121642</td><td align="left" valign="top" rowspan="1" colspan="1">JN121490</td><td align="left" valign="top" rowspan="1" colspan="1">JN121796</td><td align="left" valign="top" rowspan="1" colspan="1">JN121969</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=323.71&link_type=cbs">CBS 323.71</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium euglaucum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Argentina</td><td align="left" valign="top" rowspan="1" colspan="1">JN121644</td><td align="left" valign="top" rowspan="1" colspan="1">JN121492</td><td align="left" valign="top" rowspan="1" colspan="1">JN121798</td><td align="left" valign="top" rowspan="1" colspan="1">JN121970</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=325.48&link_type=cbs">CBS 325.48</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium expansum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 7861 = IBT 5101 = IMI 039761= MUCL 29192 = NRRL 976</td><td align="left" valign="top" rowspan="1" colspan="1">Fruit of <italic>Malus sylvestris</italic>; USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121645</td><td align="left" valign="top" rowspan="1" colspan="1">JF417427</td><td align="left" valign="top" rowspan="1" colspan="1">JF417394</td><td align="left" valign="top" rowspan="1" colspan="1">JF417527</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=229.81&link_type=cbs">CBS 229.81</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium fellutanum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10443 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=326.48&link_type=cbs">CBS 326.48</ext-link> = FRR 746 = IFO 5761 = IMI 039734 = IMI 039734iii = NRRL 746</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121605</td><td align="left" valign="top" rowspan="1" colspan="1">JN121460</td><td align="left" valign="top" rowspan="1" colspan="1">JN121767</td><td align="left" valign="top" rowspan="1" colspan="1">JN121947</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124.68&link_type=cbs">CBS 124.68</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium fractum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18567 = FRR 3448 = IMI 136701 = NRRL 3448</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Univ. Shinshu, Ueda-shi, Nagano Pref, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121582</td><td align="left" valign="top" rowspan="1" colspan="1">JN121441</td><td align="left" valign="top" rowspan="1" colspan="1">JN121748</td><td align="left" valign="top" rowspan="1" colspan="1">JN121864</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=295.62&link_type=cbs">CBS 295.62</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium fuscum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 14770 = IFO 7743 = IMI 094209 = MUCL 31196 = NRRL 3008 = WSF 15c</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>E. pinetorum</italic> and neotype of <italic>Citromyces fuscus</italic>; pine-birch forest soil, Vilas County, Wisconsin, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121633</td><td align="left" valign="top" rowspan="1" colspan="1">JN121483</td><td align="left" valign="top" rowspan="1" colspan="1">JN121789</td><td align="left" valign="top" rowspan="1" colspan="1">JN121962</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125543&link_type=cbs">CBS 125543</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium glabrum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 22658 = IMI 91944</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown</td><td align="left" valign="top" rowspan="1" colspan="1">JN121717</td><td align="left" valign="top" rowspan="1" colspan="1">JF417447</td><td align="left" valign="top" rowspan="1" colspan="1">JF417413</td><td align="left" valign="top" rowspan="1" colspan="1">JF417547</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=599.73&link_type=cbs">CBS 599.73</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium gracilentum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 28047 = ATCC 48258 = IMI 216900</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Brown River, Port Moresby, Central Dist., Papua New Guinea</td><td align="left" valign="top" rowspan="1" colspan="1">JN121704</td><td align="left" valign="top" rowspan="1" colspan="1">JN121537</td><td align="left" valign="top" rowspan="1" colspan="1">JN121843</td><td align="left" valign="top" rowspan="1" colspan="1">JN121990</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.27&link_type=cbs">CBS 185.27</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium griseofulvum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 11885 = IBT 6740 = IMI 075832 = IMI 075832ii = MUCL 28643 = NRRL 2152 = NRRL 2300</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Belgium</td><td align="left" valign="top" rowspan="1" colspan="1">JN121592</td><td align="left" valign="top" rowspan="1" colspan="1">JN121449</td><td align="left" valign="top" rowspan="1" colspan="1">JN121756</td><td align="left" valign="top" rowspan="1" colspan="1">JN121865</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=277.58&link_type=cbs">CBS 277.58</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium griseolum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18239 = IMI 071626 = NRRL 2671</td><td align="left" valign="top" rowspan="1" colspan="1">Acidic dune sand, Dorset, Stufland, England</td><td align="left" valign="top" rowspan="1" colspan="1">JN121629</td><td align="left" valign="top" rowspan="1" colspan="1">JN121480</td><td align="left" valign="top" rowspan="1" colspan="1">JN121786</td><td align="left" valign="top" rowspan="1" colspan="1">JN121959</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=336.48&link_type=cbs">CBS 336.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium herquei</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10118 = FRR 1040 = IFO 31747 = IMI 028809 = MUCL 29213 = NRRL 1040</td><td align="left" valign="top" rowspan="1" colspan="1">Leaf, France</td><td align="left" valign="top" rowspan="1" colspan="1">JN121647</td><td align="left" valign="top" rowspan="1" colspan="1">JN121494</td><td align="left" valign="top" rowspan="1" colspan="1">JN121800</td><td align="left" valign="top" rowspan="1" colspan="1">JN121972</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.68&link_type=cbs">CBS 341.68</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium idahoense</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22055 = IMI 148393</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Latàh Co., Univ. of Idaho Plant Science Farm, Idaho, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121652</td><td align="left" valign="top" rowspan="1" colspan="1">JN121499</td><td align="left" valign="top" rowspan="1" colspan="1">JN121805</td><td align="left" valign="top" rowspan="1" colspan="1">JN121976</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=351.67&link_type=cbs">CBS 351.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium inusitatum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18622 = IMI 136214</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil, Knysna Valley, Cape Province, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121658</td><td align="left" valign="top" rowspan="1" colspan="1">JN121503</td><td align="left" valign="top" rowspan="1" colspan="1">JN121809</td><td align="left" valign="top" rowspan="1" colspan="1">JN121979</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.56&link_type=cbs">CBS 247.56</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium isariiforme</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18425 = IMI 060371 = MUCL 31191 MUCL 31323 = NRRL 2638</td><td align="left" valign="top" rowspan="1" colspan="1">Woodland soil, Zaire</td><td align="left" valign="top" rowspan="1" colspan="1">JN121616</td><td align="left" valign="top" rowspan="1" colspan="1">JN121470</td><td align="left" valign="top" rowspan="1" colspan="1">JN121720</td><td align="left" valign="top" rowspan="1" colspan="1">JN121993</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=338.48&link_type=cbs">CBS 338.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium islandicum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Cape Town, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121648</td><td align="left" valign="top" rowspan="1" colspan="1">JN121495</td><td align="left" valign="top" rowspan="1" colspan="1">JN121801</td><td align="left" valign="top" rowspan="1" colspan="1">JN121906</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=339.48&link_type=cbs">CBS 339.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium italicum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10454 = FRR 983 = IBT 23029 = IMI 039760 = MUCL 15608 = NRRL 983</td><td align="left" valign="top" rowspan="1" colspan="1">Fruit, Citrus Experiment Station, Riverside, California, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121649</td><td align="left" valign="top" rowspan="1" colspan="1">JN121496</td><td align="left" valign="top" rowspan="1" colspan="1">JN121802</td><td align="left" valign="top" rowspan="1" colspan="1">JN121973</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=340.48&link_type=cbs">CBS 340.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium janthinellum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10455 = IMI 040238 = NRRL 2016</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Nicaragua</td><td align="left" valign="top" rowspan="1" colspan="1">JN131650</td><td align="left" valign="top" rowspan="1" colspan="1">JN121497</td><td align="left" valign="top" rowspan="1" colspan="1">JN121803</td><td align="left" valign="top" rowspan="1" colspan="1">JN121974</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=341.48&link_type=cbs">CBS 341.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium javanicum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 9099 = FRR 707 = IMI 039733 = MUCL 29099 = NRRL 707</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>P. javanicum, E. javanicum</italic> and <italic>P. indonesiae</italic>; root of <italic>Camellia sinensis</italic> (green tea), Buitenzorg, Java, Indonesia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121651</td><td align="left" valign="top" rowspan="1" colspan="1">JN121498</td><td align="left" valign="top" rowspan="1" colspan="1">JN121804</td><td align="left" valign="top" rowspan="1" colspan="1">JN121975</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.67&link_type=cbs">CBS 247.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium katangense</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18388 = IMI 136206 = NRRL 5182</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Katanga, Zaire</td><td align="left" valign="top" rowspan="1" colspan="1">JN121618</td><td align="left" valign="top" rowspan="1" colspan="1">JN121471</td><td align="left" valign="top" rowspan="1" colspan="1">JN121777</td><td align="left" valign="top" rowspan="1" colspan="1">JN121955</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=344.61&link_type=cbs">CBS 344.61</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium kewense</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18240 = IMI 086561= MUCL 2685 = NRRL 3332</td><td align="left" valign="top" rowspan="1" colspan="1">Culture contaminant of mineral oil, Kew, Surrey, England, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121654</td><td align="left" valign="top" rowspan="1" colspan="1">JF417428</td><td align="left" valign="top" rowspan="1" colspan="1">JF417395</td><td align="left" valign="top" rowspan="1" colspan="1">JF417528</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=106.11&link_type=cbs">CBS 106.11</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium lanosum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10458 = IMI 040224 = MUCL 29232 = NRRL 2009</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Germany</td><td align="left" valign="top" rowspan="1" colspan="1">JN121561</td><td align="left" valign="top" rowspan="1" colspan="1">JN121420</td><td align="left" valign="top" rowspan="1" colspan="1">JN121727</td><td align="left" valign="top" rowspan="1" colspan="1">JN121857</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=343.48&link_type=cbs">CBS 343.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium lapidosum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10462 = IMI 039743 = NRRL 718</td><td align="left" valign="top" rowspan="1" colspan="1">Canned blueberry, Washington, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121653</td><td align="left" valign="top" rowspan="1" colspan="1">JN121500</td><td align="left" valign="top" rowspan="1" colspan="1">JN121806</td><td align="left" valign="top" rowspan="1" colspan="1">JN121977</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=277.70&link_type=cbs">CBS 277.70</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium lassenii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22054 = IMI 148395</td><td align="left" valign="top" rowspan="1" colspan="1">Soil under conifers, Tehama Co., Lassen National Forest, 1300 m alt., California, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121630</td><td align="left" valign="top" rowspan="1" colspan="1">JN121481</td><td align="left" valign="top" rowspan="1" colspan="1">JN121787</td><td align="left" valign="top" rowspan="1" colspan="1">JN121960</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=116871&link_type=cbs">CBS 116871</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium macrosclerotiorum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">AS 3.6581</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Chongqing, Wushan County, Sichuang Province, China</td><td align="left" valign="top" rowspan="1" colspan="1">JN121573</td><td align="left" valign="top" rowspan="1" colspan="1">JN121432</td><td align="left" valign="top" rowspan="1" colspan="1">121739</td><td align="left" valign="top" rowspan="1" colspan="1">JN121860</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=647.95&link_type=cbs">CBS 647.95</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium malachiteum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 17515</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Nihondaira Pref. Park, Shimizu-shi, Shimizu-ken, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121710</td><td align="left" valign="top" rowspan="1" colspan="1">JN121543</td><td align="left" valign="top" rowspan="1" colspan="1">JN121849</td><td align="left" valign="top" rowspan="1" colspan="1">JN121991</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium marneffei<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18224 (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=334.59&link_type=cbs">CBS 334.59</ext-link> = IMI 68794)</td><td align="left" valign="top" rowspan="1" colspan="1">Bamboo rat (<italic>Rhizomys sinensis</italic>); Vietnam</td><td align="left" valign="top" rowspan="1" colspan="1">Unpublished</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=256.55&link_type=cbs">CBS 256.55</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium megasporum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 12322 = IMI 216904 = NRRL 2232</td><td align="left" valign="top" rowspan="1" colspan="1">Heath soil,Suffolk, England</td><td align="left" valign="top" rowspan="1" colspan="1">JN121621</td><td align="left" valign="top" rowspan="1" colspan="1">JN121473</td><td align="left" valign="top" rowspan="1" colspan="1">JN121779</td><td align="left" valign="top" rowspan="1" colspan="1">JN121900</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=642.68&link_type=cbs">CBS 642.68</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium minioluteum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 089377 = MUCL 28666</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown</td><td align="left" valign="top" rowspan="1" colspan="1">JN121709</td><td align="left" valign="top" rowspan="1" colspan="1">JF417443</td><td align="left" valign="top" rowspan="1" colspan="1">JF417409</td><td align="left" valign="top" rowspan="1" colspan="1">JF417543</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=353.48&link_type=cbs">CBS 353.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium namyslowskii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 11127 = IMI 040033 = MUCL 29226 = NRRL 1070</td><td align="left" valign="top" rowspan="1" colspan="1">Soil under <italic>Pinus</italic> sp.; Puszcza Bialowieska, Poland</td><td align="left" valign="top" rowspan="1" colspan="1">JN121660</td><td align="left" valign="top" rowspan="1" colspan="1">JF417430</td><td align="left" valign="top" rowspan="1" colspan="1">JF417397</td><td align="left" valign="top" rowspan="1" colspan="1">JF417530</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=203.84&link_type=cbs">CBS 203.84</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium nepalense</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">NHL 6482</td><td align="left" valign="top" rowspan="1" colspan="1">Rice soil, Boudha, Kathmandu, Nepal</td><td align="left" valign="top" rowspan="1" colspan="1">JN121596</td><td align="left" valign="top" rowspan="1" colspan="1">JN121453</td><td align="left" valign="top" rowspan="1" colspan="1">JN121760</td><td align="left" valign="top" rowspan="1" colspan="1">JN121868</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=489.66&link_type=cbs">CBS 489.66</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium ochrosalmoneum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18338 = IMI 116248ii</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>E. ochrosalmoneum</italic>; cornmeal, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121689</td><td align="left" valign="top" rowspan="1" colspan="1">JN121524</td><td align="left" valign="top" rowspan="1" colspan="1">JN121830</td><td align="left" valign="top" rowspan="1" colspan="1">JN121987</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=232.60&link_type=cbs">CBS 232.60</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium olsonii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 23473 = IMI 192502</td><td align="left" valign="top" rowspan="1" colspan="1">Root, <italic>Picea abies</italic>, alt. 1980 m., Pitztal, Austria</td><td align="left" valign="top" rowspan="1" colspan="1">JN121609</td><td align="left" valign="top" rowspan="1" colspan="1">JN121464</td><td align="left" valign="top" rowspan="1" colspan="1">JN121771</td><td align="left" valign="top" rowspan="1" colspan="1">JN121952</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=190.68&link_type=cbs">CBS 190.68</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium ornatum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18608 = IMI 137977 = NRRL 3471</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Moto-machi, Oshima Islands, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121594</td><td align="left" valign="top" rowspan="1" colspan="1">JN121451</td><td align="left" valign="top" rowspan="1" colspan="1">JN121758</td><td align="left" valign="top" rowspan="1" colspan="1">JN121867</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=462.72&link_type=cbs">CBS 462.72</ext-link><sup>HT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium osmophilum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 14679</td><td align="left" valign="top" rowspan="1" colspan="1">Agricultural soil, Wageningen, the Netherlands</td><td align="left" valign="top" rowspan="1" colspan="1">JN121683</td><td align="left" valign="top" rowspan="1" colspan="1">JN121518</td><td align="left" valign="top" rowspan="1" colspan="1">JN121824</td><td align="left" valign="top" rowspan="1" colspan="1">JN121986</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=219.30&link_type=cbs">CBS 219.30</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium oxalicum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 1126 = FRR 787 = IMI 192332 = MUCL 29047 = NRRL 787</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Connecticut</td><td align="left" valign="top" rowspan="1" colspan="1">JN121600</td><td align="left" valign="top" rowspan="1" colspan="1">JN121456</td><td align="left" valign="top" rowspan="1" colspan="1">JN131763</td><td align="left" valign="top" rowspan="1" colspan="1">JN121944</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=251.56&link_type=cbs">CBS 251.56</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium ramusculum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 12292 = IMI 063546 = NRRL 3459</td><td align="left" valign="top" rowspan="1" colspan="1">Culture contaminant, Brazil</td><td align="left" valign="top" rowspan="1" colspan="1">JN121620</td><td align="left" valign="top" rowspan="1" colspan="1">JN121472</td><td align="left" valign="top" rowspan="1" colspan="1">JN121778</td><td align="left" valign="top" rowspan="1" colspan="1">JN121956</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=367.48&link_type=cbs">CBS 367.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium restrictum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 11257 = FRR 1748 = IMI 040228 = NRRL 1748</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Honduras</td><td align="left" valign="top" rowspan="1" colspan="1">JN121662</td><td align="left" valign="top" rowspan="1" colspan="1">JN121506</td><td align="left" valign="top" rowspan="1" colspan="1">JN121812</td><td align="left" valign="top" rowspan="1" colspan="1">JN121981</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=231.61&link_type=cbs">CBS 231.61</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium sacculum</italic> (syn. <italic>Eladia saccula</italic>)<sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18350 = IMI 051498</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Madrid, Spain</td><td align="left" valign="top" rowspan="1" colspan="1">JN121607</td><td align="left" valign="top" rowspan="1" colspan="1">JN121462</td><td align="left" valign="top" rowspan="1" colspan="1">JN121769</td><td align="left" valign="top" rowspan="1" colspan="1">JN121949</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=122276&link_type=cbs">CBS 122276</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium saturniforme</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">AS 3.6886</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Jiling Province, China</td><td align="left" valign="top" rowspan="1" colspan="1">JN121580</td><td align="left" valign="top" rowspan="1" colspan="1">JN121439</td><td align="left" valign="top" rowspan="1" colspan="1">JN121746</td><td align="left" valign="top" rowspan="1" colspan="1">JN121863</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=290.48&link_type=cbs">CBS 290.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium shearii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10410 = IMI 039739 = IMI 039739iv = NRRL 715</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Tela, Honduras</td><td align="left" valign="top" rowspan="1" colspan="1">JN121631</td><td align="left" valign="top" rowspan="1" colspan="1">JN121482</td><td align="left" valign="top" rowspan="1" colspan="1">JN121788</td><td align="left" valign="top" rowspan="1" colspan="1">JN121961</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=228.89&link_type=cbs">CBS 228.89</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium shennangjianum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">AS 3.4526</td><td align="left" valign="top" rowspan="1" colspan="1">Mouldy pea, Hubei Province, Shennongjia, China</td><td align="left" valign="top" rowspan="1" colspan="1">JN121603</td><td align="left" valign="top" rowspan="1" colspan="1">JN121458</td><td align="left" valign="top" rowspan="1" colspan="1">JN121766</td><td align="left" valign="top" rowspan="1" colspan="1">JN121945</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=372.48&link_type=cbs">CBS 372.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium simplicissimum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10495 = IFO 5762 = IMI 039816</td><td align="left" valign="top" rowspan="1" colspan="1">Flannel bag, Cape, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121662</td><td align="left" valign="top" rowspan="1" colspan="1">JN121507</td><td align="left" valign="top" rowspan="1" colspan="1">JN121813</td><td align="left" valign="top" rowspan="1" colspan="1">JN121981</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=315.67&link_type=cbs">CBS 315.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium stolkiae</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18546 = IMI 136210</td><td align="left" valign="top" rowspan="1" colspan="1">Peaty forest soil, Eastern Transvaal, South Africa</td><td align="left" valign="top" rowspan="1" colspan="1">JN121640</td><td align="left" valign="top" rowspan="1" colspan="1">JN121488</td><td align="left" valign="top" rowspan="1" colspan="1">JN121794</td><td align="left" valign="top" rowspan="1" colspan="1">JN121967</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=117503&link_type=cbs">CBS 117503</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium thiersii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 27050 = NRRL 28162</td><td align="left" valign="top" rowspan="1" colspan="1">Old, black stroma, encrusting the surface of dead <italic>Acer saccharum</italic> log, alt. 300 m., New Glarus Woods State Park, Wisconsin, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121575</td><td align="left" valign="top" rowspan="1" colspan="1">JN121434</td><td align="left" valign="top" rowspan="1" colspan="1">JN121741</td><td align="left" valign="top" rowspan="1" colspan="1">JN121861</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=347.59&link_type=cbs">CBS 347.59</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium thomii</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IFO 6031 = IMI 068221</td><td align="left" valign="top" rowspan="1" colspan="1">Type of <italic>P. thomii</italic> var. <italic>flavescens</italic>; soil, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121655</td><td align="left" valign="top" rowspan="1" colspan="1">JN121501</td><td align="left" valign="top" rowspan="1" colspan="1">JN121807</td><td align="left" valign="top" rowspan="1" colspan="1">JN121978</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=430.69&link_type=cbs">CBS 430.69</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium tularense</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22056 = IMI 148394</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, under <italic>Pinus ponderosa</italic> and <italic>Quercus kelloggii</italic>, Tulare Co., Pine Flat, California</td><td align="left" valign="top" rowspan="1" colspan="1">JN121681</td><td align="left" valign="top" rowspan="1" colspan="1">JN121516</td><td align="left" valign="top" rowspan="1" colspan="1">JN121822</td><td align="left" valign="top" rowspan="1" colspan="1">JN121984</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=603.74&link_type=cbs">CBS 603.74</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium verrucosum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 48957 = FRR 965 = IBT 12809 = IBT 4733 = IMI 200310 = IMI 200310ii = MUCL 28674 = MUCL 29089 = MUCL 29186 = NRRL 965</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Belgium</td><td align="left" valign="top" rowspan="1" colspan="1">JN121706</td><td align="left" valign="top" rowspan="1" colspan="1">JN121539</td><td align="left" valign="top" rowspan="1" colspan="1">JN121845</td><td align="left" valign="top" rowspan="1" colspan="1">JN121991</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=390.48&link_type=cbs">CBS 390.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium viridicatum</italic><sup><xref ref-type="table-fn" rid="TFN3">**</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10515= IBT 23041 = IMI 039758 = IMI 039758ii = NRRL 963</td><td align="left" valign="top" rowspan="1" colspan="1">Air, District of Columbia, Washington D.C., USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121668</td><td align="left" valign="top" rowspan="1" colspan="1">JN121511</td><td align="left" valign="top" rowspan="1" colspan="1">JN121817</td><td align="left" valign="top" rowspan="1" colspan="1">JN121983</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=430.64&link_type=cbs">CBS 430.64</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialomyces macrosporus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16661 = IMI 110130 = MUCL 9776</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, near Rotorua, New Zealand</td><td align="left" valign="top" rowspan="1" colspan="1">JN121680</td><td align="left" valign="top" rowspan="1" colspan="1">JN121515</td><td align="left" valign="top" rowspan="1" colspan="1">JN121821</td><td align="left" valign="top" rowspan="1" colspan="1">JN121915</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=128032&link_type=cbs">CBS 128032</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialosimplex caninus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">UAMH 10337</td><td align="left" valign="top" rowspan="1" colspan="1">Bone marrow aspirate ex dog, San Antonio, Texas, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121587</td><td align="left" valign="top" rowspan="1" colspan="1">JN121445</td><td align="left" valign="top" rowspan="1" colspan="1">JN121752</td><td align="left" valign="top" rowspan="1" colspan="1">JN121892</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=109945&link_type=cbs">CBS 109945</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialosimplex chlamydosporus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">FMR 7371 = IMI 387422</td><td align="left" valign="top" rowspan="1" colspan="1">Disseminated infection in a dog</td><td align="left" valign="top" rowspan="1" colspan="1">JN121566</td><td align="left" valign="top" rowspan="1" colspan="1">JN121425</td><td align="left" valign="top" rowspan="1" colspan="1">JN121732</td><td align="left" valign="top" rowspan="1" colspan="1">JN121879</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=366.77&link_type=cbs">CBS 366.77</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialosimplex sclerotialis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IAM 14794</td><td align="left" valign="top" rowspan="1" colspan="1">Fodder of ray-grass and lucerne, France</td><td align="left" valign="top" rowspan="1" colspan="1">JN121661</td><td align="left" valign="top" rowspan="1" colspan="1">JN121505</td><td align="left" valign="top" rowspan="1" colspan="1">JN121811</td><td align="left" valign="top" rowspan="1" colspan="1">JN121908</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=384.61&link_type=cbs">CBS 384.61</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Polypaecilum insolitum</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18164 = IMI 075202 = MUCL 3078</td><td align="left" valign="top" rowspan="1" colspan="1">Ear of human, Leeds, Yorkshire, England, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121667</td><td align="left" valign="top" rowspan="1" colspan="1">JN121510</td><td align="left" valign="top" rowspan="1" colspan="1">JN121816</td><td align="left" valign="top" rowspan="1" colspan="1">JN121911</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101166&link_type=cbs">CBS 101166</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Polypaecilum pisci</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Yeast extract, Netherlands</td><td align="left" valign="top" rowspan="1" colspan="1">JN121555</td><td align="left" valign="top" rowspan="1" colspan="1">JN121415</td><td align="left" valign="top" rowspan="1" colspan="1">JN121722</td><td align="left" valign="top" rowspan="1" colspan="1">JN121870</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=101.69&link_type=cbs">CBS 101.69</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Rasamsonia argillacea</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">DTO 97E4 = IMI 156096 = IBT 31199</td><td align="left" valign="top" rowspan="1" colspan="1">Mine tip with a very high surface temperature; Staffordshire, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121556</td><td align="left" valign="top" rowspan="1" colspan="1">JF417415</td><td align="left" valign="top" rowspan="1" colspan="1">JF417382</td><td align="left" valign="top" rowspan="1" colspan="1">JF417515</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=413.71&link_type=cbs">CBS 413.71</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Rasamsonia byssochlamydoides</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">DTO 149D6 = IBT 11604</td><td align="left" valign="top" rowspan="1" colspan="1">Dry soil under Douglas fir; Oregon, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121675</td><td align="left" valign="top" rowspan="1" colspan="1">JF417437</td><td align="left" valign="top" rowspan="1" colspan="1">JF417403</td><td align="left" valign="top" rowspan="1" colspan="1">JF417537</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=275.58&link_type=cbs">CBS 275.58</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Rasamsonia cylindrospora</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">DTO 138F8 = IBT 31202 = ATCC 18223 = IMI 071623</td><td align="left" valign="top" rowspan="1" colspan="1">Culture contaminant; Berkshire, England, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121628</td><td align="left" valign="top" rowspan="1" colspan="1">JF417423</td><td align="left" valign="top" rowspan="1" colspan="1">JF417390</td><td align="left" valign="top" rowspan="1" colspan="1">JF417523</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=393.64&link_type=cbs">CBS 393.64</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Rasamsonia emersonii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">DTO 48I1 = IBT 21695 = ATCC 16479 = IMI 116815 = IMI 116815ii</td><td align="left" valign="top" rowspan="1" colspan="1">Compost; Italy</td><td align="left" valign="top" rowspan="1" colspan="1">JN121670</td><td align="left" valign="top" rowspan="1" colspan="1">JF417434</td><td align="left" valign="top" rowspan="1" colspan="1">JF417401</td><td align="left" valign="top" rowspan="1" colspan="1">JF417534</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=114.72&link_type=cbs">CBS 114.72</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenoma viride</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22467 = NRRL 5575</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121571</td><td align="left" valign="top" rowspan="1" colspan="1">JN121430</td><td align="left" valign="top" rowspan="1" colspan="1">JN121737</td><td align="left" valign="top" rowspan="1" colspan="1">JN121882</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=545.86&link_type=cbs">CBS 545.86</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella bohemica</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">CCF 2330 = IAM 14789</td><td align="left" valign="top" rowspan="1" colspan="1">Peloids for balneological purposes, Frantiskovy Lázne Spa, West Bohemia, Czech Republic</td><td align="left" valign="top" rowspan="1" colspan="1">JN121699</td><td align="left" valign="top" rowspan="1" colspan="1">JN121532</td><td align="left" valign="top" rowspan="1" colspan="1">JN121838</td><td align="left" valign="top" rowspan="1" colspan="1">JN121927</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=398.69&link_type=cbs">CBS 398.69</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella diversispora</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil under <italic>Populus tremuloides</italic>; Petawawa, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121673</td><td align="left" valign="top" rowspan="1" colspan="1">JF417435</td><td align="left" valign="top" rowspan="1" colspan="1">JF417402</td><td align="left" valign="top" rowspan="1" colspan="1">JF417536</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=399.69&link_type=cbs">CBS 399.69</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella diversispora</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">MUCL 15012</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil under <italic>Thuja occidentalis</italic>, Aberfoyle, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121674</td><td align="left" valign="top" rowspan="1" colspan="1">JN121513</td><td align="left" valign="top" rowspan="1" colspan="1">JN121819</td><td align="left" valign="top" rowspan="1" colspan="1">JN121913</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=426.67&link_type=cbs">CBS 426.67</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella griseoviridis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18505 = IMI 113160</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">JN121677</td><td align="left" valign="top" rowspan="1" colspan="1">JF417438</td><td align="left" valign="top" rowspan="1" colspan="1">JF417404</td><td align="left" valign="top" rowspan="1" colspan="1">JF417538</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=427.67&link_type=cbs">CBS 427.67</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella humicola</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18506 = IMI 113166</td><td align="left" valign="top" rowspan="1" colspan="1">Forest soil under <italic>Thuja occidentalis</italic>; Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121678</td><td align="left" valign="top" rowspan="1" colspan="1">JF417439</td><td align="left" valign="top" rowspan="1" colspan="1">JF417405</td><td align="left" valign="top" rowspan="1" colspan="1">JF417539</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=429.67&link_type=cbs">CBS 429.67</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella striatispora</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18510 = IMI 113163</td><td align="left" valign="top" rowspan="1" colspan="1">Soil; Guelph, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121679</td><td align="left" valign="top" rowspan="1" colspan="1">JF417440</td><td align="left" valign="top" rowspan="1" colspan="1">JF417406</td><td align="left" valign="top" rowspan="1" colspan="1">JF417540</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=414.78&link_type=cbs">CBS 414.78</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella verticillata</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IAM 14697</td><td align="left" valign="top" rowspan="1" colspan="1">Conifer forest soil, Sweden</td><td align="left" valign="top" rowspan="1" colspan="1">JN121676</td><td align="left" valign="top" rowspan="1" colspan="1">JN121514</td><td align="left" valign="top" rowspan="1" colspan="1">JN121820</td><td align="left" valign="top" rowspan="1" colspan="1">JN121914</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=124.53&link_type=cbs">CBS 124.53</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerocleista ornata</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16921 = IMI 055295 = MUCL 15643 = NRRL 2256</td><td align="left" valign="top" rowspan="1" colspan="1">Soil in oak forest, Dane Co., Madison, Wisconsin, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121581</td><td align="left" valign="top" rowspan="1" colspan="1">JN121440</td><td align="left" valign="top" rowspan="1" colspan="1">JN121747</td><td align="left" valign="top" rowspan="1" colspan="1">JN121888</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=105.25&link_type=cbs">CBS 105.25</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerocleista thaxteri</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 055296 = NRRL 2292</td><td align="left" valign="top" rowspan="1" colspan="1">Dung of caterpillar, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121560</td><td align="left" valign="top" rowspan="1" colspan="1">JN121419</td><td align="left" valign="top" rowspan="1" colspan="1">JN121726</td><td align="left" valign="top" rowspan="1" colspan="1">JN121874</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=296.48&link_type=cbs">CBS 296.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces bacillisporus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10126 = IMI 040045 = NRRL 1025</td><td align="left" valign="top" rowspan="1" colspan="1">Begonia leaf; New York City, New York, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121634</td><td align="left" valign="top" rowspan="1" colspan="1">JF417425</td><td align="left" valign="top" rowspan="1" colspan="1">JF417392</td><td align="left" valign="top" rowspan="1" colspan="1">JF417525</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100537&link_type=cbs">CBS 100537</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces convolutus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 14989</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, Kathmandu, Nepal</td><td align="left" valign="top" rowspan="1" colspan="1">JN121553</td><td align="left" valign="top" rowspan="1" colspan="1">JN121414</td><td align="left" valign="top" rowspan="1" colspan="1">JN121721</td><td align="left" valign="top" rowspan="1" colspan="1">JN121869</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=100536&link_type=cbs">CBS 100536</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces emodensis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IBT 14990</td><td align="left" valign="top" rowspan="1" colspan="1">Soil; Kathmandu, Nepal</td><td align="left" valign="top" rowspan="1" colspan="1">JN121552</td><td align="left" valign="top" rowspan="1" colspan="1">JF417445</td><td align="left" valign="top" rowspan="1" colspan="1">JF417411</td><td align="left" valign="top" rowspan="1" colspan="1">JF417545</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces flavus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 197477 = NRRL 2098</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown substrate; New Zealand</td><td align="left" valign="top" rowspan="1" colspan="1">JN121639</td><td align="left" valign="top" rowspan="1" colspan="1">JF417426</td><td align="left" valign="top" rowspan="1" colspan="1">JF417393</td><td align="left" valign="top" rowspan="1" colspan="1">JF417526</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=398.68&link_type=cbs">CBS 398.68</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces leycettanus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 22469 = IMI 178525</td><td align="left" valign="top" rowspan="1" colspan="1">Coal spoil tip soil; Leycett, Staffordshire, England, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121672</td><td align="left" valign="top" rowspan="1" colspan="1">JF417435</td><td align="left" valign="top" rowspan="1" colspan="1">JF417402</td><td align="left" valign="top" rowspan="1" colspan="1">JF417535</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=348.51&link_type=cbs">CBS 348.51</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces luteus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 089305</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, UK</td><td align="left" valign="top" rowspan="1" colspan="1">JN121656</td><td align="left" valign="top" rowspan="1" colspan="1">JF417429</td><td align="left" valign="top" rowspan="1" colspan="1">JF417396</td><td align="left" valign="top" rowspan="1" colspan="1">JF417529</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=475.71&link_type=cbs">CBS 475.71</ext-link><sup>IsoT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces purpureus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546</td><td align="left" valign="top" rowspan="1" colspan="1">Soil, near Esterel, France</td><td align="left" valign="top" rowspan="1" colspan="1">JN121687</td><td align="left" valign="top" rowspan="1" colspan="1">JN121522</td><td align="left" valign="top" rowspan="1" colspan="1">JN121828</td><td align="left" valign="top" rowspan="1" colspan="1">JN121919</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces stipitatus<sup><xref ref-type="table-fn" rid="TFN1">1</xref></sup></italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10500 (= NRRL 1006 = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=375.48&link_type=cbs">CBS 375.48</ext-link> = IMI 39805)</td><td align="left" valign="top" rowspan="1" colspan="1">Rotting wood; Louisiana, USA</td><td align="left" valign="top" rowspan="1" colspan="1">Unpublished</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=236.58&link_type=cbs">CBS 236.58</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces thermophilus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10518 = IMI 048593 = NRRL 2155</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Parthenium argentatum</italic>, decaying plant; California, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121611</td><td align="left" valign="top" rowspan="1" colspan="1">JF417420</td><td align="left" valign="top" rowspan="1" colspan="1">JF417387</td><td align="left" valign="top" rowspan="1" colspan="1">JF417520</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=373.48&link_type=cbs">CBS 373.48</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces trachyspermus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10497 = IMI 040043 = NRRL 1028</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121664</td><td align="left" valign="top" rowspan="1" colspan="1">JF417432</td><td align="left" valign="top" rowspan="1" colspan="1">JF417399</td><td align="left" valign="top" rowspan="1" colspan="1">JF4174532</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=391.48&link_type=cbs">CBS 391.48</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces wortmanii</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 10517 = IMI 040047 = NRRL 1017</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source</td><td align="left" valign="top" rowspan="1" colspan="1">JN121669</td><td align="left" valign="top" rowspan="1" colspan="1">JF417433</td><td align="left" valign="top" rowspan="1" colspan="1">JF417400</td><td align="left" valign="top" rowspan="1" colspan="1">JF417533</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=891.70&link_type=cbs">CBS 891.70</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus aurantiacus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 173037</td><td align="left" valign="top" rowspan="1" colspan="1">Wood; Firenze, Italy</td><td align="left" valign="top" rowspan="1" colspan="1">JN121719</td><td align="left" valign="top" rowspan="1" colspan="1">JF417444</td><td align="left" valign="top" rowspan="1" colspan="1">JF417410</td><td align="left" valign="top" rowspan="1" colspan="1">JF417544</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=396.78&link_type=cbs">CBS 396.78</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus aurantiacus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">JCM 12816</td><td align="left" valign="top" rowspan="1" colspan="1">Sawdust, in lumber yard, Toronto, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121671</td><td align="left" valign="top" rowspan="1" colspan="1">JN121512</td><td align="left" valign="top" rowspan="1" colspan="1">JN121818</td><td align="left" valign="top" rowspan="1" colspan="1">JN121912</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=181.67&link_type=cbs">CBS 181.67</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus crustaceus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16462 = IMI 126333</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Parthenium argentatum</italic>, decaying plant; Salinas, California, USA</td><td align="left" valign="top" rowspan="1" colspan="1">JN121591</td><td align="left" valign="top" rowspan="1" colspan="1">JF417417</td><td align="left" valign="top" rowspan="1" colspan="1">JF417384</td><td align="left" valign="top" rowspan="1" colspan="1">JF417517</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=528.71&link_type=cbs">CBS 528.71</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus themophilus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 123298 = NRRL 5208</td><td align="left" valign="top" rowspan="1" colspan="1">Wood and bark of <italic>Pinus</italic>; Sweden</td><td align="left" valign="top" rowspan="1" colspan="1">JN121697</td><td align="left" valign="top" rowspan="1" colspan="1">JF417442</td><td align="left" valign="top" rowspan="1" colspan="1">JF417408</td><td align="left" valign="top" rowspan="1" colspan="1">JF417542</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=218.34&link_type=cbs">CBS 218.34</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermomyces lanuginosus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">MUCL 8338</td><td align="left" valign="top" rowspan="1" colspan="1">Fruit shell of <italic>Theobroma</italic> cacao</td><td align="left" valign="top" rowspan="1" colspan="1">JN121599</td><td align="left" valign="top" rowspan="1" colspan="1">JF417418</td><td align="left" valign="top" rowspan="1" colspan="1">JF417385</td><td align="left" valign="top" rowspan="1" colspan="1">JF417518</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=224.63&link_type=cbs">CBS 224.63</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermomyces lanuginosus</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">MUCL 8337</td><td align="left" valign="top" rowspan="1" colspan="1">Mushroom compost; Gossau-Zürich Switzerland</td><td align="left" valign="top" rowspan="1" colspan="1">JN121602</td><td align="left" valign="top" rowspan="1" colspan="1">JF417419</td><td align="left" valign="top" rowspan="1" colspan="1">JF417386</td><td align="left" valign="top" rowspan="1" colspan="1">JF417519</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=334.68&link_type=cbs">CBS 334.68</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thysanophora canadensis</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18741 = IMI 137644 = MUCL 21216</td><td align="left" valign="top" rowspan="1" colspan="1">Needle of <italic>Tsuga canadensis</italic>, Bell's Corners, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121647</td><td align="left" valign="top" rowspan="1" colspan="1">JN121493</td><td align="left" valign="top" rowspan="1" colspan="1">JN121799</td><td align="left" valign="top" rowspan="1" colspan="1">JN121971</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=206.57&link_type=cbs">CBS 206.57</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thysanophora taxi</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 18484 = MUCL 11402</td><td align="left" valign="top" rowspan="1" colspan="1">Litter, Berlin, Germany</td><td align="left" valign="top" rowspan="1" colspan="1">JN121597</td><td align="left" valign="top" rowspan="1" colspan="1">JN121454</td><td align="left" valign="top" rowspan="1" colspan="1">JN121761</td><td align="left" valign="top" rowspan="1" colspan="1">JN121942</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.65&link_type=cbs">CBS 185.65</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Torulomyces lagena</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">MUCL 8221</td><td align="left" valign="top" rowspan="1" colspan="1">Bog soil under <italic>Thuja plicata</italic>, Guelph, Ontario, Canada</td><td align="left" valign="top" rowspan="1" colspan="1">JN121593</td><td align="left" valign="top" rowspan="1" colspan="1">JN121450</td><td align="left" valign="top" rowspan="1" colspan="1">JN121757</td><td align="left" valign="top" rowspan="1" colspan="1">JN121866</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.57&link_type=cbs">CBS 247.57</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma paradoxa</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">MUCL 39666 = IBT 31159</td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source; Hachijô, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121617</td><td align="left" valign="top" rowspan="1" colspan="1">JF417421</td><td align="left" valign="top" rowspan="1" colspan="1">JF417388</td><td align="left" valign="top" rowspan="1" colspan="1">JF417521</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=103.73&link_type=cbs">CBS 103.73</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma paradoxa</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Unknown source, Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121558</td><td align="left" valign="top" rowspan="1" colspan="1">JN121417</td><td align="left" valign="top" rowspan="1" colspan="1">JN121724</td><td align="left" valign="top" rowspan="1" colspan="1">JN121872</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=788.83&link_type=cbs">CBS 788.83</ext-link></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma paradoxa</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Rotting stump of cut down tree, Myojoji Temple near Hakui Noto Park, Ishikawa Pref., Japan</td><td align="left" valign="top" rowspan="1" colspan="1">JN121718</td><td align="left" valign="top" rowspan="1" colspan="1">JN121550</td><td align="left" valign="top" rowspan="1" colspan="1">JN121856</td><td align="left" valign="top" rowspan="1" colspan="1">JN121941</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=512.65&link_type=cbs">CBS 512.65</ext-link><sup>NT</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Warcupiella spinulosa</italic><sup><xref ref-type="table-fn" rid="TFN2">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1">ATCC 16919 = IMI 075885 = NRRL 4376</td><td align="left" valign="top" rowspan="1" colspan="1">Jungle soil; Berakas-Muara, Brunei</td><td align="left" valign="top" rowspan="1" colspan="1">JN121692</td><td align="left" valign="top" rowspan="1" colspan="1">JF417441</td><td align="left" valign="top" rowspan="1" colspan="1">JF417407</td><td align="left" valign="top" rowspan="1" colspan="1">JF417541</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=236.71&link_type=cbs">CBS 236.71</ext-link><sup>T</sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Xeromyces bisporus</italic><xref ref-type="table-fn" rid="TFN2">*</xref><xref ref-type="table-fn" rid="TFN1">1</xref></td><td align="left" valign="top" rowspan="1" colspan="1">IMI 063718</td><td align="left" valign="top" rowspan="1" colspan="1">Mouldy stick of liquorice, Homebush, New South Wales, Australia</td><td align="left" valign="top" rowspan="1" colspan="1">JN121612</td><td align="left" valign="top" rowspan="1" colspan="1">JN121466</td><td align="left" valign="top" rowspan="1" colspan="1">JN121773</td><td align="left" valign="top" rowspan="1" colspan="1">JN121898</td></tr></tbody></table><table-wrap-foot><fn id="TFN1"><label>1</label><p id="P19">Sequences derived from published full genome data.</p></fn><fn id="TFN2"><label>*</label><p id="P20">Strains used in the study of <italic>Trichocomaceae</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>)</p></fn><fn id="TFN3"><label>**</label><p id="P21">Strains used in for the preparation of Figs <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F7">7</xref>. CBS, culture collection of the CBS-KNAW Fungal Biodiversity Centre, Utrecht, Netherlands (WDCM 133) <uri xlink:type="simple" xlink:href="http://www.cbs.knaw.nl/databases/index.htm">http://www.cbs.knaw.nl/databases/index.htm</uri>; DTO, internal culture collection of CBS-KNAW Fungal Biodiversity Centre; IMI, CABI Genetic Resources Collection, Surrey, UK (WDCM 214) <uri xlink:type="simple" xlink:href="http://www.cabi.org/">http://www.cabi.org/</uri>; IBT, culture collection of Center for Microbial Biotechnology (CMB) at Department of Systems Biology, Technical University of Denmark (WDCM 758) <uri xlink:type="simple" xlink:href="http://www.biocentrum.dtu.dk/">http://www.biocentrum.dtu.dk/</uri>; NRRL, ARS Culture Collection, U.S. Department of Agriculture, Peoria, Illinois, USA (WDCM 97) <uri xlink:type="simple" xlink:href="http://nrrl.ncaur.usda.gov/">http://nrrl.ncaur.usda.gov/</uri>; ATCC, American Type Culture Collection, Manassas, VA, USA (WDCM 1) <uri xlink:type="simple" xlink:href="http://www.atcc.org/">http://www.atcc.org/</uri>; MUCL, Mycotheque de l'Universite catholique de Louvain, Leuven, Belgium (WDCM 308).</p></fn></table-wrap-foot></table-wrap></sec><sec id="S5"><title>DNA extraction, amplification and sequencing</title><p id="P22">Genomic DNA was extracted using the Ultraclean Microbial DNA isolation kit (MoBio Laboratories, Carlsbad, CA, USA), according to the manufacturer's instructions. Parts of the following loci were amplified and sequenced for the species listed in <xref ref-type="table" rid="T1">Table 1</xref>: 1. <italic>RPB1</italic>, RNA polymerase II largest subunit (regions E and F; according <xref ref-type="bibr" rid="R94">Matheny <italic>et al.</italic> 2002</xref>), 2. <italic>RPB2</italic>, RNA polymerase II second largest subunit (regions 5–7), 3. <italic>Cct8</italic>, subunit of the cytosolic chaperonin Cct ring complex, related to Tcp1p and required for the assembly of actin and tubulins <italic>in vivo</italic> (<xref ref-type="bibr" rid="R162">Stoldt <italic>et al.</italic> 1996</xref>, <xref ref-type="bibr" rid="R67">Kim <italic>et al.</italic> 1994</xref>), 4. <italic>Tsr1</italic>, protein required for processing of 20S pre-rRNA in the cytoplasm (<xref ref-type="bibr" rid="R46">Gelperin <italic>et al.</italic> 2001</xref>, <xref ref-type="bibr" rid="R78">Léger-Silvestre <italic>et al.</italic> 2004</xref>). Partial <italic>RPB2</italic> data was obtained for the majority of species listed in Table S1. Exceptions are strains used in the study of Houbraken <italic>et al</italic>. (<xref ref-type="bibr" rid="R59">2011c</xref>); in that case, published partial β-tubulin sequences were used.</p><p id="P23">The <italic>RPB1</italic> fragment was amplified using the primer pair <italic>RPB1</italic>-F1843 and R3096, and <italic>RPB1</italic>-R2623 was occasionally used as an internal primer for sequencing. A part of the <italic>RPB2</italic> locus was amplified using the primer pair <italic>RPB2</italic>-5F and <italic>RPB2</italic>-7CR (<xref ref-type="bibr" rid="R81">Liu <italic>et al.</italic> 1999</xref>) or the primer pair <italic>RPB2</italic>-5F_Eur and <italic>RPB2</italic>-7CR_Eur. The internal sequencing primers <italic>RPB2</italic>-F311 and <italic>RPB2</italic>-R310 were occasionally used when poor results were obtained with the regular forward and reverse primers. Amplification of a part of the <italic>Cct8</italic> gene was performed using the primer pair <italic>Cct8</italic>-F660 and <italic>Cct8</italic>-R1595. No amplicons could be obtained in the case of 5–10 % of the analysed strains. In those cases, amplicons were generated using the primer pair <italic>Cct8</italic>-R1595 and <italic>Cct8</italic>-F94. A part of the <italic>Tsr1</italic> gene was amplified using the forward primers <italic>Tsr1</italic>-F1526Pc or <italic>Tsr1</italic>-F1526 in combination with <italic>Tsr1</italic>-R2434. Annealing temperatures and primers used for amplification and sequencing are shown in <xref ref-type="table" rid="T2">Table 2</xref>.</p><table-wrap id="T2" position="float"><label>Table 2</label><caption><p>Primers used in this study for amplification and sequencing.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top" rowspan="1" colspan="1">Locus</th><th align="left" valign="top" rowspan="1" colspan="1">Primer</th><th align="left" valign="top" rowspan="1" colspan="1">Sequence (5′-3′)</th><th align="left" valign="top" rowspan="1" colspan="1">Annealing (°C)</th><th align="left" valign="top" rowspan="1" colspan="1">Fragment size (bp)</th><th align="left" valign="top" rowspan="1" colspan="1">References</th></tr></thead><tbody><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cct</italic>8</td><td align="left" valign="top" rowspan="1" colspan="1">F94</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) CGCAAC AAGATYGTBATYAACCA</td><td align="left" valign="top" rowspan="1" colspan="1">50–52</td><td align="left" valign="top" rowspan="1" colspan="1">F94-R1595: 1400–1450</td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">F660</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) GIGTKGTBAAGATCATGGGWGG</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">F660-R1595: 850–890</td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">R1595</td><td align="left" valign="top" rowspan="1" colspan="1">(Rev) RTCMACRCCNGTIGTCCAGTA</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB1</italic></td><td align="left" valign="top" rowspan="1" colspan="1">F1843</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) ATTTYGAYGGTGAYGARATGAAC</td><td align="left" valign="top" rowspan="1" colspan="1">48–53</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>ca</italic>. 1000</td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">R3096</td><td align="left" valign="top" rowspan="1" colspan="1">(Rev) GRACRGTDCCRTCATAYTTRACC</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">R2623</td><td align="left" valign="top" rowspan="1" colspan="1">GCRTTGTTSARATCCTTMARRCTC</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></td><td align="left" valign="top" rowspan="1" colspan="1">5F</td><td align="left" valign="top" rowspan="1" colspan="1">GAYGAYMGWGATCAYTTYGG</td><td align="left" valign="top" rowspan="1" colspan="1">48–51</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>ca</italic>. 1220</td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R81">Liu <italic>et al</italic>. 1999</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">7CR</td><td align="left" valign="top" rowspan="1" colspan="1">CCCATRGCTTGYTTRCCCAT</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R81">Liu <italic>et al</italic>. 1999</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">5F_Eur</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) GAYGAYCGKGAYCAYTTCGG</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">7CR_Eur</td><td align="left" valign="top" rowspan="1" colspan="1">(Rev) CCCATRGCYTGYTTRCCCAT</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">F311</td><td align="left" valign="top" rowspan="1" colspan="1">CATGATYCARCGIAAYATGGA</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">R310</td><td align="left" valign="top" rowspan="1" colspan="1">CCATRTTICGYTGRATCATGAA</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Tsr</italic>1</td><td align="left" valign="top" rowspan="1" colspan="1">F1526Pc</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) GARTAYCCBCARTCNGAGATGT</td><td align="left" valign="top" rowspan="1" colspan="1">48–50</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>ca</italic>. 820</td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">F1626</td><td align="left" valign="top" rowspan="1" colspan="1">(Fwd) GARTAYCCBCARTCNGAIATGT</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">This study</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">R2434</td><td align="left" valign="top" rowspan="1" colspan="1">(Rev) ASAGYTGVARDGCCTTRAACCA</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref></td></tr></tbody></table></table-wrap><p id="P24">The PCR reactions were performed in 25 μL reaction mixtures containing 1 μL genomic DNA 2.5 μL PCR buffer, 0.75 μL MgCl<sub>2</sub> (50 mM), 16.55 μL demineralised sterile water, 1.85 μL dNTP (1 mM), 0.50 μL of each primer (100 mM) and 0.1 μL Taq polymerase (5 U/μL, BioTaq, Bioline). The PCR program typically was: 5 cycles of 30 s denaturation at 94 °C, followed by primer annealing for 30 s at 51 °C, and extension for 1 min at 72 °C; followed by 5 cycles with an annealing temperature at 49 °C and 30 cycles at 47 °C, finalised with an extension for final 10 min at 72 °C. Excess primers and dNTP's were removed from the PCR product using the QIAQuick PCR purification kit (Qiagen). Purified PCR fragments were resuspended in 30–50 μL of water. PCR products were sequenced directly in both directions with the same primers and DYEnamic ET Terminator Cycle Sequencing Kit (Amersham Bioscience, Roosendaal, The Netherlands). The cycle sequencing reaction mixture had a total reaction volume of 10 μL, and contained 1 μL of template DNA, 0.85 μL BigDye reagent, 3 μL buffer, 4.75 μL demineralised water and 0.4 μL primer (10 mM).</p><p id="P25">Sequencing products were purified according to the manufacturers' recommendations with Sephadex G-50 superfine columns (Amersham Bioscience, Roosendaal, The Netherlands) in a multiscreen HV plate (Millipore, Amsterdam, The Netherlands) and with MicroAmp Optical 96-well reaction plate (AB Applied Biosystems, Nieuwerkerk a/d Yssel, The Netherlands). Contigs were assembled using the forward and reverse sequences with the programme SeqMan from the LaserGene package (DNAStar Inc., Madison, WI).</p></sec><sec id="S6"><title>Phylogenetic analysis</title><p id="P26">The protein coding nucleotide sequences were translated into amino acid data prior to alignment and subsequently aligned using the Muscle software in the MEGA5 package. After aligning, the amino acid data were translated into nucleotide data and used in the phylogenetic analysis. Combined sequence data sets were used in the study on the phylogeny of <italic>Trichocomaceae</italic> and <italic>Penicillium</italic>. Before combining the data sets, each data set was analysed using RAxML (<xref ref-type="bibr" rid="R159">Stamatakis <italic>et al.</italic> 2008</xref>). The number of bootstrap runs was set to 100. The program compat.py (from <uri xlink:type="simple" xlink:href="http://www.lutzonilab.net">http://www.lutzonilab.net</uri>) was used to detect major topological incongruences among single gene data sets (<xref ref-type="bibr" rid="R64">Kauff & Lutzoni 2002</xref>). Conflicts were considered significant when a sequence was differentially resolved between two gene trees with greater than 70 % bootstrap support. If no conflicts were detected, then the data sets were combined.</p><p id="P27">Statistical support was measured by Maximum Likelihood (ML) analysis using the RAxML (randomised axelerated maximum likelihood) software (Stamakis <italic>et al.</italic> 2008). The robustness of trees in the ML analyses was evaluated by 1000 bootstrap replications. A second measure for statistical support was performed by Bayesian tree inference (BI) analysis using MrBayes v. 3.1.2 (<xref ref-type="bibr" rid="R137">Ronquist & Huelsenbeck 2003</xref>). Prior to analysis, the most suitable substitution model was determined using MrModeltest v. 2.3 (<xref ref-type="bibr" rid="R105">Nylander 2004</xref>), utilising the Akaike Information Criterion (AIC). The Bayesian analysis was performed with two sets of four chains (one cold and three heated) and the stoprule option, stopping the analysis at an average standard deviation of split frequencies of 0.01. The sample frequency was set to 100; the first 25 percent of trees were removed as burnin. The phylograms obtained with the RAxML analysis were used for presenting the data. Bootstrap values lower than 70 % were considered unreliable because their wide range of error and Bayesian posterior probabilities are considered unreliable below 0.95 (<xref ref-type="bibr" rid="R101">Murphy <italic>et al</italic>. 2001</xref>, <xref ref-type="bibr" rid="R198">Wilcox <italic>et al</italic>. 2002</xref>, <xref ref-type="bibr" rid="R3">Alfaro & Holder 2006</xref>). Therefore, only posterior probability (pp) values higher than 0.95 and bootstrap (bs) values higher than 70 % were plotted on those phylograms. <italic>Coccidioides immitis</italic> (strain RS), a member of <italic>Onygenales</italic>, was chosen to root the phylogram used in the study on the relationships of <italic>Penicillium</italic> species among <italic>Trichocomaceae. Penicillium</italic> (= <italic>Talaromyces</italic>) <italic>marneffei</italic> ATCC 18227T was selected as an outgroup for the analysis of the phylogeny of <italic>Penicillium</italic>. Various phylograms were prepared for assignment of species to sections. All data sets were based on partial <italic>RPB2</italic> sequences and rooted with <italic>Talaromyces flavus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup>, with exception of the phylogram of sections <italic>Lanata-divaricata</italic> and <italic>Stolkia</italic>, which is based on partial β-tubulin data. <italic>Penicillium glabrum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125543&link_type=cbs">CBS 125543</ext-link><sup>T</sup> was used as an outgroup.</p></sec></sec><sec id="S7"><title>RESULTS</title><sec id="S8"><title>Phylogeny of <italic>Trichocomaceae</italic></title><p id="P28">A phylogenetic study using four combined loci (<italic>RPB1, RPB2, Cct8</italic> and <italic>Tsr1</italic>) was conducted to determine the relationship among members of <italic>Trichocomaceae</italic>. A total of 157 species were included in the analysis and the total length of the alignment was 3 111 characters, 1 939 of those characters were variable. The length of the <italic>Cct8, Tsr1, RPB1</italic> and <italic>RPB2</italic> partitions were 714, 669, 768, 960 base pairs long, respectively. The GTR+I+G model was optimal for all four partitions.</p><p id="P29">The result of the analysis is shown in <xref ref-type="fig" rid="F1">Fig. 1</xref> and indicates that <italic>Trichocomaceae</italic> can be divided into three lineages. Lineage 1 is divided into seven clades (clades 1–7) and these clades are on a well-supported branch (100 % bs, 1.00 pp). The type species of the genera <italic>Chromocleista</italic> (<italic>C. malachitea</italic>), <italic>Eladia</italic> (<italic>E. saccula</italic>),<italic>Eupenicillium</italic> (<italic>E. crustaceum</italic>), <italic>Hemicarpenteles</italic> (<italic>H. paradoxus</italic>), <italic>Penicillium</italic> (<italic>P. expansum</italic>), <italic>Thysanophora</italic> (<italic>T. penicillioides</italic>) and <italic>Torulomyces</italic> (<italic>T. lagena</italic>) belong to clade 1. This clade is named <italic>Penicillium sensu stricto</italic> and is divided into two subclades: clade 1A and 1B. The types of subgenera <italic>Aspergilloides</italic> and <italic>Furcatum</italic> are accommodated in clade 1A and the type of subgenus <italic>Penicillium</italic> belongs to clade 1B. Clade 2 is moderately supported (< 70 % bs, 1.00 pp) and contains the type species of the genera <italic>Aspergillus</italic> (<italic>A. glaucus</italic>), <italic>Cristaspora</italic> (<italic>C. arxii</italic>), <italic>Phialosimplex</italic> (<italic>P. caninus</italic>), <italic>Polypaecilum</italic> (<italic>P. insolitum</italic>) and the teleomorphs of <italic>Aspergillus</italic> (<italic>Fennellia, Eurotium, Emericella, Neocarpenteles, Dichotomyces, Neosartorya, Sclerocleista</italic>). Not all teleomorph genera of <italic>Aspergillus</italic> are represented in our analysis; however, previous data has shown the genera <italic>Chaetosartorya, Neopetromyces</italic> and <italic>Petromyces</italic> also belong to this lineage (<xref ref-type="bibr" rid="R115">Peterson 2008</xref>). This clade is subdivided into six groups. Four of the six groups represent the <italic>Aspergillus</italic> subgenera as defined by Peterson (<xref ref-type="bibr" rid="R115">2008</xref>). In addition, also <italic>Aspergillus</italic> section <italic>Cremei</italic> and a clade with <italic>Phialosimplex</italic> and <italic>Polypaecilum</italic> are present. Clade 3 comprises the type species of <italic>Hamigera</italic> (<italic>H. avellanea</italic>), <italic>Warcupiella</italic> (<italic>W. spinosa</italic>) and <italic>Raperia</italic> (<italic>R. spinulosa</italic>) but this clade is poorly supported (< 70 % bs, < 0.95 pp). Clade 4 contains <italic>P. clavariiformis</italic>, the type species <italic>Penicilliopsis</italic>. The type species of the genera <italic>Basipetospora</italic> (<italic>B. rubra</italic>), <italic>Fraseriella</italic> (<italic>F. bisporus</italic>), <italic>Leiothecium</italic> (<italic>L. ellipsoideum</italic>), <italic>Monascus</italic> (<italic>M. ruber</italic>), <italic>Xeromyces</italic> (<italic>X. bisporus</italic>) cluster together in clade 5. <italic>Phialomyces</italic> (<italic>P. macrosporus</italic>) and <italic>Sclerocleista</italic> (<italic>S. ornata</italic>) belong to clade 6 and 7, respectively. Lineage 2 is subdivided into two clades: the type species of <italic>Thermoascus, Coonemeria</italic> and <italic>Dactylomyces</italic> belong to clade 8, and the types of the genera <italic>Byssochlamys</italic> (<italic>B. nivea</italic>) and <italic>Paecilomyces</italic> (<italic>P. variotii</italic>) belong to clade 9. The posterior probability value indicates a strong relationship between these two clades (0.99); however, the maximum likelihood analysis resulted in a bootstrap value lower than 70 % (67 %). The posterior probability and bootstrap values are also contradictory regarding the relationship between lineages 1 and 2 (< 70 % bs, 1.00 pp). Lineage 3 is subdivided into five clades (clades 10–14) and these clades are on a strongly supported branch (100 % bs, 1.00 pp). Clade 10 is centered on the type species of <italic>Talaromyces, T. flavus</italic>, and the type species of <italic>Sagenoma</italic> (<italic>S. viride</italic>) also belongs in this clade. The type species of <italic>Thermomyces</italic> (<italic>T. lanuginosus</italic>), <italic>Sagenomella</italic> (<italic>S. diversispora</italic>), <italic>Rasamsonia</italic> (<italic>R. emersonii</italic>) and<italic> Trichocoma</italic> (<italic>T. paradoxa</italic>) belong in clades 11–14, respectively.</p><fig id="F1" position="float"><label>Fig. 1.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on combined data set of partial <italic>Cct8</italic>, <italic>Tsr1, RPB1 and RPB2</italic> sequences showing the relationship among members of <italic>Trichocomaceae</italic>. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 50 % supported in the ML or less than 0.90 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate full support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Coccidioides immitis</italic> (strain RS).</p></caption><graphic xlink:href="1fig1a"></graphic><graphic xlink:href="1fig1b"></graphic></fig></sec><sec id="S9"><title>Phylogeny of <italic>Penicillium sensu stricto</italic></title><p id="P30">The phylogenetic relationship among members of <italic>Penicillium s. str.</italic> was studied using the same four combined loci (<italic>RPB1, RPB2, Cct8</italic> and <italic>Tsr1</italic>). In total, 72 strains were included in the analysis and the total length of the alignment was 3 393 characters, and 1 805 of them were variable. <italic>Penicillium</italic> (= <italic>Talaromyces</italic>) <italic>marneffei</italic> was used as an outgroup. The length of the <italic>Cct8, Tsr1, RPB1</italic> and <italic>RPB2</italic> partitions were 723, 759, 955, 957 base pairs, respectively. The best-fit model GTR+I+G was optimal for all four partitions. The result of the analysis is shown in <xref ref-type="fig" rid="F7">Fig. 7</xref> and confirms the result above that <italic>Penicillium s. str.</italic> can be divided into two distinct lineages. Similarly, the type species of subgenus <italic>Aspergilloides, P. aurantiobrunneum</italic> (= <italic>P. glabrum</italic>) and <italic>Furcatum</italic> (<italic>P. citrinum</italic>), belong to lineage 1 and the type of subgenus <italic>Penicillium</italic> belongs to lineage 2. Lineage 1 is subdivided in 14 clades (<xref ref-type="fig" rid="F7">Fig. 7</xref>). These clades (1–14) were in most cases supported with a bootstrap value higher than 95 % and a posterior probability of 1.00. Lineage 2 is subdivided into 11 clades (15–25). Clades 20–25 are on well-supported branches; however, the overall bootstrap and posterior probability values of clades 15–19 are low. The numbering of the clades is therefore based on the analysis of the partial β-tubulin data in Samson <italic>et al</italic>. (<xref ref-type="bibr" rid="R147">2004</xref>), because well-supported clades (sections) were present in that phylogenetic treatment. Five separate phylograms (Figs <xref ref-type="fig" rid="F8">8</xref>, <xref ref-type="fig" rid="F10">10</xref>, <xref ref-type="fig" rid="F11">11</xref>, <xref ref-type="fig" rid="F12">12</xref>, <xref ref-type="fig" rid="F13">13</xref>) were prepared in order to determine which species belong to which clade (section). Details of these analyses are summarised in <xref ref-type="table" rid="T3">Table 3</xref>.</p><fig id="F7" position="float"><label>Fig. 7.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on combined data set of partial <italic>Cct8</italic>, <italic>Tsr1, RPB1 and RPB2</italic> sequences showing the relationship among members of <italic>Penicillium</italic> <italic>s</italic>. <italic>str</italic>. <italic>Penicillium</italic> <italic>s</italic>. <italic>str</italic>. is divided in two lineages (s/g <italic>Aspergilloides</italic> and <italic>Penicillium</italic>) and 25 sections. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (bs/pp). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Penicillium</italic> (= <italic>Talaromyces</italic>) <italic>marneffei</italic> ATCC 18227<sup>T</sup>.</p></caption><graphic xlink:href="1fig7"></graphic></fig><fig id="F8" position="float"><label>Fig. 8.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on partial <italic>RPB2</italic> sequences and giving an overview of the members accommodated in sections <italic>Aspergilloides</italic>, <italic>Sclerotiora</italic> and <italic>Charlesii</italic>. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Talaromyces flavus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup>.</p></caption><graphic xlink:href="1fig8"></graphic></fig><fig id="F10" position="float"><label>Fig. 10.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on partial <italic>RPB2</italic> sequences and giving an overview of the members accommodated in sections <italic>Exilicaulis</italic>, <italic>Cinnamopurpurea</italic>, <italic>Ramigena</italic> and <italic>Gracilenta</italic>. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Talaromyces flavus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup>.</p></caption><graphic xlink:href="1fig10"></graphic></fig><fig id="F11" position="float"><label>Fig. 11.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on partial β-tubulin sequences and giving an overview of the members accommodated in sections <italic>Lanata-divaricata</italic> and <italic>Stolkia</italic>. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Penicillium glabrum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=125543&link_type=cbs">CBS 125543</ext-link><sup>T</sup>.</p></caption><graphic xlink:href="1fig11"></graphic></fig><fig id="F12" position="float"><label>Fig. 12.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on partial <italic>RPB2</italic> sequences and giving an overview of the members accommodated in sections <italic>Citrina</italic> and <italic>Ochrosalmonea</italic>. The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Talaromyces flavus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup>.</p></caption><graphic xlink:href="1fig12"></graphic></fig><fig id="F13" position="float"><label>Fig. 13.</label><caption><p>Best-scoring Maximum Likelihood tree using RAxML based on partial <italic>RPB2</italic> sequences and giving an overview of the members accommodated in subgenus Penicillium (clades 15–25). The BI posterior probabilities (pp) values and bootstrap (bs) percentages of the maximum likelihood (ML) analysis are presented at the nodes (pp/bs). Values less than 70 % supported in the ML or less than 0.95 in the Bayesian analysis are indicated with a hyphen, whereas asterisks indicate good support (100 % bs or 1.00 pp). The branches with more than 95 % bootstrap support and 1.00 posterior probability values are thickened. The bar indicates the number of substitutions per site. The tree is rooted with <italic>Talaromyces flavus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=310.38&link_type=cbs">CBS 310.38</ext-link><sup>NT</sup>.</p></caption><graphic xlink:href="1fig13a"></graphic><graphic xlink:href="1fig13b"></graphic></fig><table-wrap id="T3" position="float"><label>Table 3</label><caption><p>Details of each analysis of the data sets used for generating Figs 8, 10–13.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top" rowspan="1" colspan="1">Figure</th><th align="left" valign="top" rowspan="1" colspan="1">Clades, acc. Fig. 7</th><th align="left" valign="top" rowspan="1" colspan="1">Locus</th><th align="left" valign="top" rowspan="1" colspan="1">No. isolates</th><th align="left" valign="top" rowspan="1" colspan="1">Length alignment</th><th align="left" valign="top" rowspan="1" colspan="1">Best-fit model</th></tr></thead><tbody><tr><td align="left" valign="top" rowspan="1" colspan="1">8</td><td align="left" valign="top" rowspan="1" colspan="1">1, 2, 3</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></td><td align="left" valign="top" rowspan="1" colspan="1">50</td><td align="left" valign="top" rowspan="1" colspan="1">916</td><td align="left" valign="top" rowspan="1" colspan="1">SYM+I+G</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">10</td><td align="left" valign="top" rowspan="1" colspan="1">6, 7, 10 and 13</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></td><td align="left" valign="top" rowspan="1" colspan="1">69</td><td align="left" valign="top" rowspan="1" colspan="1">916</td><td align="left" valign="top" rowspan="1" colspan="1">GTR+I+G</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">11</td><td align="left" valign="top" rowspan="1" colspan="1">11, 12</td><td align="left" valign="top" rowspan="1" colspan="1">β-tubulin</td><td align="left" valign="top" rowspan="1" colspan="1">45</td><td align="left" valign="top" rowspan="1" colspan="1">528</td><td align="left" valign="top" rowspan="1" colspan="1">HKY+I+G</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">12</td><td align="left" valign="top" rowspan="1" colspan="1">5, 14</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></td><td align="left" valign="top" rowspan="1" colspan="1">44</td><td align="left" valign="top" rowspan="1" colspan="1">849</td><td align="left" valign="top" rowspan="1" colspan="1">SYM+I+G</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">13</td><td align="left" valign="top" rowspan="1" colspan="1">15–25</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>RPB2</italic></td><td align="left" valign="top" rowspan="1" colspan="1">86</td><td align="left" valign="top" rowspan="1" colspan="1">916</td><td align="left" valign="top" rowspan="1" colspan="1">GTR+I+G</td></tr></tbody></table></table-wrap></sec></sec><sec id="S10"><title>DISCUSSION</title><sec id="S11"><title>Part one: Phylogenetic analysis of <italic>Trichocomaceae</italic></title><sec id="S12"><title>Choice of genes</title><p id="P31">Parts of the <italic>RPB1, RPB2, Tsr1</italic> and <italic>Cct8</italic> genes were only used for the construction of the phylogenetic relationships among members of <italic>Trichocomaceae</italic> and <italic>Penicillium</italic> species, and the ability of these genes for species recognition remains largely unexplored. The regions E and F (according <xref ref-type="bibr" rid="R94">Matheny <italic>et al</italic>. 2002</xref>) of the <italic>RPB1</italic> gene were analysed. No additional sequence data of <italic>Trichocomaceae</italic> were published on this part of the <italic>RPB1</italic> gene and comparison with other studies is therefore difficult. The regions 5–7 of the <italic>RPB2</italic> gene are commonly used in taxonomic studies of <italic>Penicillium</italic> and <italic>Aspergillus</italic> and proved to be a good marker for species recognition (<italic>e.g.</italic> <xref ref-type="bibr" rid="R115">Peterson 2008</xref>, Serra <italic>et al</italic>. 2008, <xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>, <xref ref-type="bibr" rid="R119">Peterson <italic>et al</italic>. 2010</xref>, <xref ref-type="bibr" rid="R12">Barreto <italic>et al</italic>. 2011</xref>). However, <italic>RPB1</italic> and <italic>RPB2</italic>, as well as TEF1α, β-tubulin, and γ-actin, were not found among the best performing genes for fungal systematics (<xref ref-type="bibr" rid="R2">Aguileta <italic>et al</italic>. 2008</xref>). Aguileta <italic>et al</italic>. (<xref ref-type="bibr" rid="R2">2008</xref>) studied, using a bioinformatics approach, the performance of single-copy protein-coding genes for fungal phylogenetics. Their analyses of 30 published fungal genomes revealed that <italic>MCM7</italic> (= MS456), <italic>Tsr1</italic> (= MS277) and <italic>Cct8</italic> (= FG610) were among the best single-copy genes in phylogenetic utility. <italic>MCM7</italic>, the best gene for recovering a larger-scale phylogeny across fungal groups, was excluded in the current study since it was not variable enough within the genus <italic>Penicillium</italic> (<xref ref-type="bibr" rid="R93">Marthey <italic>et al</italic>. 2008</xref>). <italic>Tsr1</italic> and <italic>Cct8</italic> were also used in other (phylogenetic) studies of groups belonging to <italic>Trichocomaceae</italic> (<xref ref-type="bibr" rid="R87">López-Villavicencio <italic>et al</italic>. 2010</xref>, <xref ref-type="bibr" rid="R119">Peterson <italic>et al</italic>. 2010</xref>). Analysis of the <italic>Tsr1</italic> gene generated the best resolved trees, when compared with <italic>Cct8, MCM7</italic> and ITS (<xref ref-type="bibr" rid="R87">López-Villavicencio <italic>et al</italic>. 2010</xref>). The sequenced parts of the <italic>RPB1, RPB2, Tsr1</italic> and <italic>Cct8</italic> genes mainly contain exons, and the alignment of these loci is therefore unambiguous. This is the main advantage over ITS regions where alignment above genus can be difficult. Furthermore, the ITS region is generally considered unreliable as a phylogenetic marker, especially above genus rank. β-tubulin and calmodulin sequences are often used in taxonomical studies of <italic>Penicillium, Paecilomyces</italic> and <italic>Aspergillus</italic> (<italic>e.g.</italic> <xref ref-type="bibr" rid="R147">Samson <italic>et al</italic>. 2004</xref>, <xref ref-type="bibr" rid="R54">Houbraken <italic>et al</italic>. 2007</xref>, <xref ref-type="bibr" rid="R145">Samson <italic>et al</italic>. 2009</xref>, <xref ref-type="bibr" rid="R191">Varga <italic>et al</italic>. 2011</xref>). However, a large part of these genes consists of intron data and these regions cannot be aligned above genus level, resulting in loss of information in these data sets. In addition, there is evidence that β-tubulins are present in the genome in multiple copies and thus have the potential of being phylogenetically misleading (<xref ref-type="bibr" rid="R75">Landvik <italic>et al</italic>. 2001</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>).</p></sec><sec id="S13"><title>Phylogenetic analysis of Trichocomaceae</title><p id="P32">Three lineages are recognised in <italic>Trichocomaceae</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>) and we propose to treat these three lineages as distinct families: <italic>Trichocomaceae, Aspergillaceae</italic> and <italic>Thermoascaceae</italic>. Lineage 1 corresponds with <italic>Aspergillaceae</italic> and this name is the oldest available family name within the analysed group of related genera. Malloch & Cain (<xref ref-type="bibr" rid="R92">1972</xref>) did not accept this family name since it was based on the asexual (anamorph) form-genus <italic>Aspergillus</italic> and therefore not applicable for ascomycete perfect (sexual) states. Because we are applying a single-name system and give priority to the oldest name, the family name <italic>Aspergillaceae</italic> is re-instated. Phylogenetically, <italic>Monascaceae</italic> belong to <italic>Aspergillaceae</italic> and this is in agreement with other studies that show that <italic>Monascus</italic> (type genus of <italic>Monascaceae</italic>) is related to <italic>Penicillium</italic> and/or <italic>Aspergillus</italic> (<xref ref-type="bibr" rid="R15">Berbee <italic>et al</italic>. 1995</xref>, <xref ref-type="bibr" rid="R106">Ogawa <italic>et al</italic>. 1997</xref>, <xref ref-type="bibr" rid="R107">Ogawa & Sugiyama 2000</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>, <xref ref-type="bibr" rid="R124">Pettersson <italic>et al</italic>. 2011</xref>). In contrast, Stchigel <italic>et al</italic>. (<xref ref-type="bibr" rid="R160">2004</xref>), who used ITS sequence data to determine the molecular relationships of <italic>Monascaceae</italic> taxa, concluded that <italic>Monascus</italic> and <italic>Xeromyces</italic> form a well-supported, monophyletic clade (81 % bs), separate from <italic>Eurotiales</italic> (<xref ref-type="bibr" rid="R161">Stchigel & Guarro 2007</xref>). These contradictory results can be explained by a deeper taxon sampling in this study combined with a phylogeny based on sequences of four protein-coding genes instead of ITS sequences alone. The <italic>Thermoascaceae</italic> (= lineage 2) were introduced by Apinis (<xref ref-type="bibr" rid="R5">1967</xref>) and typified by <italic>Thermoascus</italic>. Lineage 3 corresponds to <italic>Trichocomaceae</italic> and this family was introduced by Fischer (<xref ref-type="bibr" rid="R30">1897</xref>) (as <italic>Trichocomataceae</italic>) and is typified by <italic>Trichocoma</italic>. The <italic>Eurotiaceae</italic> were placed in synonymy with this family because the name <italic>Trichocomaceae</italic> predates <italic>Eurotiaceae</italic> (<xref ref-type="bibr" rid="R92">Malloch & Cain 1972</xref>). The current analysis shows that <italic>Eurotiaceae</italic> (type genus <italic>Eurotium</italic>) should be placed in synonymy with <italic>Aspergillaceae</italic>. The family names <italic>Hemicarpenteleaceae, Penicilliopsidaceae, Phialomycetaeae, Warcupiellaceae, Xeromycetaceae</italic> and <italic>Talaromycetaceae</italic> were introduced by Locquin (<xref ref-type="bibr" rid="R85">1972</xref>, <xref ref-type="bibr" rid="R86">1984</xref>) but all lack a Latin description and are invalidly published.</p></sec><sec id="S14"><title>Phenotypic classification and delimitation of Aspergillaceae, Trichocomaceae and Thermoascaceae</title><p id="P33">Several studies on the classification of <italic>Trichocomaceae</italic> and <italic>Eurotiales</italic> based on phenotypic characters were published (<xref ref-type="bibr" rid="R92">Malloch & Cain 1972</xref>, <xref ref-type="bibr" rid="R29">Fennell 1973</xref>, <xref ref-type="bibr" rid="R14">Benny & Kimbrough 1980</xref>, <xref ref-type="bibr" rid="R90">Malloch 1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>, von Arx 1987) and an overview of selected studies is shown in <xref ref-type="table" rid="T4">Table 4</xref>. Some of these classifications differ significantly from each other. We compared the results of these studies with the current proposed phylogenetic classification and this showed that our phylogenetic classification largely corresponds with the phenotypic classification described by Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>). Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>) divided <italic>Trichocomaceae</italic> into two subfamilies, <italic>Trichomoideae</italic> and <italic>Dichlaenoideae</italic>, based on phenotypic characters including cleistothecial initials, peridium, ascus structure and ascospore morphology. Malloch's list of genera belonging to <italic>Dichlaenoideae</italic> largely corresponds with the genera we place in <italic>Aspergillaceae</italic> and his definition of <italic>Trichomoideae</italic> is comparable with our phylogenetically defined <italic>Trichocomaceae</italic>. There are two main differences: a) <italic>Monascus</italic> is treated here in <italic>Aspergillaceae</italic> and b) the genera <italic>Byssochlamys</italic> and <italic>Thermoascus</italic> are accommodated in <italic>Thermoascaceae</italic>; these were treated by Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>) in <italic>Trichomoideae</italic> and <italic>Dichlaenoideae</italic>, respectively. Using the characters proposed by Malloch in his classification, <italic>Aspergillaceae</italic> are characterised by the production of asci inside cleistothecia, stromata, or are surrounded by Hülle cells and mainly have oblate to ellipsoidal ascospores with a furrow or slit. The conidia are mostly formed on flask shaped or cylindrical phialides. The <italic>Trichocomaceae</italic> are defined by having asci borne within a tuft or layer of loose hyphae, and ascospores are lacking slits or furrows. The phialides of species belonging to this family are mostly lanceolate or cylindrical. Apinis (<xref ref-type="bibr" rid="R5">1967</xref>) introduced <italic>Thermoascaceae</italic> and noted that the common essential character of genera of this family is the production of firm, somewhat sclerotioid, pseudoparenchymatous cleistothecia. The inclusion of <italic>Byssochlamys</italic> in this family does not fit in that description because it produces almost naked ascomata. Based on the relative branch length in <xref ref-type="fig" rid="F1">Fig. 1</xref>, another possibility would be to delimit the <italic>Thermoascus</italic> clade (clade 8) and the <italic>Byssochlamys</italic>/<italic>Paecilomyces</italic> clade (clade 9) as separate families. However, there are characters shared by <italic>Thermoascus</italic> and <italic>Byssochlamys</italic> including the production of asci in croziers and the formation of smooth or finely roughened ascospores lacking a furrow or slit. The relationship between these two genera is also illustrated by <italic>Byssochlamys verrucosa</italic> and <italic>Thermoascus crustaceus. Byssochlamys verrucosa</italic> phenotypically belongs to <italic>Byssochlamys</italic>, but is positioned phylogenetically in <italic>Thermoascus</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>) and <italic>Therm. crustaceus</italic> shares a <italic>Paecilomyces</italic> anamorph with members of the <italic>Byssochlamys</italic>/<italic>Paecilomyces</italic> clade. In addition, most members of both genera are thermotolerant or thermophilic.</p><table-wrap id="T4" position="float"><label>Table 4</label><caption><p>Overview of the classifications of the <italic>Trichocomaceae</italic> and <italic>Eurotiaceae</italic> by Benny & Kimbrough (1980), Malloch (1985b), von Arx (1987) and the current study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" valign="top" rowspan="1" colspan="1">Benny & Kimbrough (<xref ref-type="bibr" rid="R14">1980</xref>)</th><th align="left" valign="top" rowspan="1" colspan="1">von Arx (1987)</th><th align="left" valign="top" rowspan="1" colspan="1">Malloch (<xref ref-type="bibr" rid="R91">1985b</xref>)</th><th align="left" valign="top" rowspan="1" colspan="1">Current study</th></tr></thead><tbody><tr><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Trichocomaceae</italic>:</bold></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Eurotiaceae</italic>:</bold></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Trichomoideae</italic>:</bold></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Aspergillaceae</italic>:</bold></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aphanoascus</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Chaetosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Byssochlamys</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergillus</italic> (incl. teleomorphs, syn. <italic>Stilbothamnium</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Byssochlamys</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cristaspora</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dendrosphaera</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hamigera</italic> (incl. <italic>Merimbla</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Chaetosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichlaena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenoma</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Leiothecium</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichotomomyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Monascus</italic> (incl. <italic>Basipetospora</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichleana</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Emericella</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicilliopsis</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Edyuillia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eupenicillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Dichlaenoideae</italic>:</bold></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium</italic> (syn. <italic>Chromocleista, Eladia, Eupenicillium, Hemicarpenteles, Thysanophora, Torulomyces</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Emericella</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eurotium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Chaetosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialomyces</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eupenicillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fennellia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cristaspora</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Phialosimplex</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eurotium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemicarpenteles</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichlaena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Polypaecilum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fennellia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Mallochia</italic><sup><xref ref-type="table-fn" rid="TFN5">2</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichotomomyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerocleista</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hamigera</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Neosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eupenicillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Warcupiella</italic> (incl. <italic>Raperia</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemicarpenteles</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Saitoa</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Edyuillia</italic> (=<italic>Eurotium</italic>)</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Xeromyces</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemisartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Emericella</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Thermoascaceae</italic>:</bold></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Neosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eurotium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Paecilomyces</italic> (incl. <italic>Byssochlamys</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicilliopsis</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fennellia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus</italic> (syn. <italic>Coonemeria, Dactylomyces</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Petromyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hamigera</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Trichocomaceae</italic>:</bold></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Roumegueriella</italic><sup><xref ref-type="table-fn" rid="TFN4">1</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemicarpenteles</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dendrosphaera</italic> (tentatively, <italic>fide</italic> Malloch 1985b)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenoma</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Hemisartorya</italic><sup><xref ref-type="table-fn" rid="TFN6">3</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Rasamsonia</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerocleista</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Neosartorya</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sagenomella</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicilliopsis</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Talaromyces</italic> (syn. <italic>Sagenoma, Erythrogymnotheca</italic>)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Petromyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermomyces</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Warcupiella</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerocleista</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Trichocoma</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><bold><italic>Monascaceae</italic>:</bold></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thermoascus</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><bold>Unknown status:</bold></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ascorhiza</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Warcupiella</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ascorhiza</italic> (no strains available/studied)</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Leiothecium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Pseudocordyceps</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Monascus</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sarophorum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Xeromyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Dichleana</italic></td></tr></tbody></table><table-wrap-foot><fn id="TFN4"><label>1</label><p id="P34">Benny & Kimbrough (<xref ref-type="bibr" rid="R14">1980</xref>) accommodated <italic>Roumegueriella</italic> in the <italic>Trichocomaceae</italic>; however, Sung <italic>et al</italic>. (<xref ref-type="bibr" rid="R174">2007</xref>) showed that this genus belongs to the <italic>Bionectriaceae</italic> (<italic>Hypocreales</italic>) and is excluded in our study of the <italic>Trichocomaceae.</italic></p></fn><fn id="TFN5"><label>2</label><p id="P35">The type species <italic>Mallochia, M. echinulata</italic>, has a close relationship with <italic>Amaurascopsis reticulata</italic> and both species belong to the <italic>Onygenales</italic> (<xref ref-type="bibr" rid="R158">Solé <italic>et al</italic>. 2002</xref>).</p></fn><fn id="TFN6"><label>3</label><p id="P36">Comparison of the ITS sequence of the type strain of the type of <italic>Hemisartorya, H. maritima</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=186.77&link_type=cbs">CBS 186.77</ext-link>), showed to have a 100 % homology with the type of <italic>A. versicolor</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=583.65&link_type=cbs">CBS 583.65</ext-link> (J. Houbraken, unpubl. data).</p></fn></table-wrap-foot></table-wrap><p id="P37">The genera <italic>Chaetosartorya, Cristaspora, Dichlaena, Dichotomomyces, Eupenicillium, Edyuillia, Emericella, Eurotium, Hamigera, Hemicarpenteles, Hemisartorya, Neosartorya, Penicilliopsis, Petromyces, Sclerocleista, Thermoascus</italic> and <italic>Warcupiella</italic> were placed by Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>) in <italic>Aspergillaceae</italic> (as subfamily <italic>Dichlaenoideae</italic>). The majority of these genera are also included in our classification, and exceptions are <italic>Edyuilla</italic>, which is synonymised with <italic>Eurotium</italic> (<xref ref-type="bibr" rid="R7">von Arx 1974</xref>) and <italic>Thermoascus</italic>, which is classified in <italic>Thermoascaceae</italic>. The main difference is the placement of <italic>Monascaceae</italic> is <italic>Aspergillaceae</italic>. Benny & Kimbrough (<xref ref-type="bibr" rid="R14">1980</xref>) placed the genera <italic>Ascorhiza, Leiothecium, Monascus</italic> and <italic>Xeromyces</italic> in <italic>Monascaceae</italic> and suggested a relationship with <italic>Ascosphaerales.</italic> Later, several authors included this family in <italic>Pezizales</italic> (<xref ref-type="bibr" rid="R89">Malloch 1981</xref>, <xref ref-type="bibr" rid="R52">Hawksworth & Pitt 1983</xref>). Von Arx (1987), in his revision of <italic>Eurotiales</italic>, included <italic>Monascus</italic> in <italic>Onygenaceae</italic>, and reduced <italic>Monascaceae</italic> to synonymy. More recently, <italic>Monascaceae</italic> was placed in <italic>Eurotiales</italic> (<xref ref-type="bibr" rid="R84">LoBuglio <italic>et al</italic>. 1993</xref>, <xref ref-type="bibr" rid="R51">Hawksworth <italic>et al.</italic> 1995</xref>). Fennell (<xref ref-type="bibr" rid="R29">1973</xref>) noted that species of both <italic>Monascaceae</italic> and <italic>Eurotiaceae</italic>, which approximates our definition <italic>Aspergillaceae</italic>, form a distinct cleistothecial wall. Nevertheless, Fennell (<xref ref-type="bibr" rid="R29">1973</xref>) separated these families based on the formation of aleurioconidia by members of <italic>Monascaceae</italic>, but our results show that this feature is insufficient for family delimitation. Anamorph genera were not treated by Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>) and <xref ref-type="fig" rid="F1">Fig. 1</xref> shows that the genera <italic>Aspergillus, Basipetospora, Eladia, Fraseriella, Penicillium, Phialomyces, Phialosimplex, Polypaecilum, Thysanophora</italic> and <italic>Torulomyces</italic> are classified in <italic>Aspergillaceae</italic>. The teleomorph genera <italic>Chromocleista, Fennellia, Neocarpenteles</italic> and <italic>Neopetromyces,</italic> which were not treated in Malloch's study (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>), also belong to this family.</p><p id="P38">The genera <italic>Byssochlamys, Dendrosphaera, Sagenoma, Talaromyces</italic> and <italic>Trichocoma</italic> were placed by Malloch (<xref ref-type="bibr" rid="R90">1985a</xref>, <xref ref-type="bibr" rid="R91">b</xref>) in <italic>Trichocomaceae</italic> (as subfamily <italic>Trichomoideae</italic>), and anamorphs in <italic>Paecilomyces</italic> or <italic>Penicillium</italic> were linked to it. The results of our phylogenetic analysis (<xref ref-type="fig" rid="F1">Fig. 1</xref>) confirm the positioning of the genera <italic>Sagenoma, Talaromyces</italic> and <italic>Trichocoma</italic> in this family. In addition, the recently described genus <italic>Rasamsonia</italic> (<xref ref-type="bibr" rid="R60">Houbraken <italic>et al</italic>. 2011d</xref>), and the asexual genera <italic>Thermomyces</italic> and<italic>Sagenomella</italic> are classified in this family. Phylogenetic analysis shows that <italic>Byssochlamys</italic> is more closely related to <italic>Thermoascus</italic>. Fennell (<xref ref-type="bibr" rid="R29">1973</xref>) also observed the relationship between these two genera and stated that <italic>Byssochlamys</italic> is transitional between <italic>Thermoascaceae</italic> and <italic>Aspergillaceae</italic> (as <italic>Eurotiaceae</italic>). No strains of the genus <italic>Dendrosphaera</italic> were available and its position remains questionable. Kobayasi (<xref ref-type="bibr" rid="R68">1971</xref>) described an aleurioconidial state in <italic>Dendrosphaera eberhardtii</italic> and Benny & Kimbrough (<xref ref-type="bibr" rid="R14">1980</xref>) therefore suggested placing this species in <italic>Onygenales</italic> (which makes <italic>Dendrosphaeraceae</italic> a family of <italic>Onygenales</italic>). On the other hand, Malloch (<xref ref-type="bibr" rid="R91">1985b</xref>) noted that <italic>D. eberhardtii</italic> and <italic>T. paradoxa</italic>produce similar brushes of soft hyphae bearing asci and ascospores suggesting the placement in <italic>Trichocomaceae</italic>. Following Malloch (<xref ref-type="bibr" rid="R91">1985b</xref>), we tentatively place this genus in <italic>Trichocomaceae</italic>, and consequently, <italic>Dendrosphaeraceae</italic> are synonymised with <italic>Trichocomaceae</italic>.</p></sec><sec id="S15"><title>Phylogeny of Aspergillaceae</title><p id="P39">Seven clades (<xref ref-type="fig" rid="F1">Fig. 1</xref>, clades 1–7) can be distinguished in <italic>Aspergillaceae</italic>. Each clade is discussed and phenotypic characters of the members belonging to those clades are compared with those of <italic>Penicillium</italic>.</p><sec id="S16"><title>Clade 1: <italic>Penicillium sensu stricto</italic></title><p id="P40"><italic>Penicillium sensu lato</italic> is polyphyletic and species of this genus occur in the phylogenetically redefined families <italic>Aspergillaceae</italic> and <italic>Trichocomaceae</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>). The type species of <italic>Penicillium, Penicillium expansum</italic>, and the type species of <italic>Eupenicillium, E. crustaceum</italic>, form a clade within <italic>Aspergillaceae</italic>, defined here as <italic>Penicillium sensu stricto</italic>. The Penicillia not belonging to <italic>Penicillium</italic> <italic>s. str.</italic> are mainly classified in <italic>Trichocomaceae</italic>, in a clade together with the type species of <italic>Talaromyces, T. flavus</italic> (clade 10). The presence of two major clades in <italic>Penicillium</italic> is concordant with earlier studies using rDNA sequences (<xref ref-type="bibr" rid="R15">Berbee & Taylor 1995</xref>, <xref ref-type="bibr" rid="R106">Ogawa <italic>et al</italic>. 1997</xref>, <xref ref-type="bibr" rid="R172">Sugiyama 1998</xref>, <xref ref-type="bibr" rid="R107">Ogawa & Sugiyama 2000</xref>, <xref ref-type="bibr" rid="R176">Tamura <italic>et al.</italic> 2000</xref>). More recently, Wang & Zhuang (<xref ref-type="bibr" rid="R192">2007</xref>) used partial calmodulin sequences for the phylogenetic analysis of <italic>Penicillium</italic> and their data also supported the presence of two lineages in <italic>Trichocomaceae</italic>. However, their placement of <italic>Talaromyces trachyspermus</italic> on a single lineage is contradictory with our data. The <italic>Penicillium s. str.</italic> clade is most closely related to the <italic>Aspergillus</italic> clade (clade 2) and is phylogenetically more distant from genera with similar anamorphs such as <italic>Paecilomyces, Merimbla</italic> and the <italic>Penicillium</italic> species assigned to <italic>Trichocomaceae</italic> in this study. The phylogenetic study shows that various other genera belong to <italic>Penicillium s. str.</italic> The type species of the genera <italic>Chromocleista, Torulomyces, Thysanophora, Hemicarpenteles</italic> and <italic>Eladia</italic> are positioned in <italic>Penicillium s. str.</italic> These genera are considered here as synonyms of <italic>Penicillium,</italic> and the species are transferred as appropriate. Two well-supported subclades (<xref ref-type="fig" rid="F1">Fig. 1A, B</xref>) can be distinguished within <italic>Penicillium s. str.</italic> Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) classified <italic>Penicillium</italic> in four subgenera: <italic>Aspergilloides, Furcatum, Penicillium</italic> and <italic>Biverticillium</italic>. This system was mainly based on conidiophore branching and shape of the phialides. The type species of subgenus <italic>Penicillium</italic> (<italic>P. expansum</italic>) belongs to clade 1B and mainly comprises the species which are ter- and/or quarterverticillate. The type species of the subgenera <italic>Aspergilloides</italic> and <italic>Furcatum</italic> (<italic>P. aurantiobrunneum</italic> (= <italic>P. glabrum</italic>) and <italic>P. citrinum</italic>, respectively) are positioned in clade 1A, and monoverticillate and biverticillate species with flask shaped phialides more frequently occur in this clade. The type species of subgenus <italic>Biverticillium, Penicillium minioluteum</italic>, does not belong to <italic>Penicillium s. str.</italic> and is recombined as <italic>Talaromyces minioluteus</italic> elsewhere (<xref ref-type="bibr" rid="R149">Samson <italic>et al.</italic> 2011</xref>). Species with symmetrical biverticillate conidiophores and lanceolate phialdes belong to this clade. These observations confirm other studies that also showed that the current phenotype-based subgeneric classification, which is mainly based on the branching system of the <italic>Penicillium</italic> conidiophores, is incongruent with the molecular phylogeny (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, <xref ref-type="bibr" rid="R192">Wang & Zhuang 2007</xref>). It is proposed here to abandon the current subgeneric classification and to synonymise subgenus <italic>Furcatum</italic> with <italic>Aspergilloides</italic>, because the latter is an older name. The subgenera <italic>Aspergilloides</italic> and <italic>Penicillium</italic> correspond to clades 1A and 1B, respectively. The phylogenetic structure within these clades is examined with more depth in Part 3 of the discussion.</p></sec><sec id="S17"><title>Clade 2: <italic>Aspergillus</italic></title><p id="P41">A limited number of <italic>Aspergillus</italic> species and related teleomorphs are included in this study. The majority of the studied <italic>Aspergillus</italic> strains form a clade with 51 % bootstrap and 1.00 posterior probability support and this clade is defined here as <italic>Aspergillus sensu stricto. Aspergillus s. str.</italic> is phylogenetically closely related to <italic>Penicillium s. str.</italic> (77 % bs, 1.00 pp). These genera are morphologically distinct. <italic>Aspergillus</italic> forms nonseptate stipes, which often terminate in a distinct inflated part (vesicle) and have a foot-cell (<xref ref-type="bibr" rid="R133">Raper & Fennell 1965</xref>). Furthermore, the phialides are produced synchronously from the vesicle in <italic>Aspergillus</italic>. The distinction between these two genera is largely supported by the phylogeny. However, there are a few exceptions. <italic>Aspergillus paradoxus, A. crystallinus</italic> and <italic>A. malodoratus</italic> phylogenetically belong to <italic>Penicillium</italic> (R.A. Samson, unpubl. data). However, Raper & Fennell (<xref ref-type="bibr" rid="R133">1965</xref>) also noted that <italic>A. crystallinus</italic> and <italic>A. malodoratus</italic> produce triseriate structures that resemble <italic>Penicillium</italic>. In addition, there are also Aspergilli, which look similar to <italic>Penicillium</italic>. An example is <italic>Penicillium inflatum</italic>, which phylogenetically belongs to <italic>Aspergillus</italic> section <italic>Cremei</italic> and will be transferred from <italic>Penicillium</italic> to <italic>Aspergillus</italic> (R.A. Samson, unpubl. data). In addition, <italic>Aspergillus sydowii</italic> regularly produces small penicilli, and <italic>A. restrictus</italic> can produce diminutive vesiculate monoverticillate stipes, similar in appearance to those of some <italic>Penicillium</italic> species.</p><p id="P42">The classification of the genus <italic>Aspergillus</italic> is traditionally based on morphological characters. Raper & Fennell (<xref ref-type="bibr" rid="R133">1965</xref>) divided the genus into 18 groups. More recently, Peterson (<xref ref-type="bibr" rid="R115">2008</xref>) studied the relationship among Aspergilli using a multigene phylogeny and accepted 5 subgenera (<italic>Aspergillus, Circumdati, Fumigati, Nidulantes</italic> and <italic>Ornati</italic>) and 16 sections. Our data largely corresponds with Peterson's phylogeny, and four of the six subclades in <xref ref-type="fig" rid="F1">Fig. 1</xref> represent the <italic>Aspergillus</italic> subgenera as defined by Peterson (<xref ref-type="bibr" rid="R115">2008</xref>). However, there are some discrepancies. Sections <italic>Restricti</italic> and <italic>Aspergillus</italic> of the subgenus <italic>Aspergillus</italic> are on a well supported branch (100 % bs, 1.00 pp), confirming Peterson's data. Peterson (<xref ref-type="bibr" rid="R115">2008</xref>) placed sections <italic>Clavati</italic> and <italic>Fumigati</italic> in a single subgenus and, because of lack of statistical support, tentatively placed section <italic>Cervini</italic> in this subgenus. The representatives of section <italic>Cervini</italic> (<italic>Aspergillus cervinus, A. kanagawaensis</italic>) used in our study show that this section is basal to sections <italic>Fumigati</italic> and <italic>Clavati</italic> and belongs in the subgenus <italic>Fumigati</italic>. This confirms the phenotypic data of Gams <italic>et al.</italic> (<xref ref-type="bibr" rid="R44">1985</xref>), who placed sections <italic>Fumigati</italic> and <italic>Cervini</italic> in subgenus <italic>Fumigati</italic>. Phylogenetically, the monophyletic subgenus <italic>Circumdati</italic> as proposed by Peterson (<xref ref-type="bibr" rid="R115">2008</xref>) contains sections <italic>Circumdati, Candidi, Flavi, Flavipedes, Nigri, Terrei</italic> and <italic>Cremei</italic>. The relationship between the former six sections is poorly supported in our analysis (30 % bs, 0.94 pp) and more studies on the phylogenetic structure of <italic>Aspergillus</italic> are needed. In contrast to previous published results (<xref ref-type="bibr" rid="R111">Peterson 1995</xref>, <xref ref-type="bibr" rid="R115">2008</xref>), section <italic>Cremei</italic> appeared to be unrelated to the other sections of subgenus <italic>Circumdati</italic>. The studied members of section <italic>Cremei</italic> (<italic>A. pulvinus, A. wentii, A. brunneouniseriatus</italic>) formed a well supported clade with the type species of <italic>Cristaspora</italic> (<italic>C. arxii</italic>) and this clade is more closely related to members of the subgenus <italic>Aspergillus</italic> (64 % bs, 1.00 pp) than to subgenus <italic>Circumdati</italic>. The subgenus <italic>Nidulantes</italic> contains sections <italic>Nidulantes, Ochraceorosei, Usti, Sparsi</italic> and <italic>Aeni</italic> (<xref ref-type="bibr" rid="R40">Frisvad <italic>et al.</italic> 2005</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>, <xref ref-type="bibr" rid="R190">Varga <italic>et al.</italic> 2010</xref>). These results were confirmed in our study, with exception of section <italic>Aeni</italic>, because no representatives were included in our study. Section <italic>Ornati</italic> in subgenus <italic>Ornati</italic> is not positioned in <italic>Aspergillus s. str.</italic> and species belonging to this section are placed in the clade 7. Peterson (<xref ref-type="bibr" rid="R115">2008</xref>) suggested that it would be possible to change the classification of <italic>Aspergillus</italic> by splitting the genus based on teleomorphic states associated with particular monophyletic groups. However, he advocated keeping <italic>Aspergillus</italic> as a monophyletic genus, since this would reflect the actual relationships of species displaying an aspergillum whereas dividing the form genus into several genera based on teleomorphs would de-emphasise the relationships for most biologists not intimately familiar with the genus. Teleomorph genera associated with <italic>Aspergillus</italic> anamorphs include <italic>Chaetosartorya, Dichotomomyces, Emericella, Eurotium, Fennellia, Neocarpenteles, Neopetromyces, Neosartorya</italic> and <italic>Petromyces</italic>.</p><p id="P43">The type species of the genera <italic>Polypaecilum</italic> and <italic>Phialosimplex</italic> and the ex-type strain of <italic>Basipetospora halophilica</italic> form a strongly supported clade (100 % bs, 1.00 pp) within <italic>Aspergillus s. str.</italic> This clade is related to <italic>Aspergillus</italic> sections <italic>Cremei, Aspergillus</italic> and <italic>Restricti</italic> (64 % bs, 1.00 pp). Recently, <italic>Phialosimplex</italic> was introduced for species with simple phialides borne laterally on vegetative hyphae. These phialides form chains of conidia and are mostly monophialidic, but a second opening can also be formed (polyphialides). <italic>Sagenomella chlamydosporus</italic> and <italic>S. sclerotialis</italic> were transferred to this genus and <italic>Phialosimplex canicus</italic> was described as a new species (<xref ref-type="bibr" rid="R155">Sigler <italic>et al</italic>. 2010</xref>). The transfer of <italic>S. sclerotialis</italic> to <italic>Phialosimplex</italic> created a paraphyletic genus with <italic>Polypaecilum</italic> embedded in it. The type species of <italic>Polypaecilum, P. insolitum</italic>, produces its conidia on polyphialides and this feature is shared with members of <italic>Phialosimplex</italic> (<xref ref-type="bibr" rid="R156">Smith 1961a</xref>). The formation of chlamydospores and the occurrence in patient material are also shared features of both genera. This indicates that these genera could be congeneric and more research is needed to clarify their taxonomic status. <italic>Basipetospora halophilica</italic> also belongs to this diverse clade. The production of short solitary conidiophores or conidiogeneous cells by this species is a shared character with members of <italic>Phialosimplex, Polypaecilum</italic> and many other genera; however, formation of polyphialides by this species was not described (<xref ref-type="bibr" rid="R127">Pitt & Hocking 1985</xref>). Furthermore, <italic>Polypaecilum</italic> morphs related to <italic>Thermoascus</italic> and <italic>Dichotomyces</italic> are not part of this clade and this genus is polyphyletic.</p></sec><sec id="S18"><title>Clade 3: <italic>Hamigera</italic></title><p id="P44"><italic>Hamigera, Warcupiella</italic> and the related anamorphs <italic>Merimbla</italic> and <italic>Raperia</italic> are positioned in clade 3. The statistical support of this clade is low (< 70 % bs, < 0.90 pp) and the studied species might not be related. We decided to place the species <italic>Hamigera avellanea, Hamigera striata, Penicillium megasporum, Talaromyces leycettanus</italic> and <italic>Warcupiella spinosa</italic> in our taxon sampling based on data presented in previous studies, in which it was demonstrated that these species are related (<xref ref-type="bibr" rid="R107">Ogawa & Sugiyama 2000</xref>, <xref ref-type="bibr" rid="R176">Tamura <italic>et al.</italic> 2000</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>, <xref ref-type="bibr" rid="R119">Peterson <italic>et al.</italic> 2010</xref>). <italic>Penicillium giganteum, Merimbla ingelheimensis, Hamigera paravellanea, H. insecticola, H. inflata, H. terricola, H. pallida, H. fusca</italic> were not included in our study, but are also members of this clade (<xref ref-type="bibr" rid="R107">Ogawa & Sugiyama 2000</xref>, <xref ref-type="bibr" rid="R119">Peterson <italic>et al.</italic> 2010</xref>). <italic>Hamigera striata</italic> and <italic>Talaromyces leycettanus</italic> are on a strongly supported branch (94 % bs, 1.00 pp). Ogawa & Sugiyama (<xref ref-type="bibr" rid="R107">2000</xref>) showed in their 18S rDNA analysis that both species are related (83 % bs), confirming our data. Peterson <italic>et al.</italic> (<xref ref-type="bibr" rid="R119">2010</xref>) did not accept <italic>H. striata</italic> in <italic>Hamigera</italic> because of lack of statistical support and followed Benjamin's (<xref ref-type="bibr" rid="R13">1955</xref>) placement of this species in <italic>Talaromyces</italic>. Our results indicate that <italic>Talaromyces</italic> is phylogenetically distant and we therefore maintain <italic>H. striata</italic> in <italic>Hamigera. Talaromyces leycettanus</italic> also warrants further attention. Stolk & Samson (<xref ref-type="bibr" rid="R165">1972</xref>) noted that the anamorph of <italic>T. leycettanus, Paecilomyces leycettanus</italic>, seems to occupy an intermediate form between <italic>Penicillium</italic> and <italic>Paecilomyces</italic>. The complex conidiophore of <italic>T. leycettanus</italic> resembles <italic>Merimbla</italic> (= anamorph of <italic>Hamigera</italic>) (<xref ref-type="bibr" rid="R119">Peterson <italic>et al.</italic> 2010</xref>), supporting its placement in this diverse clade. <italic>Warcupiella</italic> is monotypic, represented by <italic>Warcupiella spinulosa</italic> (<xref ref-type="bibr" rid="R169">Subramanian 1972</xref>) and this species was originally described as <italic>Aspergillus spinulosus</italic> (<xref ref-type="bibr" rid="R133">Raper & Fennell 1965</xref>). Later, <italic>Raperia</italic> was introduced by Subramanian & Rajendran (<xref ref-type="bibr" rid="R170">1979</xref>) to accommodate the anamorph of <italic>W. spinulosa</italic> (<xref ref-type="bibr" rid="R8">von Arx 1986</xref>). Our results and others (<xref ref-type="bibr" rid="R176">Tamura <italic>et al.</italic> 2000</xref>, <xref ref-type="bibr" rid="R115">Peterson 2008</xref>) show that <italic>W. spinulosa</italic> does not belong to <italic>Penicillium</italic> or <italic>Aspergillus</italic>, and is more closely related to <italic>Hamigera avellanea</italic>. The relationship between <italic>Warcupiella</italic>/<italic>Raperia</italic> and <italic>Hamigera</italic> was also noted by von Arx (<xref ref-type="bibr" rid="R8">1986</xref>), and he transferred <italic>W. spinulosa</italic> to <italic>Hamigera. Penicillium megasporum</italic>, another member of this clade, has little affinity with <italic>Penicillium s. str.</italic> as noted by Pitt (<xref ref-type="bibr" rid="R126">1980</xref>), who created <italic>Penicillium</italic> series <italic>Megaspora</italic> for this species and <italic>P. asperosporum</italic>. Peterson <italic>et al.</italic> (<xref ref-type="bibr" rid="R119">2010</xref>) described the penicillus structure of <italic>P. megasporum</italic> as similar as that of <italic>Merimbla</italic>, but that phylogenetic analysis did not support inclusion of <italic>P. megasporum</italic> in the <italic>Hamigera</italic> clade. Our analysis lacks high bootstrap support to confidentially place <italic>P. megasporum, W. spinulosa</italic> and <italic>T. leycettanus</italic> in <italic>Hamigera</italic>. More research is needed to elucidate the classification of this diverse clade.</p></sec><sec id="S19"><title>Clade 4: <italic>Penicilliopsis</italic></title><p id="P45">Clade 4 comprises <italic>Aspergillus zonatus</italic> and <italic>Penicilliopsis clavariiformis</italic> and these two species form a strongly supported clade. <italic>Penicilliopsis</italic> is typified by <italic>P. clavariiformis</italic> and characterised by seed-borne, stipitate stromata. The anamorph genera <italic>Pseudocordyceps, Sarophorum</italic> and <italic>Stilbodendron</italic> are phenotypically related (<xref ref-type="bibr" rid="R146">Samson & Seifert 1985</xref>, <xref ref-type="bibr" rid="R61">Hsieh & Ju 2002</xref>). The former two genera have conidiogenous structures similar to those of <italic>Penicillium</italic> and the latter has <italic>Aspergillus</italic>-like conidiogenous structures. The sclerotia of <italic>Stilbothamnium</italic> morphologically resemble ascomata of <italic>Penicilliopsis</italic>. However, phylogenetically, the type species of <italic>Stilbothamnium, Aspergillus togoensis</italic>, belongs to <italic>Aspergillus</italic> subgenus <italic>Circumdati</italic> section <italic>Flavi</italic> and is unrelated to <italic>Penicilliopsis</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>). More research is needed to clarify the relationship between <italic>Penicillium, Penicilliopsis</italic> and the associated anamorph genera <italic>Pseudocordyceps</italic> and <italic>Sarophorum</italic>.</p></sec><sec id="S20"><title>Clade 5: <italic>Monascus, Xeromyces</italic> and <italic>Leiothecium</italic></title><p id="P46">The teleomorph genera <italic>Monascus, Xeromyces</italic> and <italic>Leiothecium</italic> belong in clade 5, as do the anamorph genera <italic>Fraseriella</italic> and <italic>Basipetospora</italic> (<xref ref-type="bibr" rid="R124">Pettersson <italic>et al.</italic> 2011</xref>, our data). Benny & Kimbrough (<xref ref-type="bibr" rid="R14">1980</xref>) placed <italic>Monascus, Xeromyces</italic> and <italic>Leiothecium</italic> in <italic>Monascaceae</italic> and this family is transferred here to <italic>Aspergillaceae</italic> (see part 1, phylogeny of <italic>Aspergillaceae</italic>). These genera have similar phenotypic characters including the formation of stalked ascomata and the production of aleurioconidia from undifferentiated conidiogenous cells. These features clearly set these genera apart from <italic>Penicillium s. str</italic>. and <italic>Aspergillus</italic>. Our results confirm those of Pettersson <italic>et al.</italic> (<xref ref-type="bibr" rid="R124">2011</xref>) and we follow their opinion in retaining <italic>Xeromyces</italic> for xerophilic <italic>Monascus</italic>-like species and <italic>Monascus</italic> for the species that grow at higher water activities. In addition, Pettersson <italic>et al.</italic> (<xref ref-type="bibr" rid="R124">2011</xref>) suggested that <italic>Chrysosporium inops</italic> should be transferred to a new genus. However, <xref ref-type="fig" rid="F1">Fig. 1</xref> shows that this species is closely related to <italic>X. bisporus</italic> and the xerophilic nature of boths species indicates a close relationship (<xref ref-type="bibr" rid="R128">Pitt & Hocking 2009</xref>, <xref ref-type="bibr" rid="R124">Pettersson <italic>et al.</italic> 2011</xref>). <italic>Leiothecium</italic> is basal to <italic>Monascus</italic> and the connection between these two genera was also noted by Samson & Mouchacca (<xref ref-type="bibr" rid="R143">1975</xref>). <italic>Aspergillus clavatoflavus</italic> is basal to this clade, but the relationship lacks statistical support. The micromorphology is <italic>A. clavatoflavus</italic> differs from the members of clade 5 and therefore this species is placed outside this clade, awaiting more conclusive data.</p></sec><sec id="S21"><title>Clade 6: <italic>Phialomyces</italic></title><p id="P47">The type species of <italic>Phialomyces, Phial. macrosporus</italic> (<xref ref-type="bibr" rid="R99">Misra & Talbot 1968</xref>), is positioned in clade 6 and is closely related to <italic>Penicillium arenicola</italic> (100 % bs, 1.00 pp). <italic>Merimbla humicoloides</italic> (= <italic>Penicillium humicoloides sensu</italic> <xref ref-type="bibr" rid="R119">Peterson <italic>et al</italic>. 2010</xref>) also belongs to this clade (R.A. Samson, unpubl. data). All three species are phylogenetically distinct form <italic>Penicillium s. str.</italic> Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) placed <italic>P. arenicola</italic> in a separate section and series and noted that this species may not be a true <italic>Penicillium</italic>. Phenotypically, <italic>Phial. macrosporus, M. humicoloides</italic> and <italic>P. arenicola</italic> form conidia in shades of gold-brown, a feature uncommon for <italic>Penicillium</italic> species. These species can produce terverticillate conidiophores, a character also present in subgenus <italic>Penicillium</italic> (clade 1B). Our results indicate that <italic>P. arenicola</italic> and <italic>M. humicoloides</italic> should be transferred to another genus.</p></sec><sec id="S22"><title>Clade 7: <italic>Sclerocleista</italic></title><p id="P48"><italic>Sclerocleista ornata</italic> and <italic>S. thaxteri</italic> are basal to <italic>Phial. macrosporus</italic> and <italic>P. arenicola</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>). <italic>Sclerocleista ornata</italic> was originally described as <italic>Aspergillus ornatus</italic> (<xref ref-type="bibr" rid="R134">Raper <italic>et al.</italic> 1953</xref>), and later transferred to <italic>Sclerocleista</italic> (<xref ref-type="bibr" rid="R169">Subramanian 1972</xref>). <italic>Sclerocleista thaxteri</italic> was originally described in <italic>Sclerocleista</italic> and later von Arx (<xref ref-type="bibr" rid="R7">1974</xref>) transferred this species to <italic>Hemicarpenteles</italic>. The two species are closely related, and phylogenetically distant from <italic>H. paradoxus,</italic> the type species of <italic>Hemicarpenteles</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>, <italic>Penicillium s. str</italic>.). Peterson (<xref ref-type="bibr" rid="R115">2008</xref>) placed <italic>Sclerocleista</italic> basal to the Aspergilli, suggesting a monophyletic <italic>Aspergillus</italic> clade; however, our data do not support this conclusion. <italic>Sclerocleista</italic> differs from <italic>Penicillium s. str.</italic> in having an <italic>Aspergillus</italic>-type anamorph and purple coloured cleistothecia filled with lenticular ascospores (<xref ref-type="bibr" rid="R133">Raper & Fennell 1965</xref>).</p></sec></sec><sec id="S23"><title>Phylogeny of Thermoascaceae</title><p id="P49"><xref ref-type="fig" rid="F1">Figure 1</xref> shows that two clades (clade 8 and 9) are present in <italic>Thermoascaceae</italic> (= lineage 2). The phylogeny of these two clades and the comparison of the species belonging to these two clades with <italic>Penicillium s. str</italic>. is discussed below.</p><sec id="S24"><title>Clade 8: <italic>Thermoascus</italic></title><p id="P50"><italic>Thermoascus aurantiacus, T. crustaceus</italic> and <italic>T. thermophilus</italic> are together with <italic>Byssochlamys verrucosa</italic> in a separate clade. The taxonomy of <italic>Thermoascus</italic> is treated in various studies. Apinis (<xref ref-type="bibr" rid="R5">1967</xref>) split <italic>Thermoascus</italic> in two: <italic>Thermoascus</italic> was retained for its type species <italic>T. aurantiacus</italic>, and <italic>T. thermophilus</italic> and <italic>T. crustaceus</italic> were transferred to <italic>Dactylomyces</italic>. Later, Mouchacca (1997) divided <italic>Dactylomyces</italic> further in two, creating <italic>Coonemeria</italic> for <italic>T. crustaceus</italic>. Although these species have different anamorphs (<italic>Paecilomyces</italic>/<italic>Polypaecilum</italic>), our phylogenetic study (<xref ref-type="fig" rid="F1">Fig. 1</xref>) shows that these three species are closely related and should be retained in <italic>Thermoascus</italic>. Samson <italic>et al.</italic> (<xref ref-type="bibr" rid="R145">2009</xref>) noted that <italic>Byssochlamys verrucosa</italic> is misidentified in <italic>Byssochlamys</italic> but related to <italic>Thermoascus</italic>, and this observation is confirmed here. <italic>Thermoascus</italic> has a similar type of sclerotioid cleistothecium as members of <italic>Penicillium s. str.</italic> (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>). These two genera differ mainly in ascomatal development. Ascomata of <italic>Thermoascus</italic> are initiated by an ascogonial coil (<xref ref-type="bibr" rid="R163">Stolk 1965</xref>, <xref ref-type="bibr" rid="R171">Subramanian & Rajendran 1980</xref>), whereas in <italic>Penicillium s. str</italic>. the formation begins with sclerotium-like bodies inside which the ascogonia develop. Furthermore, the anamorphs of <italic>Thermoascus</italic> are not of the <italic>Penicillium</italic> type, but can be similar to <italic>Paecilomyces</italic>.</p></sec><sec id="S25"><title>Clade 9: <italic>Paecilomyces</italic></title><p id="P51">The types of <italic>Paecilomyces</italic> (<italic>P. variotii</italic>) and <italic>Byssochlamys</italic> (<italic>B. nivea</italic>) occur together on a branch with 100
% bootstrap support. Using a polyphasic approach, Samson <italic>et al.</italic> (<xref ref-type="bibr" rid="R145">2009</xref>) showed that the genera <italic>Byssochlamys</italic> and <italic>Paecilomyces s. str.</italic> are closely related and form a monophyletic group. <italic>Paecilomyces</italic> was introduced by Bainier (<xref ref-type="bibr" rid="R10">1907</xref>) and has priority over <italic>Byssochlamys</italic> (<xref ref-type="bibr" rid="R196">Westling 1909</xref>). Phylogenetic analysis of the 18S rDNA demonstrated that <italic>Paecilomyces sensu</italic> Samson (<xref ref-type="bibr" rid="R142">1974</xref>) is polyphyletic across two subclasses, <italic>Sordariomycetidae</italic> and <italic>Eurotiomycetidae</italic>. The type species of this genus, <italic>Paecilomyces variotii</italic>, and its thermophilic relatives belong in the <italic>Eurotiales</italic> (<xref ref-type="bibr" rid="R82">Luangsa-ard <italic>et al.</italic> 2004</xref>). <xref ref-type="fig" rid="F1">Figure 1</xref> shows that <italic>Paecilomyces s. str.</italic> is also phylogenetically distinct from <italic>Penicillium</italic>. Morphological characters also support this conclusion. The conidia of <italic>Paecilomyces s. str.</italic> are olive-brown and formed in phialides that have a broad base and end in a long and slender neck, while the conidia of <italic>Penicillium</italic> species are green and formed in flask or cylindrical shaped phialides. In addition, the conidiophores of <italic>Paecilomyces s. str.</italic> are more irregularly branched than those of <italic>Penicillium</italic>. The teleomorphs are also different: those of <italic>Paecilomyces</italic> (formerly known as <italic>Byssochlamys</italic>) are almost naked while <italic>Penicillium s. str</italic>. produces cleistothecia with a distinct wall.</p></sec></sec><sec id="S26"><title>Phylogeny of Trichocomaceae</title><p id="P52">Five clades (clades 10–15) can be recognised in the more narrowly delimited <italic>Trichocomaceae</italic>. The species treated in these clades are phylogenetically distinct from <italic>Penicillium s. str</italic>., but some are phenotypically similar.</p><sec id="S27"><title>Clade 10: <italic>Talaromyces</italic></title><p id="P53">The majority of <italic>Penicillium</italic> species assigned to the subgenus <italic>Biverticillium</italic> belong in clade 10 (incl. type of subgenus <italic>Biverticillium, P. minioluteum</italic>) together with the type species of the genera <italic>Talaromyces</italic> and <italic>Sagenoma</italic>. These species are phylogenetically distant from <italic>Penicillium s. str.</italic> and therefore these species are transferred to the genus <italic>Talaromyces</italic> (<xref ref-type="bibr" rid="R149">Samson <italic>et al.</italic> 2011</xref>, this study). Phenotypically, <italic>Talaromyces</italic> differs from <italic>Penicillium s. str.</italic> by the formation of symmetrically branched conidiophores with lanceolate phialides, and the production of soft ascomata without a well-defined, persistant wall. Members of the <italic>Talaromyces</italic> clade grow slower on the agar medium G25N than <italic>Penicillium s. str</italic>. members (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>). Also differences in ubiquinones and extrolites patterns are observed between <italic>Penicillium sensu stricto</italic> and <italic>Talaromyces</italic>. The Q9 ubiquinone system was present in most <italic>Penicillium sensu stricto</italic> species, while nearly all <italic>Talaromyces</italic> have Q10(H<sub>2</sub>) (<xref ref-type="bibr" rid="R109">Paterson 1998</xref>). In addition, extrolites such as mitorubrins, certain bisanthraquinones (rugulosin, skyrin), duclauxin and glauconic acide were detected in <italic>Talaromyces</italic>, but never found in <italic>Penicillium sensu stricto</italic> (<xref ref-type="bibr" rid="R32">Frisvad <italic>et al</italic>. 1998</xref>). The taxonomic and phylogenetic structure of <italic>Talaromyces</italic> is considered further by Samson <italic>et al</italic>. (<xref ref-type="bibr" rid="R149">2011</xref>).</p><p id="P54">The neotype strain of <italic>Aphanoascus cinnabarinus sensu</italic> Udagawa and Takada also belongs to this clade. Much taxonomic confusion followed after the proposal of <italic>Aphanoascus</italic> by Zukal (<xref ref-type="bibr" rid="R203">1890</xref>). Most authors follow Apinis (<xref ref-type="bibr" rid="R6">1968</xref>) and maintain <italic>Aphanoascus</italic> that is typified by <italic>A. fulvescens</italic>. In addition, the neotypification of <italic>A. cinnabarinus</italic> by Udagawa & Takada (<xref ref-type="bibr" rid="R184">1973</xref>) was incorrect, because their neotype strain had a <italic>Paecilomyces</italic> anamorph, while Zukal's original description and illustrations showed structures of a <italic>Chrysosporium</italic> anamorph (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>). Based on morphological characters, Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>) suggested that <italic>Chromocleista cinnabarina</italic> (as <italic>A. cinnabarinus sensu</italic> Udagawa & Takada) belongs to <italic>Eurotiales,</italic> and that this species occupies an intermediate position between the genera <italic>Thermoascus</italic> and <italic>Talaromyces</italic>. The result of our multigene phylogeny shows that <italic>C. cinnabarina</italic> belongs to <italic>Talaromyces s. str.</italic></p><p id="P55">This data is in concordance with the 18S rDNA sequence data of Ogawa & Sugiyama (<xref ref-type="bibr" rid="R107">2000</xref>), which shows that <italic>C. cinnabarina</italic> forms a monophyletic group with <italic>T. macrosporus</italic> and <italic>T. bacillisporus</italic>. No specimens of <italic>Erythrogymnotheca</italic> were studied, but an ITS sequence of the type species of this genus (<italic>E. paucispora</italic>) is deposited GenBank (AB176603) and a BLAST search on GenBank and internal CBS databases shows that this sequence belongs to <italic>Talaromyces s. str</italic>.</p></sec><sec id="S28"><title>Clade 11: <italic>Thermomyces</italic></title><p id="P56"><italic>Talaromyces thermophilus</italic> belongs to the same clade as the type of <italic>Thermomyces, T. lanuginosus. Talaromyces thermophilus</italic> and <italic>Therm. lanuginosus</italic> share similar characters, including their ability to grow at high temperatures and the formation of thick-walled chlamydospores or chlamydospore-like conidia. These characters are not shared by members of <italic>Penicillium s. str. Talaromyces luteus</italic> is basal to this clade. This species is not thermophilic and phenotypically different from <italic>Thermomyces</italic> and <italic>Tal. thermophilus</italic>, and it is therefore excluded from clade 11.</p></sec><sec id="S29"><title>Clade 12: <italic>Sagenomella</italic></title><p id="P57">Clade 12 is centered around the type species of <italic>Sagenomella, S. diversispora</italic>, and this genus is phylogenetically unrelated to <italic>Penicillium s. str. Sagenomella</italic> was described by Gams (<xref ref-type="bibr" rid="R43">1978</xref>) for <italic>Acremonium</italic>-like fungi and is characterised by connected conidial chains and sympodially proliferating, often centrally swollen phialides. These characters are not present in <italic>Penicillium s. str.</italic> Molecular data showed that <italic>Sagenomella sensu</italic> Gams is polyphyletic (<xref ref-type="bibr" rid="R27">Endo <italic>et al.</italic> 1998</xref>, <xref ref-type="bibr" rid="R177">Thanh <italic>et al.</italic> 1998</xref>, our results). Sigler <italic>et al.</italic> (<xref ref-type="bibr" rid="R155">2010</xref>) transferred <italic>S. chlamydospora</italic> and <italic>S. sclerotialis</italic> to the new genus <italic>Phialosimplex</italic> and <italic>Sagenomella bohemica</italic> belongs in <italic>Talaromyces</italic> (<xref ref-type="bibr" rid="R149">Samson <italic>et al.</italic> 2011</xref>). The close relationship of this genus with <italic>Talaromyces</italic> indicates that <italic>Sagenomella</italic> is a reduced form of <italic>Talaromyces</italic>.</p></sec><sec id="S30"><title>Clade 13: <italic>Rasamsonia</italic></title><p id="P58">The thermophiles <italic>Talaromyces emersonii</italic> and <italic>T. byssochlamydoides</italic> were transferred to <italic>Rasamsonia</italic> (<xref ref-type="bibr" rid="R60">Houbraken <italic>et al.</italic> 2011d</xref>), leaving <italic>T. thermophilus</italic> as sole thermophile in <italic>Talaromyces</italic>. However, our phylogenetic analysis shows that this species belongs to <italic>Thermomyces</italic> and not to <italic>Talaromyces</italic>. The genus <italic>Rasamsonia</italic> was erected for thermotolerant or thermophilic species, which have cylindrical phialides usually gradually tapering towards the apices, conidiophores with distinctly rough walled stipes, olive-brown conidia and ascomata, if present, with a scanty covering. This clade contains the species <italic>R. argillacea, R. brevistipitata, R. byssochlamydoides, R. cylindrospora, R. eburnea</italic> and <italic>R. emersonii</italic> (<xref ref-type="bibr" rid="R60">Houbraken <italic>et al.</italic> 2011d</xref>).</p></sec><sec id="S31"><title>Clade 14: <italic>Trichocoma</italic></title><p id="P59">The monotypic genus <italic>Trichocoma</italic> is typified by <italic>Trichocoma paradoxa</italic> and is characterised by asci born in hyphal masses or tufts that can be up to 10–20 mm long (<xref ref-type="bibr" rid="R72">Kominami <italic>et al</italic>. 1952</xref>, <xref ref-type="bibr" rid="R91">Malloch 1985b</xref>). The anamorph of this species resembles an anamorph of <italic>Talaromyces</italic>. However, <italic>Trichocoma</italic> produces conidia in shades of brown. <italic>Rasamsonia</italic> is phylogenetically related to <italic>Trichocoma</italic>, and can be differentiated by the presence of scanty ascomatal coverings and its ability to grow at temperatures above 40 °C.</p></sec></sec><sec id="S32"><title>Excluded genera: Geosmithia, Phialotubus and Yunnania</title><p id="P60">The genera <italic>Geosmithia, Phialotubus</italic> and <italic>Yunnania</italic> have sometimes been hypothesised to be related to <italic>Penicillium</italic> (<xref ref-type="bibr" rid="R43">Gams 1978</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R73">Kong 1998</xref>). Our data shows that these genera do not belong to the <italic>Eurotiales</italic> and details are provided below.</p><sec id="S33"><title>Geosmithia</title><p id="P61">The genus <italic>Geosmithia</italic> is typified by <italic>G. lavendula</italic> (<xref ref-type="bibr" rid="R125">Pitt 1978</xref>) and is a polyphyletic morphogenus introduced to classify <italic>Penicillium</italic> species, which are characterised by: a) cylindroidal phialides and conidia, b) rugulose to rugose conidiophores walls, metulae and phialides and c) conidial colour other than green (with the exception of <italic>G. namyslowskii</italic>). Anamorphs of <italic>Geosmithia</italic> have affinities with hypocrealean (<italic>Hypocreales</italic>: <italic>Bionectriaceae</italic>) and eurotialean (<italic>Eurotiales</italic>: <italic>Trichocomaceae</italic>) fungi, and the type species of <italic>Geosmithia, G. lavendula</italic>, is related to <italic>Acremonium alternatum</italic>, the type species of <italic>Acremonium</italic> (<xref ref-type="bibr" rid="R106">Ogawa <italic>et al.</italic> 1997</xref>, <xref ref-type="bibr" rid="R138">Rossman <italic>et al</italic>. 2001</xref>, <xref ref-type="bibr" rid="R173">Summerbell <italic>et al.</italic> 2011</xref>). Currently, there are 16 described species (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R199">Yaguchi <italic>et al.</italic> 1993</xref>, <xref ref-type="bibr" rid="R200">1994</xref>, <xref ref-type="bibr" rid="R130">Pitt <italic>et al.</italic> 2000</xref>, <xref ref-type="bibr" rid="R71">Kolařík <italic>et al.</italic> 2004</xref>, <xref ref-type="bibr" rid="R70">2005</xref>, <xref ref-type="bibr" rid="R69">2010</xref>), and eight of these species (<italic>G. fassatiae, G. flava, G. langdonii, G. lavendula, G. morbida, G. obscura, G. pallida</italic>, and <italic>G. putterillii</italic>) belong to the <italic>Hypocreales. Geosmithia argillacea</italic> (teleomorph <italic>Talaromyces eburneus sensu</italic> <xref ref-type="bibr" rid="R201">Yaguchi <italic>et al</italic>. 2005</xref>), <italic>G. eburnea</italic> (teleomorph <italic>Talaromyces eburneus sensu</italic> <xref ref-type="bibr" rid="R200">Yaguchi <italic>et al</italic>. 1994</xref>), <italic>G. emersonii</italic> (teleomorph <italic>Talaromyces emersonii</italic>) and <italic>G. cylindrospora</italic> are closely related to each other and were recently transferred to <italic>Rasamsonia</italic> (<xref ref-type="bibr" rid="R60">Houbraken <italic>et al.</italic> 2011d</xref>, see clade 13 above). <italic>Geosmithia swiftii</italic> (teleomorph <italic>Talaromyces bacillisporus</italic>) and <italic>G. viridis</italic> belong to <italic>Talaromyces s. str.</italic> and <italic>G. namyslowskii</italic> and <italic>G. malachiteum</italic> (described as the anamorph of <italic>Chromocleista malachitea</italic>) belong to <italic>Penicillium s. str.</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>). Zaleski (<xref ref-type="bibr" rid="R202">1927</xref>) originally described <italic>Geosmithia namyslowskii</italic> as <italic>Penicillium namyslowskii</italic> and the new combination of <italic>Penicillium malachiteum</italic> is made elsewhere in this article.</p></sec><sec id="S34"><title>Phialotubus</title><p id="P62"><italic>Phialotubus</italic> (<xref ref-type="bibr" rid="R140">Roy & Leelavathy 1966</xref>) is monotypic with <italic>Phialotubus microsporus</italic> as the type. This species is characterised by the formation of cylindrical phialides with long hyaline thread-like projections, which get prolonged into the hyaline tube-like projection when conidia are formed (<xref ref-type="fig" rid="F2">Fig. 2</xref>). The conidia are fusiform in shape and produced in chains (<xref ref-type="bibr" rid="R140">Roy & Leelavathy 1966</xref>, <xref ref-type="bibr" rid="R43">Gams 1978</xref>, Arx 1981). These characters suggest a close connection with the <italic>Eurotiales</italic>, for example with <italic>Paecilomyces, Phialomyces,Sagenomella</italic> and <italic>Torulomyces</italic>. However, a BLAST search on GenBank with an ITS sequence of strain <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=861.70&link_type=cbs">CBS 861.70</ext-link><sup>isoT</sup> (GenBank no. JN831360) did not retrieve any high similarity matches with members of the <italic>Eurotiales</italic>. The overall similarity matches were low and this species probably belongs to the class <italic>Sordariomycetes</italic>.</p><fig id="F2" position="float"><label>Fig. 2.</label><caption><p><italic>Phialotubus microsporus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=861.70&link_type=cbs">CBS 861.70</ext-link><sup>isoT</sup>. A. Colonies grown for 7 d at 25 °C, from left to right: CYA, MEA, OA. B–D. Conidiophores and conidia. Scale bar = 10 μm.</p></caption><graphic xlink:href="1fig2"></graphic></fig></sec><sec id="S35"><title>Yunnania</title><p id="P63">Kong (<xref ref-type="bibr" rid="R73">1998</xref>) proposed the genus <italic>Yunnania</italic> and typified it with <italic>Y. penicillata</italic>. The truncated conidia and the black or brownish black colonies resemble those of <italic>Scopulariopsis</italic>. In addition, the conidia are produced by annelides (<xref ref-type="fig" rid="F3">Fig. 3</xref>). Examination of the type strain of <italic>Y. penicillata</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130296&link_type=cbs">CBS 130296</ext-link><sup>T</sup>) showed that this species is morphologically related to <italic>Scedosporium</italic>. A BLAST search on GenBank with an ITS sequence of this species (GenBank no. JN831361) did not retrieve a high similarity match, but showed that this species belongs to the order <italic>Microascales</italic>.</p><fig id="F3" position="float"><label>Fig. 3.</label><caption><p><italic>Yunnania penicillata</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130296&link_type=cbs">CBS 130296</ext-link><sup>T</sup>. A. Colonies grown for 7 d at 25 °C, from left to right: MEA, OA, CYA. B–D. Conidiophores and conidia.</p></caption><graphic xlink:href="1fig3"></graphic></fig></sec></sec><sec id="S36"><title>Taxonomic implications</title><p id="P64"><bold><italic>Aspergillaceae</italic></bold> Link, Abh. dt. Akad. Wiss. Berlin 1824: 165. 1826.</p><p id="P65"><list list-type="simple"><list-item><p id="P66">= <italic>Eurotiaceae</italic> Clements and Shear, Gen. Fung. 50. 1931.</p></list-item><list-item><p id="P67">= <italic>Monascaceae</italic> J. Schröter, Nat. Pflanzenfamilien 1: 148. 1894.</p></list-item><list-item><p id="P68">= <italic>Hemicarpenteleaceae</italic> Locquin, Tribune Méd. (Paris) 1. 1972. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item><list-item><p id="P69">= <italic>Penicilliaceae</italic> Vuillemin, Pl. Jungh. 10: 172. 1910. (as Penicilliacées <italic>nom</italic>. <italic>inval</italic>. Art. 32.1b).</p></list-item><list-item><p id="P70">= <italic>Penicilliopsidaceae</italic> Locquin, Tribune Méd. (Paris) 1. 1972. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item><list-item><p id="P71">= <italic>Phialomycetaeae</italic> Locquin, Mycologie générale et structurale: 212. 1984. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item><list-item><p id="P72">= <italic>Warcupiellaceae</italic> Locquin, Mycologie générale et structurale: 167. 1984. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item><list-item><p id="P73">= <italic>Xeromycetaceae</italic> Locquin, Tribune Méd. (Paris) 1. 1972. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item></list></p><p id="P74">Type: <italic>Aspergillus</italic> Fr: Fr.</p><p id="P75"><bold><italic>Thermoascaceae</italic></bold> Apinis, Trans. Br. Mycol. Soc 50: 581. 1967.</p><p id="P76">Type: <italic>Thermoascus</italic> Miehe</p><p id="P77"><bold><italic>Trichocomaceae</italic></bold> E. Fischer, Nat. Pflanzenfam. 1: 310. 1897. (as <italic>Trichocomataceae</italic>)</p><p id="P78"><list list-type="simple"><list-item><p id="P79">= <italic>Talaromycetaceae</italic> Locquin, Mycologie générale et structurale: 176. 1984. <italic>nom</italic>. <italic>inval</italic>. (Art. 36).</p></list-item><list-item><p id="P80">= <italic>Dendrosphaeraceae</italic> Ciferri ex Benny & Kimbrough, Mycotaxon 12: 22. 1980.</p></list-item></list></p><p id="P81">Type: <italic>Trichocoma</italic> Junghuhn</p></sec></sec><sec id="S37"><title>Part Two: delimitation of <italic>Penicillium</italic></title><sec id="S38"><title>Authority</title><p id="P82">The generic name <italic>Penicillium</italic> is attributed to Link (<xref ref-type="bibr" rid="R80">1809</xref>). Link included three species within <italic>Penicillium, P. glaucum, P. candidum</italic> and <italic>P. expansum</italic>. He illustrated <italic>P. candidum</italic>, which clearly shows structures of a <italic>Penicillium</italic> species. Later, <italic>Penicillium expansum</italic> was selected by Thom (1910) and later (co-)authors as the lectotype of <italic>Penicillium</italic>. The generic name <italic>Penicillium</italic> was attributed by Fries (<xref ref-type="bibr" rid="R31">1832</xref>: 406) to Link (<xref ref-type="bibr" rid="R80">1809</xref>). Hawksworth <italic>et al.</italic> (<xref ref-type="bibr" rid="R49">1976</xref>) proposed to conserve the generic name <italic>Penicillium</italic> as <italic>Penicillium</italic> Link ex Grey over <italic>Penicillium</italic> <xref ref-type="bibr" rid="R31">Fries 1832</xref> (proposal no. 420), and lectotypified the genus with <italic>Penicillium expansum</italic> Link ex Grey. This proposal was countered by Jørgensen & Gunnerbeck (<xref ref-type="bibr" rid="R63">1977</xref>) because Fries listed “<italic>Mucor crustaceus</italic> L.” as a typical species of <italic>Penicillium</italic> and not as the type species of this genus. The proposal of Hawksworth <italic>et al</italic>. (<xref ref-type="bibr" rid="R49">1976</xref>) was therefore rejected (<xref ref-type="bibr" rid="R110">Petersen 1980</xref>). The general starting point for fungal names is Linnaeus 1753, but there are a few exceptions and these are mentioned in the ICBN under art. 13e. One exception is that names used by E.M. Fries' “Systema mycologicum” 1821–1832 have a protected status. These names are sanctioned and have priority over older synonyms and homonyms. The authority used here is therefore <italic>Penicillium</italic> Link: Fries.</p></sec><sec id="S39"><title>Generic diagnosis</title><p id="P83">The concept of <italic>Penicillium</italic> has been refined and restated often in mycological history. The concept of Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>) is followed here; however, there are some emendations. In our concept, <italic>Penicillium</italic> includes species with pigmented stipes (<italic>Thysanophora</italic> species, <italic>P. stolkiae</italic> and related species), as well as species formerly ascribed to the genera <italic>Eladia, Torulomyces, Chromocleista</italic> and <italic>Hemicarpenteles</italic>. Details regarding the position of these genera in <italic>Penicillium</italic> are presented below. Another important difference between our and Raper & Thom's (<xref ref-type="bibr" rid="R135">1949</xref>) concept is the exclusion of <italic>Talaromyces</italic> and related <italic>Penicillium</italic> species. In our concept, only teleomorphs producing pseudoparenchymatous and sclerotioid ascomata are included (“Eupenicillium-type”), and <italic>Talaromyces</italic> species, with soft ascomata without a well-defined, persistent wall, are excluded (<xref ref-type="bibr" rid="R149">Samson <italic>et al</italic>. 2011</xref>). Also the <italic>Penicillium</italic> species, which have lanceolate phialides and metulae with equal lengths as the phialides, are excluded. These species are also phylogenetically distinct (<xref ref-type="fig" rid="F1">Fig. 1</xref>). Our emended generic diagnosis is derived from Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>) and is presented here:</p><p id="P84"><bold><italic>Penicillium</italic></bold> Link: Fries, Systema Mycologicum 3: 406. 1832.</p><p id="P85"><italic>Vegetative mycelium</italic> abundant, entirely submerged or more or less effused, irregularly branching, septate, hyaline or brightly coloured and forming a dense and compact mycelia colony with well-defined margins. <italic>Conidiophores</italic> borne from undifferentiated subsurface, superficial or aerial hyphae, rarely subapically proliferation under terminal penicillus. <italic>Stipes</italic> relatively narrow and thin walled, 2–5 μm, and in some species apically swollen, hyaline, in some species brown. <italic>Conidial apparatus</italic> usually a well defined structure (brush or broom), named the Penicillus; penicilli comprised of phialides born directly on the stipe, or with one, two or rarely more verticils of metulae and rami as supporting cells. <italic>Conidiogenous cells</italic> phialides, borne in succession, <italic>i.e.</italic> not synchronouse, rarely exceeding 15 μm in length, ampulliform, rarely cylindrical. <italic>Conidia</italic> in unbranched chains, borne basipetally, single celled, commonly between 2–5 μm in diameter, rarely exceeding 6 μm, <italic>en masse</italic> coloured in shades of green, rarely white, olive or brown. <italic>Chlamydospores</italic> absent. <italic>Sclerotia</italic> occasionally produced, composed of thick-walled cells, usually hard. <italic>Cleistothecia</italic>, if produced, usually hard, globose to subglobse, pseudoparenchymatous or sclerochymatous, ripening from the center outward and often tardily; white, pale, yellow, orange or brown coloured, occasionally black or red. <italic>Asci</italic> ellipsoidal to globose, usually 8-spored, 5–15 μm. <italic>Ascospores</italic> lenticular, usually with equatorial ridges, 2–5 μm.</p></sec><sec id="S40"><title>Synonyms of Penicillium</title><p id="P86">The re-definition of the genus <italic>Penicillium</italic> has several taxonomic implications. Based on the phylogenetic data presented in <xref ref-type="fig" rid="F1">Fig. 1</xref> in combination with a review of literature, we place the genera <italic>Chromocleista, Carpenteles, Citromyces, Eladia, Eupenicillium, Hemicarpenteles, Thysanophora</italic> and <italic>Torulomyces</italic> in synonymy with <italic>Penicillium</italic>. More genera are congeneric with <italic>Penicillium</italic> and a more extended list can be found in Seifert <italic>et al</italic>. (<xref ref-type="bibr" rid="R152">2011</xref>: 333). Each genus is discussed here and new combinations are proposed below for the species accommodated in these genera.</p><p id="P87"><bold><italic>Penicillium</italic></bold> Link: Fries, Systema Mycologicum 3: 406. 1832.</p><p id="P88"><list list-type="simple"><list-item><p id="P89">= <italic>Penicillium</italic> Link, Obs. Mycol 1: 16. 1809 (<italic>nom. inval.</italic>, Art. 13e).</p></list-item><list-item><p id="P90">= <italic>Coremium</italic> Link ex Gray, Nat. Arr. Br. Pl. 1: 563. 1821.</p></list-item><list-item><p id="P91">= <italic>Eupenicillium</italic> Ludwig, Lehrb. Nied. Kryptog.: 263. 1892.</p></list-item><list-item><p id="P92">= <italic>Citromyces</italic> Wehmer, Bleitr. Kennt. Pilze 1: 1. 1893.</p></list-item><list-item><p id="P93">= <italic>Carpenteles</italic> Langeron, C.r. Séanc. Soc. Boil. Paris 87: 344. 1922.</p></list-item><list-item><p id="P94">= <italic>Torulomyces</italic> Delitsch, Systematik der Schimmelpilze: 91. 1943.</p></list-item><list-item><p id="P95">= <italic>Thysanophora</italic> Kendrick, Can. J. Bot. 39: 820. 1961.</p></list-item><list-item><p id="P96">= <italic>Eladia</italic> Smith, Trans. Brit. Mycol. Soc. 44: 47. 1961.</p></list-item><list-item><p id="P97">= <italic>Hemicarpenteles</italic> Sarbhoy & Elphick, Trans. Brit. Mycol. Soc. 51: 156. 1968.</p></list-item><list-item><p id="P98">= <italic>Penicillium</italic> Link ex Gray <italic>sensu</italic> Pitt, The Genus <italic>Penicillium</italic>: 154. 1980 (<italic>nom</italic>. <italic>inval</italic>., art 13e).</p></list-item><list-item><p id="P99">= <italic>Chromocleista</italic> Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 101. 1993.</p></list-item></list></p><p id="P100">Subgenus <bold><italic>Aspergilloides</italic></bold> Dierckx, Annls. Soc. Scient. Brux. 25: 85. 1901.</p><p id="P101"><list list-type="simple"><list-item><p id="P102">= Subgenus <italic>Monoverticillium</italic> Biourge, Cellule 33: 265. 1923.</p></list-item><list-item><p id="P103">= Subgenus <italic>Furcatum</italic> Pitt, The Genus <italic>Penicillium</italic>: 233. 1980.</p></list-item></list></p><p id="P104">Subgenus <bold><italic>Penicillium</italic></bold></p><p id="P105"><list list-type="simple"><list-item><p id="P106">= Subgenus <italic>Eupenicillium</italic> Dierckx, Annls Soc. Scient. Brux. 25: 85. 1901.</p></list-item></list></p><sec id="S41"><title>Chromocleista</title><p id="P107">The genus <italic>Chromocleista</italic>, defined by the type species <italic>C. malachitea</italic>, belongs to <italic>Penicillium</italic> and is related to <italic>P. herquei</italic> (see Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F7">7</xref>). This genus was created by Yaguchi <italic>et al.</italic> (<xref ref-type="bibr" rid="R199">1993</xref>) for species that form bright coloured sclerotioid cleistothecia with a <italic>Geosmithia</italic> anamorph (<xref ref-type="fig" rid="F4">Fig. 4</xref>). The close relationship with <italic>Eupenicillium</italic> was noted in the original description, but the presence of the <italic>Geosmithia</italic> anamorph was, according to the authors, sufficient to create a new genus. Using 18S rDNA sequence data, Ogawa & Sugiyama (<xref ref-type="bibr" rid="R107">2000</xref>) showed that <italic>C. malachitea</italic> groups with <italic>Eupenicillium javanicum, E. crustaceum, P. chrysogenum</italic> and <italic>Geo. namyslowskii</italic>. Furthermore, they indicated that the <italic>Geosmithia</italic>-anamorph of <italic>Chromocleista malachitea</italic> resembles <italic>P. herquei</italic> and the former species could be placed in synonymy. Comparison of the β-tubulin sequences and <italic>RPB2</italic> sequences of the (neo)type cultures of <italic>P. herquei</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=336.48&link_type=cbs">CBS 336.48</ext-link><sup>NT</sup> and <italic>C. malachitea</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=647.95&link_type=cbs">CBS 647.95</ext-link><sup>T</sup> showed homologies of 92.8 % and 94.7 % respectively. Furthermore, a BLAST search with the ITS, <italic>RPB2</italic> and β-tubulin sequence data of <italic>C. malachitea</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=647.95&link_type=cbs">CBS 647.95</ext-link><sup>T</sup> on GenBank and local databases did not retrieve any high similarity matches with other described species and therefore this species is combined with <italic>Penicillium</italic> below.</p><fig id="F4" position="float"><label>Fig. 4.</label><caption><p>A–F. <italic>Penicillium malachiteum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=647.95&link_type=cbs">CBS 647.95</ext-link><sup>HT</sup>. A. Colonies grown for 7 d at 25 °C, from left to right: MEA, CYA, YES, DG18. B–D. Conidiophores. E. Immature cleistothecia. F. Conidia. G–K. <italic>Penicillium sacculum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=123567&link_type=cbs">CBS 123567</ext-link>. G. Colonies grown for 7 d at 25 °C, from left to right: MEA, CYA, YES, DG18. H–J. Conidiophores. K. Conidia. Scale bar = 10 μm.</p></caption><graphic xlink:href="1fig4"></graphic></fig></sec><sec id="S42"><title>Citromyces</title><p id="P108"><italic>Citromyces</italic> was introduced by Wehmer (<xref ref-type="bibr" rid="R195">1893</xref>) for monoverticillate <italic>Penicillium</italic> species. Many authors have agreed that this genus is a synonym of <italic>Penicillium</italic> (<xref ref-type="bibr" rid="R197">Westling 1911</xref>, <xref ref-type="bibr" rid="R18">Biourge 1923</xref>, <xref ref-type="bibr" rid="R178">Thom 1930</xref>, <xref ref-type="bibr" rid="R135">Raper & Thom 1949</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>). <italic>Citromyces</italic> largely encompasses subgenus <italic>Aspergilloides</italic> as defined by Pitt (<xref ref-type="bibr" rid="R126">1980</xref>). In our classification system, <italic>Citromyces</italic> corresponds with section <italic>Aspergilloides</italic>.</p></sec><sec id="S43"><title>Eladia</title><p id="P109">Thom (<xref ref-type="bibr" rid="R178">1930</xref>) and Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>) regarded <italic>Penicillium sacculum</italic> Dale as a <italic>Scopulariopsis</italic>, and Smith (<xref ref-type="bibr" rid="R157">1961b</xref>) introduced the genus <italic>Eladia</italic> to accommodate this species and typified it with <italic>E. saccula</italic>. Smith (<xref ref-type="bibr" rid="R157">1961b</xref>) did not indicate why this species should not be considered a <italic>Penicillium</italic>. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) accepted the positioning of <italic>E. saccula</italic> in a separate genus and he noted that this genus is closely related to <italic>Penicillium</italic>, but differing in three features (<xref ref-type="fig" rid="F4">Fig. 4</xref>): a) the phialides are born irregularly on stipes, b) phialides have a short collula and distinct thickening of the wall; c) the conidial chains are very short. Stolk & Samson (<xref ref-type="bibr" rid="R167">1985</xref>) did not accept this genus and transferred <italic>E. saccula</italic> to <italic>Penicillium</italic> and this position was retained in the list of accepted species in <italic>Trichocomaceae</italic> (<xref ref-type="bibr" rid="R130">Pitt <italic>et al.</italic> 2000</xref>). Our molecular data support the positioning of Smith's neotype of <italic>Eladia succula</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=231.61&link_type=cbs">CBS 231.61</ext-link><sup>NT</sup>) in <italic>Penicillium</italic> (Figs <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F7">7</xref>). This species is most closely related to <italic>P. canescens</italic> and <italic>P. atrovenetum</italic> (<xref ref-type="fig" rid="F7">Fig. 7</xref>, clades 24, 25). The relationship of <italic>P. sacculum</italic> with these species (and also with <italic>e.g. P. janczewskii</italic>) was also suggested by Stolk & Samson (<xref ref-type="bibr" rid="R167">1985</xref>), who emphasised that all these species have swollen phialides with an abruptly narrowed neck and often short conidial chains.</p><p id="P110">Six species were described in <italic>Eladia</italic>: <italic>E. saccula, E. inflata, E. minima, E. striatispora, E. pachyphialis</italic> and <italic>E. tibetensis</italic>. The current name for <italic>Eladia saccula</italic> is <italic>Penicillium sacculum</italic> Dale (<xref ref-type="bibr" rid="R23">1926</xref>). Ex-type strains of <italic>E. inflata</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127833&link_type=cbs">CBS 127833</ext-link>) and <italic>E. minima</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=127834&link_type=cbs">CBS 127834</ext-link>) were examined and comparison of the <italic>RPB2</italic> region (<xref ref-type="fig" rid="F8">Fig. 8</xref>) showed that <italic>E. inflata</italic> and <italic>P. fuscum</italic> (= <italic>E. pinetorum</italic>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=295.62&link_type=cbs">CBS 295.62</ext-link><sup>T</sup>) are closely related. <italic>Eladia minima</italic> is closely related to <italic>P. heteromorphum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=226.89&link_type=cbs">CBS 226.89</ext-link><sup>T</sup>) and <italic>P. philippinense</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=623.72&link_type=cbs">CBS 623.72</ext-link><sup>T</sup>). <italic>Eladia minima</italic> is closely related to <italic>P. heteromorphum, P. restrictum, Eup. katangense</italic> and <italic>Eup. philippinense</italic> (data not shown). More research is needed to determine species boundaries in this group of phylogenetical related species. No living ex-type material could be obtained for <italic>Eladia striatispora</italic>. Drawings of <italic>E. striatispora</italic> show a clear resemblance with <italic>P. striatisporum</italic>, and therefore <italic>E. striatispora</italic> is regarded as a synonym of <italic>P. striatisporum</italic> (<xref ref-type="bibr" rid="R164">Stolk 1969</xref>, <xref ref-type="bibr" rid="R95">Matsushima 1971</xref>, <xref ref-type="bibr" rid="R68">Kobayasi 1971</xref>). No type material could be obtained from <italic>E. pachyphialis</italic> and <italic>E. tibetensis</italic> and their taxonomic position remains uncertain.</p></sec><sec id="S44"><title><italic>Eupenicillium</italic> and <italic>Carpenteles</italic></title><p id="P111">The genus <italic>Eupenicillium</italic> was introduced by Ludwig (<xref ref-type="bibr" rid="R88">1892</xref>) for an ascomycete species that Brefeld (<xref ref-type="bibr" rid="R19">1874</xref>) described and illustrated as <italic>P. crustaceum</italic>. Unaware of Ludwig's publication, Langeron (<xref ref-type="bibr" rid="R76">1922</xref>) introduced the genus <italic>Carpenteles</italic> for ascus-producing <italic>Penicillium</italic> species. Because we include sexual and asexual species in our definition of <italic>Penicillium, Eupenicillium</italic> and <italic>Carpenteles</italic> are considered synonyms of <italic>Penicillium</italic>. In most cases a <italic>Penicillium</italic> anamorph name is already available for these <italic>Eupenicillium</italic> species; however, in the case of <italic>E. bovifimosum</italic> and <italic>E. saturniforme</italic>, only the teleomorph was described and no <italic>Penicillium</italic> names linked to these species exist (<xref ref-type="bibr" rid="R180">Tuthill & Frisvad 2002</xref>, <xref ref-type="bibr" rid="R193">Wang & Zhuang 2009</xref>). The new combinations <italic>Penicillium bovifimosum</italic> and <italic>Penicillium saturniforme</italic> are proposed below for these two species.</p></sec><sec id="S45"><title>Hemicarpenteles</title><p id="P112">The genus <italic>Hemicarpenteles</italic> was created by Sarbhoy & Elphick (<xref ref-type="bibr" rid="R150">1968</xref>) and <italic>H. paradoxus</italic> was designated as type (IMI 117502<sup>T</sup> = <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=793.68&link_type=cbs">CBS 793.68</ext-link><sup>T</sup>). This species is characterised by the presence of an <italic>Aspergillus</italic> anamorph and sclerotioid ascomata (<xref ref-type="fig" rid="F5">Fig. 5</xref>). This unique combination led to the proposition of a new genus. If only ascoma development and characteristics were considered, then <italic>H. paradoxus</italic> is most similar to <italic>Eupenicillium</italic>, because both genera form sclerotioid cleistothecia that ripen from the centre outwards (<xref ref-type="bibr" rid="R150">Sarbhoy & Elphick 1968</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>). <xref ref-type="fig" rid="F1">Figure 1</xref> shows the phylogenetic positioning of <italic>H. paradoxus</italic> in the genus <italic>Penicillium</italic>. The placement of this species in <italic>Penicillium</italic> is remarkable, since this species has an <italic>Aspergillus</italic> anamorph. The positioning of <italic>H. paradoxus</italic> in <italic>Penicillium</italic> is also supported by analysis of the ITS and D1/D2 regions of the 28S rDNA and partial calmodulin and β-tubulin data (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, <xref ref-type="bibr" rid="R115">2008</xref>) and the name <italic>Penicillium paradoxum</italic> will therefore be proposed (R.A. Samson, unpubl. data). The placement of an <italic>Aspergillus</italic>-type anamorph in the genus <italic>Penicillium</italic> might be confusing, when using solely phenotypic characters for identification. Three other species are described in <italic>Hemicarpenteles</italic>: <italic>H. acanthosporus, H. ornatus</italic> and <italic>H. thaxteri</italic>. The former species was transferred to <italic>Neocarpenteles acanthosporus</italic> (<xref ref-type="bibr" rid="R185">Udagawa & Uchiyama 2002</xref>) and phylogenetic studies showed that this species is related to <italic>Aspergillus</italic> section <italic>Clavati</italic> (<xref ref-type="bibr" rid="R176">Tamura <italic>et al.</italic> 2000</xref>, <xref ref-type="bibr" rid="R189">Varga <italic>et al.</italic> 2007</xref>, <xref ref-type="bibr" rid="R113">Peterson 2000b</xref>, <xref ref-type="bibr" rid="R115">2008</xref>). <italic>Hemicarpenteles ornatus</italic> and <italic>H. thaxteri</italic> are currently classified in <italic>Sclerocleista</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>, clade 7) (<xref ref-type="bibr" rid="R130">Pitt <italic>et al.</italic> 2000</xref>).</p><fig id="F5" position="float"><label>Fig. 5.</label><caption><p>A–G. <italic>Penicillium kewense</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=344.61&link_type=cbs">CBS 344.61</ext-link><sup>T</sup>. A. Colonies grown for 7 d at 25 °C, from left to right: MEA, CYA, YES, DG18. B–C. Cleistothecia. D-E. Conidiophores. F. Conidia. G. Ascospores. H–N. <italic>Aspergillus paradoxus</italic> (= <italic>P. paradoxum</italic>, R.A. Samson unpubl. results) <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=130295&link_type=cbs">CBS 130295</ext-link>. H. Colonies grown for 7 d at 25 °C, from left to right: MEA (14 d), CYA, YES, DG18. I. Detail of conidiophores. J. Cleistothecia. K–L. Conidiophores. M. Ascospores. N. Conidia. Scale bar = 10 μm.</p></caption><graphic xlink:href="1fig5"></graphic></fig></sec><sec id="S46"><title>Thysanophora</title><p id="P113"><italic>Thysanophora</italic> was proposed by Kendrick (<xref ref-type="bibr" rid="R65">1961</xref>), based on <italic>Haplographium penicillioides. Haplographium penicillioides</italic> was transferred to <italic>Thysanophora</italic> because this species produces conidia from phialides in a basipetal succession and in dry chains, while <italic>Haplographium</italic> species produce ameroconidia in slime. Roumeguère (<xref ref-type="bibr" rid="R139">1890</xref>) noted in his description of <italic>H. penicillioides</italic> that this species also forms <italic>Penicillium</italic>-like conidiophores (“l'appareil fructifère ressemble à celui d'un <italic>Penicillium</italic>”). Preuss' description of three new <italic>Penicillium</italic> species (<italic>P. finitimum, P. flexuosum</italic> and <italic>P. fuscipes</italic>) in 1851 from pine needles might be the first report of members <italic>Thysanophora</italic>. The habitat and descriptions certainly indicate this placement, but unfortunately, no type specimens were maintained (<xref ref-type="bibr" rid="R65">Kendrick 1961</xref>).</p><p id="P114"><italic>Thysanophora</italic> species produce dark coloured colonies, have dark and stout conidiophores and the majority of species have secondary growth of the stipe by means of the proliferation of an apical penicillius (<xref ref-type="fig" rid="F6">Fig. 6</xref>). Based on the combined <italic>RPB1, RPB2, Tsr1</italic> and <italic>Cct8</italic> data, it is clear that members of the genus <italic>Thysanophora</italic> belong to <italic>Penicillium.</italic> Members of this genus form a separate clade within this genus (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F7">7</xref>), confirming earlier results using rDNA sequences (<xref ref-type="bibr" rid="R62">Iwamoto <italic>et al</italic>. 2002</xref>, <xref ref-type="bibr" rid="R122">Peterson & Sigler 2002</xref>). Although stipe pigmentation of <italic>Thysanophora</italic> species is brown, this feature is thus not a useful phylogenetic character for separating this genus from <italic>Penicillium</italic> (<xref ref-type="bibr" rid="R62">Iwamoto <italic>et al</italic>. 2002</xref>). Melanised conidiophores appear in two separated lineages in <italic>Penicillium</italic>, namely in <italic>Thysanophora,</italic> and in a second lineage centered on <italic>P. stolkiae</italic> (<xref ref-type="bibr" rid="R122">Peterson & Sigler 2002</xref>). Another characteristic of <italic>Thysanophora</italic> is the secondary growth of the stipes. This character is not present in any other <italic>Penicillium</italic> species and could be argued as a feature sufficient to keep <italic>Thysanophora</italic> as a separate genus. However, that would create a paraphyletic clade in <italic>Penicillium</italic> or the need for at least eight genera to restore monophyly. To avoid both scenarios it is chosen here to transfer this genus to <italic>Penicillium. Thysanophora</italic> comprises eight accepted species, namely <italic>T. longispora, T. canadensis, T. taxi, T. striatispora, T. asymmetrica, T. verrucosa, T. glaucoalbida</italic> and <italic>T. taiwanensis</italic> (<xref ref-type="bibr" rid="R98">Minter 2007</xref>). <italic>Thysanophora penicillioides</italic> is regarded as a synonym of <italic>T. glauco-albida</italic>, because following the ICBN, the latter epithet has priority (<xref ref-type="bibr" rid="R100">Morelet 1968</xref>, <xref ref-type="bibr" rid="R98">Minter 2007</xref>). No type material was present in the CBS culture collection of <italic>T. striatispora, T. asymmetrica, T. verrucosa, T. glaucoalbida</italic> and <italic>T. taiwanensis</italic>. Only the species descriptions were studied and the species delimitation of Mercado-Sierra (<xref ref-type="bibr" rid="R97">1998</xref>) is largely followed. With exception of <italic>T. taxi</italic>, which was originally described as <italic>Penicillium taxi</italic> (<xref ref-type="bibr" rid="R151">Schneider 1956</xref>), all accepted species of <italic>Thysanophora</italic> are transferred here to <italic>Penicillium</italic> and new combinations are proposed below.</p><fig id="F6" position="float"><label>Fig. 6.</label><caption><p>A–H. <italic>Penicillium glaucoalbidum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=292.60&link_type=cbs">CBS 292.60</ext-link>. A. Colonies grown for 7 d at 25 °C, from left to right: MEA, CYA, YES, DG18. B–D. Conidiophores. E. Conidia. F–J. <italic>Penicillium lagena</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=337.97&link_type=cbs">CBS 337.97</ext-link>. F. Colonies grown for 7 d at 25 °C, from left to right: MEA, CYA, YES, DG18. G–I. Conidiophores. J. Conidia. Scale bar = 10 μm.</p></caption><graphic xlink:href="1fig6"></graphic></fig></sec><sec id="S47"><title>Torulomyces</title><p id="P115">The genus <italic>Torulomyces</italic> was erected for two species (<italic>T. lagena</italic> and <italic>T. viscosus</italic>) which form dry connected chains in a basipetal manner (<xref ref-type="bibr" rid="R24">Delitsch 1943</xref>). Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>) transferred <italic>Torulomyces lagena</italic>, the type species, to <italic>Penicillium</italic>. This transfer was based on morphological similarities, such as the phialide shape and cultural appearances (<xref ref-type="fig" rid="F6">Fig. 6</xref>). Later, Pitt & Samson (<xref ref-type="bibr" rid="R129">1993</xref>) did not accept this transfer to <italic>Penicillium</italic>, and <italic>Torulomyces</italic> was re-instated. Our phylogenetic data support Stolk & Samson's (<xref ref-type="bibr" rid="R166">1983</xref>) proposal to transfer <italic>Torulomyces</italic> to <italic>Penicillium</italic> and other species described in <italic>Torulomyces</italic> need to be combined with <italic>Penicillium</italic>.</p><p id="P116">Currently, eight species are described in <italic>Torulomyces</italic>: <italic>T. brunneus, T. indicus, T. laevis, T. lagena, T. macrosporus, T. ovatus, T. parviverrucosus</italic> and <italic>T. viscosus</italic>. Isolate <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.65&link_type=cbs">CBS 185.65</ext-link> was designated as the neotype of <italic>P. lagena</italic>, and <italic>Eupenicillium limoneum</italic> was considered to be the teleomorph of this species (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>). Unfortunately, the ex-type culture of <italic>E. limoneum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=650.82&link_type=cbs">CBS 650.82</ext-link><sup>T</sup>) maintained in the CBS collection is dead. Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>) are followed here and <italic>E. limoneum</italic> is kept in synonymy with <italic>P. lagena</italic>. Delitsch's species <italic>Torulomyces viscosus</italic> remains doubtful since no type material is available and the diagnosis lacks critical details (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>, <xref ref-type="bibr" rid="R4">Ando <italic>et al.</italic> 1998</xref>). No ex-type material of <italic>Torulomyces macrosporus</italic> could be obtained; based on its protologue (<xref ref-type="bibr" rid="R96">Matsushima 1987</xref>), <italic>T. macrosporum</italic> may belong to <italic>Monocillium</italic> (<xref ref-type="bibr" rid="R4">Ando <italic>et al.</italic> 1998</xref>).<italic>Torulomyces laevis, T. ovatus</italic> and <italic>T. parviverrucosus</italic> were described by Ando <italic>et al.</italic> (<xref ref-type="bibr" rid="R4">1998</xref>) and in the same publication <italic>Monocillium humicola</italic> var. <italic>brunneum</italic> was combined with <italic>T. brunneus</italic>. The type strain of <italic>T. brunneus</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=382.64&link_type=cbs">CBS 382.64</ext-link><sup>T</sup> is closely related to <italic>Torulomyces lagena</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.65&link_type=cbs">CBS 185.65</ext-link><sup>NT</sup>; these isolates have identical ITS sequences, but differ in their partial β-tubulin, calmodulin and <italic>RPB2</italic> sequences (ITS 100 %; calmodulin 98.3 % and β-tubulin 98.4 % and <italic>RPB2</italic> 98.3 %; unpubl. data). Ando <italic>et al.</italic> (<xref ref-type="bibr" rid="R4">1998</xref>) is followed here and this species is kept as separate. No type material of <italic>T. laevis, T. ovatus</italic> and <italic>T. parviverrucosus</italic> was available for analysis, but a detailed study of the species descriptions suggests they warrant separate species status. New combinations in <italic>Penicillium</italic> are proposed below. Various isolates with similar morphology to <italic>P. lagena</italic> are maintained in the CBS collections (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=185.65&link_type=cbs">CBS 185.65</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=382.64&link_type=cbs">CBS 382.64</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=287.66&link_type=cbs">CBS 287.66</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=337.97&link_type=cbs">CBS 337.97</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=120415&link_type=cbs">CBS 120415</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=110532&link_type=cbs">CBS 110532</ext-link>, DTO 82A8, DTO 92D1), and preliminary sequencing results show a sequence variation among these strains, suggesting the presence of multiple species (unpubl. data). A thorough taxonomic study should be preformed to elucidate the species diversity in this clade.</p><p id="P117">The genus <italic>Monocillium</italic> needs further attention. This genus was established for a single species, <italic>M. indicum</italic> (<xref ref-type="bibr" rid="R141">Saksena 1955</xref>). Based on conidium morphogenesis, Hashmi <italic>et al.</italic> (<xref ref-type="bibr" rid="R48">1972</xref>) placed <italic>Monocillium</italic> in synonymy with <italic>Torulomyces</italic>, and later Kendrick & Carmichael (<xref ref-type="bibr" rid="R66">1973</xref>) made the combination <italic>Torulomyces indicus</italic>. However, a BLAST search with the ITS sequence of the type strain of <italic>M. indicum</italic> (UAMH 1499, GenBank GQ169328) showed that the closest relatives are among <italic>Hypocreaceae</italic> (<xref ref-type="bibr" rid="R155">Sigler <italic>et al.</italic> 2010</xref>). This is in agreement with Gams (<xref ref-type="bibr" rid="R42">1971</xref>), who showed that <italic>Monocillium</italic> species are anamorphs related to <italic>Niesslia</italic> species.</p></sec></sec></sec><sec id="S48"><title>Part Three: sectional delimitation within <italic>Penicillium s. str.</italic></title><sec id="S49"><title>Classification</title><p id="P118">Dierckx (<xref ref-type="bibr" rid="R25">1901</xref>) proposed the first infrageneric classification of <italic>Penicillium</italic> and introduced the subgenera <italic>Aspergilloides, Biverticillium</italic> and <italic>Eupenicillium</italic> (<xref ref-type="bibr" rid="R18">Biourge 1923</xref>). Biourge (<xref ref-type="bibr" rid="R18">1923</xref>) expanded this subdivision and accepted two subgenera, two sections, four series and six subsections. The sections <italic>Bulliardium</italic> (<italic>Asymetrica</italic>) was introduced by Biourge (<xref ref-type="bibr" rid="R18">1923</xref>) and in this section <italic>Penicillium</italic> species with branched conidiophores were included. No type species was designated and species with terverticillate conidiophores belong to Biourge's definition of his section <italic>Bulliardium</italic> (<italic>Asymetrica</italic>). We decided to synonymise this section with section <italic>Penicillium</italic>. The section <italic>Biverticillium</italic> belongs to <italic>Talaromyces s. str.</italic> and is not treated here (<xref ref-type="fig" rid="F1">Fig. 1</xref>). In the classical work of Thom (<xref ref-type="bibr" rid="R178">1930</xref>: 155–159), <italic>Penicillium</italic> is divided in four subgenera (although not named as such), and 12 sections and 17 subsections. Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>) introduced various new sections, subsections and series and Ramírez (<xref ref-type="bibr" rid="R132">1982</xref>) largely followed Raper and Thom's classification. Neither provided Latin descriptions for their newly introduced sections (and series), and these names are therefore regarded as <italic>nomen invalidum</italic> are not considered further here. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) divided <italic>Penicillium</italic> into four subgenera, 10 sections and 21 series. Five years later, Stolk & Samson (<xref ref-type="bibr" rid="R167">1985</xref>) proposed another taxonomic scheme for <italic>Penicillium</italic> anamorphs. In the latter taxonomic scheme, both sexual and asexual species were treated. More recently, Samson & Frisvad (<xref ref-type="bibr" rid="R147">2004</xref>) revised subgenus <italic>Penicillium</italic> and five sections and 17 series were recognised. An overview of sections and their type species of the studies of Thom (<xref ref-type="bibr" rid="R178">1930</xref>), Pitt (<xref ref-type="bibr" rid="R126">1980</xref>), Stolk & Samson (<xref ref-type="bibr" rid="R167">1985</xref>) and Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) is shown in <xref ref-type="table" rid="T5">Table 5</xref>.</p><table-wrap id="T5" position="float"><label>Table 5</label><caption><p>Overview of sectional classification in different studies of <italic>Penicillium</italic></p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="2" align="center" valign="top" rowspan="1">Thom (<xref ref-type="bibr" rid="R178">1930</xref>)</th><th colspan="2" align="center" valign="top" rowspan="1">Pitt (<xref ref-type="bibr" rid="R126">1980</xref>)</th><th colspan="2" align="center" valign="top" rowspan="1">Stolk & Samson (<xref ref-type="bibr" rid="R167">1985</xref>)</th><th colspan="2" align="center" valign="top" rowspan="1">Current study</th></tr><tr><th align="left" valign="top" rowspan="1" colspan="1">Section</th><th align="left" valign="top" rowspan="1" colspan="1">Type species</th><th align="left" valign="top" rowspan="1" colspan="1">Section</th><th align="left" valign="top" rowspan="1" colspan="1">Type species</th><th align="left" valign="top" rowspan="1" colspan="1">Section</th><th align="left" valign="top" rowspan="1" colspan="1">Type species</th><th align="left" valign="top" rowspan="1" colspan="1">Section</th><th align="left" valign="top" rowspan="1" colspan="1">Type species</th></tr></thead><tbody><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ascogena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. luteum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergilloides</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. aurantiobrunneum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergilloides</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. glabrum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Aspergilloides</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. aurantiobrunneum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Brevi-compacta</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. brevicompactum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Coremigenum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. duclauxii</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Biverticillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. minioluteum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Brevicompacta</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. olsonii</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Coremigena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. duclauxii</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Coronatum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. olsonii</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Coremigenum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. duclauxii</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Canescentia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. canescens</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fasciculata</italic></td><td align="left" valign="top" rowspan="1" colspan="1">Fasiculate Penicillia e.g. <italic>P. hirsutum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cylindrosporum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. italicum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Divaricatum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. janthinellum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Charlesii</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. charlesii</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Funiculosa</italic></td><td align="left" valign="top" rowspan="1" colspan="1">Undefined; similar to <italic>Lanata-divaricata</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Divaricatum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. janthinellum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eladia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. sacculum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Chrysogena</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. chrysogenum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Lanata-divaricata</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. janthinellum</italic>-type</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Exilicaulis</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. restrictum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Geosmithia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. lavendulum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Cinnamopurpurea</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. cinnamopurpureum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Lanata-typica</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. camemberti</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Furcatum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. oxalicum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Inordinate</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. arenicola</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Citrina</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. citrinum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Luteo-virida</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. minioluteum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Inordinate</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. arenicola</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. expansum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Digitata</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. digitatum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Miscellanea</italic></td><td align="left" valign="top" rowspan="1" colspan="1">Miscellaneous species and genera</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. expansum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ramosum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. lanosum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Eladia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. sacculum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">(Monoverticillata)-<italic>stricta</italic></td><td align="left" valign="top" rowspan="1" colspan="1">Undefined section</td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Simplicium</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. minioluteum</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Torulomyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. lagena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Exilicaulis</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. restrictum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">(Monoverticillata)-<italic>Ramigena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Citromyces</italic> species</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fasciculata</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. viridicatum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"><italic>Velutina</italic></td><td align="left" valign="top" rowspan="1" colspan="1">Undefined section</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Fracta</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. fractum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Gracilenta</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. gracilentum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Lanata-divaricata</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. janthinellum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ochrosalmonea</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. ochrosalmoneum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Paradoxa</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>A. paradoxus</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Penicillium</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. expansum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ramigena</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. cyaneum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Ramosa</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. lanosum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Roguefortorum</italic><sup><xref ref-type="table-fn" rid="TFN7">*</xref></sup></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. roqueforti</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Sclerotiora</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. sclerotiorum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Stolkia</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. stolkiae</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Thysanophora</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>S. glauco-albidum</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Torulomyces</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. lagena</italic></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>Turbata</italic></td><td align="left" valign="top" rowspan="1" colspan="1"><italic>P. turbatum</italic></td></tr></tbody></table><table-wrap-foot><fn id="TFN7"><label>*</label><p id="P119">Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) divided subgenus <italic>Penicillium</italic> in six sections. This sectional classification is supported by extrolite, phenotypic and physiological data and their subdivision is followed here. The results of our analysis based on partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) do not confirm these sections; however, partial β-tubulin data largely confirmed their polyphasic classification (Samson <italic>et al</italic>. 2004).</p></fn></table-wrap-foot></table-wrap><p id="P120">The classification of <italic>Eupenicillium</italic> does not have such a long history: Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) was the first, and introduced eight series. In the monograph of Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>), four sections were introduced for the grouping of the <italic>Eupenicillium</italic> species and Pitt's infrageneric concept of classifying species in series was abandoned.</p></sec><sec id="S50"><title>Accepted species and their position in the sections of Penicillium</title><p id="P121">The phylogenetic relationship among <italic>Penicillium s. str.</italic> was studied using combined sequence data of four loci. Based on these results (<xref ref-type="fig" rid="F7">Fig. 7</xref>), <italic>Penicillium</italic> is subdivided into two subgenera and 25 sections. An overview of these sections is presented in <xref ref-type="table" rid="T5">Table 5</xref>, together with the type species of each section. In our study, a new sectional subdivision is proposed and older names at different ranks (<italic>e.g.</italic> subgeneric, subsection and series names) and invalid names (<xref ref-type="bibr" rid="R135">Raper & Thom 1949</xref>, <xref ref-type="bibr" rid="R132">Ramírez 1982</xref>) are not considered. Assignment of the species to the various sections was mainly based on the overviews presented in Figs <xref ref-type="fig" rid="F8">8</xref> and <xref ref-type="fig" rid="F10">10</xref>, <xref ref-type="fig" rid="F11">11</xref>, <xref ref-type="fig" rid="F12">12</xref>, <xref ref-type="fig" rid="F13">13</xref> and other published molecular-based data. The accepted <italic>Penicillium</italic> and <italic>Eupenicillium</italic> species mentioned in the list of “accepted species and their synonyms in <italic>Trichocomaceae</italic>” (<xref ref-type="bibr" rid="R130">Pitt <italic>et al</italic>. 2000</xref>) were used as a starting point for dividing the species among the various sections, updated species described after 2000. In various cases, the same <italic>Penicillium</italic> and <italic>Eupenicillium</italic> species share the same ex-type specimen. However, if the type material of the <italic>Penicillium</italic> morph differs from the <italic>Eupenicillium</italic> morph, then both ex-type strains were included in the study and additional comments are given in the text.</p><sec id="S51"><title>Clade 1: section <italic>Aspergilloides</italic></title><p id="P122"><list list-type="simple"><list-item><p id="P123">= <italic>Eupenicillium</italic> sect. <italic>Pinetorum</italic> (Pitt) Stolk & Samson, Stud. Mycol. 23: 88. 1983.</p></list-item></list></p><p id="P124">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic>.</p><p id="P125">Type: <italic>Penicillium aurantiobrunneum</italic> Dierckx</p><p id="P126">Most members of this section grow quickly on agar media, form velvety colonies and are predominantly monoverticillate. This section corresponds to group 2 of Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>). Two teleomorph species are positioned in this section: <italic>P. fuscum</italic> and <italic>P. saturniforme</italic>. Stolk (1968) found ascospores in an old culture of the type strain of <italic>P. pinetorum</italic> and described the ascosporic state as <italic>Eupenicillium pinetorum</italic>. Later, the anamorph of <italic>E. pinetorum</italic> was linked to <italic>P. fuscum</italic> (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>); the latter name is older than <italic>P. pinetorum</italic> and therefore used here. The taxonomic position of <italic>P. lapidosum</italic> warrants further attention. Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>) suggested that this species is conspecific with <italic>P. thomii</italic>. However, our results show that the type strain of this species (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=343.48&link_type=cbs">CBS 343.48</ext-link><sup>T</sup>) is phylogenetically related to <italic>P. namyslowskii</italic> (<xref ref-type="fig" rid="F7">Fig. 7</xref>, clade 10) and therefore unrelated to section <italic>Aspergilloides</italic>. Based on the data presented in <xref ref-type="fig" rid="F8">Fig. 8</xref> and literature (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, <xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>, <xref ref-type="bibr" rid="R193">Wang & Zhuang 2009</xref>, <xref ref-type="bibr" rid="R12">Barreto <italic>et al.</italic> 2011</xref>), we place the following species in section <italic>Aspergilloides</italic>:</p><p id="P127"><list list-type="simple"><list-item><p id="P128"><italic>Penicillium ardesiacum</italic> Novobranova, Novosti Sist. Nizs. Rast. 11: 228. 1974.</p></list-item><list-item><p id="P129"><italic>Penicillium asperosporum</italic> Smith, Trans. Br. Mycol. Soc. 48: 275. 1965.</p></list-item><list-item><p id="P130"><italic>Penicillium crocicola</italic> Yamamoto, Scient. Rep. Hyogo Univ. Agric., Agric. Biol. Ser. 2, 2: 28. 1956.</p></list-item><list-item><p id="P131"><italic>Penicillium fuscum</italic> (Sopp) Biourge, Cellule 33: 103. 1923 (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>).</p></list-item><list-item><p id="P132"><italic>Penicillium georgiense</italic> Peterson & Horn, Mycologia 101: 79. 2009.</p></list-item><list-item><p id="P133"><italic>Penicillium glabrum</italic> (Wehmer) Westling, Ark. Bot. 11: 131. 1911 (syn. <italic>P. terlikowskii</italic>; <xref ref-type="bibr" rid="R12">Barreto <italic>et al</italic>. 2011</xref>).</p></list-item><list-item><p id="P134"><italic>Penicillium kananaskense</italic> Seifert, Frisvad & McLean, Can. J. Bot. 72: 20. 1994 (unpubl. data, K.A. Seifert).</p></list-item><list-item><p id="P135"><italic>Penicillium lapatayae</italic> Ramírez, Mycopathol. 91: 96. 1985 (<xref ref-type="bibr" rid="R39">Frisvad <italic>et al</italic>. 1990c</xref>).</p></list-item><list-item><p id="P136"><italic>Penicillium lividum</italic> Westling, Ark. Bot. 11: 134. 1911.</p></list-item><list-item><p id="P137"><italic>Penicillium montanense</italic> Christensen & Backus, Mycologia 54: 574. 1963.</p></list-item><list-item><p id="P138"><italic>Penicillium odoratum</italic> Christensen & Backus, Mycologia 53: 459. 1962 (this study, <xref ref-type="fig" rid="F8">Fig. 8</xref>).</p></list-item><list-item><p id="P139"><italic>Penicillium palmense</italic> Ramírez & Martínez, Mycopathol. 66: 80. 1978.</p></list-item><list-item><p id="P140"><italic>Penicillium patens</italic> Pitt & Hocking, Mycotaxon 22: 197. 1985.</p></list-item><list-item><p id="P141"><italic>Penicillium quercetorum</italic> Baghdadi, Nov. Sist. Niz. Rast. 5: 110. 1968.</p></list-item><list-item><p id="P142"><italic>Penicillium saturniforme</italic> (Wang & Zhuang) Houbraken & Samson, Stud. Mycol. 70: 48. 2011 (this study).</p></list-item><list-item><p id="P143"><italic>Penicillium spinulosum</italic> Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 76. 1910.</p></list-item><list-item><p id="P144"><italic>Penicillium subericola</italic> Barreto, Frisvad & Samson, Fungal Diversity 49: 32. 2011.</p></list-item><list-item><p id="P145"><italic>Penicillium thiersii</italic> Peterson, Bayer & Wicklow, Mycologia 96: 1283. 2004.</p></list-item><list-item><p id="P146"><italic>Penicillium thomii</italic> Maire, Bull. Soc. Hist. Nat. Afrique N. 8: 189. 1917.</p></list-item></list></p></sec><sec id="S52"><title>Clade 2: section <italic>Sclerotiora</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563124&link_type=mb">MB563124</ext-link>.</title><p id="P147">Sectio in Penicillio subgen. Aspergilloide. Mycelio saepe colorato, plus minusve flavido et/vel aurantiaco. Sclerotis/cleistotheciis claris. colore.</p><p id="P148">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P149">Type: <italic>Penicillium sclerotiorum</italic> van Beyma</p><p id="P150">Members of section <italic>Sclerotiora</italic> generally have monoverticillate conidiophores; however, exceptions are <italic>P. malachiteum, P. nodositatum</italic> and <italic>P. herquei</italic>, which form symmetrically biverticillate conidiophores. The mycelium of members of sect. <italic>Sclerotiora</italic> is pigmented in shades of yellow and/or orange, reverse colony colours in shades of yellow, orange or red, and sclerotia and cleistothecia are, if present, bright coloured. Species belonging to this section occur regularly in and are abundant upon substrata exposed to soil. This section corresponds with group 3 of Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>). Our list of species belonging to this section was composed based on the data presented in <xref ref-type="fig" rid="F8">Fig. 8</xref> and studies by Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>), Peterson <italic>et al.</italic> (<xref ref-type="bibr" rid="R121">2003</xref>, <xref ref-type="bibr" rid="R116">2004</xref>), Peterson & Horn (<xref ref-type="bibr" rid="R118">2009</xref>), Nonaka <italic>et al</italic>. (<xref ref-type="bibr" rid="R103">2011</xref>) and Rivera & Seifert (<xref ref-type="bibr" rid="R136">2011</xref>). Isolate NRRL 2060 is included in <xref ref-type="fig" rid="F8">Fig. 8</xref> and Peterson & Horn (<xref ref-type="bibr" rid="R118">2009</xref>) treated this strain as the type of <italic>P. multicolor</italic>. However, Raper & Thom's (<xref ref-type="bibr" rid="R135">1949</xref>) isolates of <italic>P. multicolor</italic> differ in significant features from the original description of Grigorieva-Manoilova & Poradielova (<xref ref-type="bibr" rid="R47">1915</xref>) (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>), and Rivera & Seifert (<xref ref-type="bibr" rid="R136">2011</xref>) treated this species as a synonym of <italic>P. fellutanum. Penicillium nodositatum</italic> shares identical partial <italic>RPB2</italic> sequences with <italic>P. bilaiae</italic> and might be conspecific with the latter species. More research is needed because the former species produces biverticillate conidiophores and the latter strictly monoverticillate structures (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R188">Valla <italic>et al.</italic> 1989</xref>).</p><p id="P151"><list list-type="simple"><list-item><p id="P152"><italic>Penicillium adametzii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 507. 1927.</p></list-item><list-item><p id="P153"><italic>Penicillium adametzioides</italic> Abe ex Smith, Trans. Br. Mycol. Soc. 46: 335. 1963.</p></list-item><list-item><p id="P154"><italic>Penicillium angulare</italic> Peterson, Bayer & Wicklow, Mycologia 96: 1289. 2004.</p></list-item><list-item><p id="P155"><italic>Penicillium bilaiae</italic> Chalabuda, Bot. Mater. Otd. Sporov. Rast. 6: 165. 1950.</p></list-item><list-item><p id="P156"><italic>Penicillium brocae</italic> Peterson, Pérez, Vega & Infante, Mycologia 95: 143. 2003.</p></list-item><list-item><p id="P157"><italic>Penicillium cainii</italic> Rivera & Seifert, Stud. Mycol. 70: 147. 2011.</p></list-item><list-item><p id="P158"><italic>Penicillium guanacastense</italic> Rivera, Urb & Seifert, Mycotaxon, <italic>in press</italic>. 2011.</p></list-item><list-item><p id="P159"><italic>Penicillium herquei</italic> Bainier & Sartory, Bull. Soc. Mycol. France 28: 121. 1912.</p></list-item><list-item><p id="P160"><italic>Penicillium hirayamae</italic> Udagawa, J. Agric. Sci. Tokyo Nogyo Daigaku 5: 6. 1959.</p></list-item><list-item><p id="P161"><italic>Penicillium jacksonii</italic> Rivera & Seifert, Stud. Mycol. 70: 151. 2011.</p></list-item><list-item><p id="P162"><italic>Penicillium johnkrugii</italic> Rivera & Seifert, Stud. Mycol. 70: 151. 2011.</p></list-item><list-item><p id="P163"><italic>Penicillium jugoslavicum</italic> Ramírez & Muntañola-Cvetkovic, Mycopathol. 88: 65. 1984.</p></list-item><list-item><p id="P164"><italic>Penicillium malachiteum</italic> (Yaguchi & Udagawa) Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P165"><italic>Penicillium mallochii</italic> Rivera, Urb & Seifert Mycotaxon, <italic>in press</italic>. 2011.</p></list-item><list-item><p id="P166"><italic>Penicillium nodositatum</italic> Valla, Plant and Soil 114: 146. 1989.</p></list-item><list-item><p id="P167"><italic>Penicillium sclerotiorum</italic> van Beyma, Zentralbl. Bakteriol., 2. Abt., 96: 418. 1937.</p></list-item><list-item><p id="P168"><italic>Penicillium viticola</italic> Nonaka & Masuma, Mycoscience 52: 339. 2011.</p></list-item></list></p></sec><sec id="S53"><title>Clade 3: section <italic>Charlesia</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563125&link_type=mb">MB563125</ext-link>.</title><p id="P169">Sectio in Penicillio subgen. Aspergilloide. Solum in CYA, conidiophoris ad apicem inflatis.</p><p id="P170">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P171">Type: <italic>Penicillium charlesii</italic> Smith</p><p id="P172">The phylogeny of this section was studied by Peterson <italic>et al</italic>. (<xref ref-type="bibr" rid="R123">2005</xref>). In the same study, an overview was presented of phenotypic characters to differentiate species within section <italic>Charlesii</italic>. It was stated that the overall phenotypic similarity of these species is striking; however, no shared characters were given. With exception of <italic>P. indicum</italic>, all members of section <italic>Charlesii</italic> grow restricted on CYA and have conidiophores with an apical swelling. Species of this section can be strictly monoverticillate, but <italic>P. charlesii</italic> and <italic>P. fellutanum</italic> can also be irregularly biverticillate. Included species are based on the data presented in <xref ref-type="fig" rid="F8">Fig. 8</xref> and Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>) and Peterson <italic>et al</italic>. (<xref ref-type="bibr" rid="R123">2005</xref>).</p><p id="P173"><list list-type="simple"><list-item><p id="P174"><italic>Penicillium charlesii</italic> Smith, Trans. Br. Mycol. Soc. 18: 90. 1933.</p></list-item><list-item><p id="P175"><italic>Penicillium coffeae</italic> Peterson, Vega, Posada & Nagai, Mycologia 97: 662. 2005.</p></list-item><list-item><p id="P176"><italic>Penicillium fellutanum</italic> Biourge, Cellule 33: 262. 1923.</p></list-item><list-item><p id="P177"><italic>Penicillium georgiense</italic> Peterson & Horn, Mycologia 101: 79. 2009.</p></list-item><list-item><p id="P178"><italic>Penicillium indicum</italic> Sandhu & Sandhu, Can. J. Bot. 41: 1273. 1963 (syn. <italic>P. gerundense</italic>, <xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>).</p></list-item><list-item><p id="P179"><italic>Penicillium phoeniceum</italic> van Beyma, Zentralbl. Bakteriol., 2. Abt., 88: 136. 1933.</p></list-item></list></p></sec><sec id="S54"><title>Clade 4: section <italic>Thysanophora</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563126&link_type=mb">MB563126</ext-link>.</title><p id="P180">Sectio in Penicillio subgen. Aspergilloide. Coloniis pullis, conidiophoris pigmentatis, compactis et incremento secundario stipitis per proliferationem penicillii apicali.</p><p id="P181">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P182">Type: <italic>Sclerotium glauco-albidum</italic> Desmazières</p><p id="P183">The genus <italic>Thysanophora</italic> is placed in synonymy with <italic>Penicillium</italic> (see above). The section is characterised by the formation dark coloured colonies, pigmented and stout conidiophores and the majority of species have secondary growth of the stipe by means of the proliferation of an apical penicillius. Nine specific epithets have been combined with <italic>Thysanophora</italic>, and eight are accepted species. Mercado-Sierra <italic>et al.</italic> (<xref ref-type="bibr" rid="R97">1998</xref>) is largely followed here and the following species belong in section <italic>Thysanophora</italic>:</p><p id="P184"><list list-type="simple"><list-item><p id="P185"><italic>Penicillium asymmetricum</italic> (Subramanian & Sudha) Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P186"><italic>Penicillium coniferophilum</italic> Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P187"><italic>Penicillium glaucoalbidum</italic> (Desmazières) Houbraken & Samson, Stud. Mycol.70: 47. 2011 (this study).</p></list-item><list-item><p id="P188"><italic>Penicillium hennebertii</italic> Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P189"><italic>Penicillium longisporum</italic> (Kendrick) Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P190"><italic>Penicillium melanostipe</italic> Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P191"><italic>Penicillium taiwanense</italic> (Matsushima) Houbraken & Samson, Stud. Mycol. 70: 48. 2011 (this study).</p></list-item><list-item><p id="P192"><italic>Penicillium taxi</italic> Schneider, Zentralblatt für Bakteriologie und Parasitenkunde, Abteilung 2, 110: 43. 1956.</p></list-item></list></p></sec><sec id="S55"><title>Clade 5: section <italic>Ochrosalmonea</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563127&link_type=mb">MB563127</ext-link>.</title><p id="P193">Sectio in Penicillio subgen. Aspergilloide. Mycelio conspicue pigmentoso, flavido; phialidibus ampulliformibus vel acerosis; conidiis apiculatis.</p><p id="P194">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P195">Type: <italic>Penicillium ochrosalmoneum</italic> Udagawa</p><p id="P196"><italic>Penicillium ochrosalmoneum</italic> and <italic>P. isariiforme</italic> are accommodated in section <italic>Ochrosalmonea</italic> (<xref ref-type="fig" rid="F5">Fig. 5</xref>, clade 5). Both species seem macroscopically dissimilar. <italic>Penicillium isariiforme</italic> grows quickly on agar media MEA and CYA (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>) and forms characteristic feather-like synnemata (<xref ref-type="bibr" rid="R148">Samson <italic>et al</italic>. 1976</xref>, <xref ref-type="fig" rid="F9">Fig. 9</xref>). In contrast, <italic>P. ochrosalmoneum</italic> isolates grow slowly on agar media and forms a velutinous colony surface (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>). However, both species form conspicuous yellow coloured mycelium, ampulliform to acerose shaped phialides and apiculate conidia. The classification of <italic>P. isariiforme</italic> in <italic>Penicillium</italic> was subject of various studies. This species was classified in subgenus <italic>Biverticillium</italic> (= <italic>Talaromyces s. str</italic>.) (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R33">Frisvad & Filtenborg 1983</xref>), but also in subgenus <italic>Penicillium</italic> (= <italic>Penicillium s. str</italic>.) (<xref ref-type="bibr" rid="R132">Ramírez 1982</xref>, <xref ref-type="bibr" rid="R148">Samson <italic>et al</italic>. 1976</xref>). <xref ref-type="fig" rid="F7">Figure 7</xref> shows that <italic>P. isariiforme</italic> phylogenetically belongs to subgenus <italic>Aspergilloides</italic> in <italic>Penicillium s. str</italic>.</p><fig id="F9" position="float"><label>Fig. 9.</label><caption><p><italic>Penicillium isariiforme</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=247.56&link_type=cbs">CBS 247.56</ext-link>. A. Colonies grown for 14 d at 25 °C, from left to right: MEA, YES, CYA. B–D. Conidiophores. E. Conidia. Scale bar = 10 μm.</p></caption><graphic xlink:href="1fig9"></graphic></fig><p id="P197">The holotype of <italic>Eup. ochrosalmoneum</italic> is <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=489.66&link_type=cbs">CBS 489.66</ext-link> and <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=231.60&link_type=cbs">CBS 231.60</ext-link> is the ex-type of <italic>P. ochrosalmoneum</italic>. The strains share identical partial <italic>RPB2</italic> sequences and therefore <italic>E. ochrosalmoneum</italic> is regarded as conspecific with <italic>P. ochrosalmoneum</italic> (<xref ref-type="fig" rid="F12">Fig. 12</xref>). Based on the data presented in <xref ref-type="fig" rid="F12">Fig. 12</xref>, the following species belong in section <italic>Ochrosalmonea</italic>.</p><p id="P198"><list list-type="simple"><list-item><p id="P199"><italic>Penicillium isariiforme</italic> Stolk & Meyer, Trans. Br. Mycol. Soc. 40: 187. 1957.</p></list-item><list-item><p id="P200"><italic>Penicillium ochrosalmoneum</italic> Udagawa, J. Agric. Sci. Tokyo Nogyo Daigaku 5: 10. 1959.</p></list-item></list></p></sec><sec id="S56"><title>Clade 6: section <italic>Cinnamopurpurea</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563128&link_type=mb">MB563128</ext-link>.</title><p id="P201">Sectio in Penicillio subgen. Aspergilloide. Sect. Ornatis similis, sed conidiophoris semper simplicibus vel biverticillate divaricates; stipitibus cum conidiophoris distincte vesiculosis.</p><p id="P202">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P203">Type: <italic>Penicillium cinnamopurpureum</italic> Udagawa</p><p id="P204">Members of section <italic>Cinnamopurpurea</italic> grow slowly on MEA and CYA and can be strictly monoverticillate, but species with biverticillate conidiophores are also present in this section. The majority of the species have distinct vesicular conidiophores. This section is phenotypically related to section <italic>Ornata</italic>; however, statistical support for this relationship is lacking in our phylogenetic analysis (<xref ref-type="fig" rid="F7">Fig. 7</xref>).</p><p id="P205"><italic>Penicillium cinnamopurpureum</italic> was originally described by Abe (<xref ref-type="bibr" rid="R1">1956</xref>) without a Latin diagnosis, and validated by Udagawa (<xref ref-type="bibr" rid="R181">1959</xref>). Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>) considered <italic>P. dierckxii</italic> the anamorph of <italic>Eupenicillium cinnamopurpureum</italic> and Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) linked <italic>P. phoeniceum</italic> to <italic>E. cinnamopurpureum</italic>. Our data show that <italic>P. phoeniceum</italic> (sect. <italic>Charlesii</italic>, <xref ref-type="fig" rid="F8">Fig. 8</xref>) and <italic>P. dierckxii</italic> (sect. <italic>Ramigena</italic>, <xref ref-type="fig" rid="F10">Fig. 10</xref>) are phylogenetically distinct from <italic>P. cinnamopurpureum</italic>. Furthermore, partial <italic>RPB2</italic> data show that the type strains of <italic>P. cinnamopurpureum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=847.68&link_type=cbs">CBS 847.68</ext-link>) and <italic>E. cinnamopurpureum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=490.66&link_type=cbs">CBS 490.66</ext-link>) are similar (<xref ref-type="fig" rid="F10">Fig. 10</xref>).</p><p id="P206"><italic>Penicillium chermesinum</italic> is also placed in this section. This species was neotypified with NRRL 2048 (= <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=231.81&link_type=cbs">CBS 231.81</ext-link>), because the type culture, NRRL 735, no longer adequately represented Biourge's protologue (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>). Molecular analysis shows that these two species are phylogenetically unrelated. The ITS-partial 28S rDNA sequences of NRRL 735<sup>T</sup> (= GenBank no. AF033413) is related to <italic>P. cinnamopurpureum</italic> (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>) while the neotype of this species, NRRL 2048<sup>NT</sup>, (AY742693) is related to <italic>P. indicum</italic> in section <italic>Charlesii</italic>. Based on the data presented in <xref ref-type="fig" rid="F10">Fig. 10</xref> and Peterson & Horn (<xref ref-type="bibr" rid="R118">2009</xref>), the following species are accommodated in <italic>Cinnamopurpurea.</italic></p><p id="P207"><list list-type="simple"><list-item><p id="P208"><italic>Penicillium chermesinum</italic> Biourge, Cellule 33: 284. 1923.</p></list-item><list-item><p id="P209"><italic>Penicillium cinnamopurpureum</italic> Udagawa, J. Agric. Food Sci., Tokyo 5: 1. 1959.</p></list-item><list-item><p id="P210"><italic>Penicillium ellipsoideosporum</italic> Wang & Kong, Mycosystema 19: 463. 2000.</p></list-item><list-item><p id="P211"><italic>Penicillium idahoense</italic> Paden, Mycopath. Mycol. Appl. 43: 261. 1971 (<xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>, this study).</p></list-item><list-item><p id="P212"><italic>Penicillium incoloratum</italic> Huang & Qi, Acta Mycol. Sin. 13: 264. 1994.</p></list-item><list-item><p id="P213"><italic>Penicillium malacaense</italic> Ramírez & Martínez, Mycopathologia 72: 186. 1980 (syn. <italic>P. ovetense</italic>, this study) (<xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>).</p></list-item><list-item><p id="P214"><italic>Penicillium nodulum</italic> Kong & Qi, Mycosystema 1: 108. 1988.</p></list-item><list-item><p id="P215"><italic>Penicillium parvulum</italic> Peterson & Horn, Mycologia 101: 75. 2009.</p></list-item><list-item><p id="P216"><italic>Penicillium shennangjianum</italic> Kong & Qi, Mycosystema 1: 110. 1988.</p></list-item></list></p></sec><sec id="S57"><title>Clade 7: section <italic>Ramigena</italic> Thom, The Penicillia: 225. 1930. In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></title><p id="P217">Type: <italic>Penicillium cyaneum</italic> (Bainier & Sartory) Biourge</p><p id="P218">This section is based on Thom's section <italic>Ramigena</italic>. Thom (<xref ref-type="bibr" rid="R178">1930</xref>) introduced this section for species where monoverticillate conidiophores are evident, but divaricate branching at various levels without a definiteness of organisation or arrangement is consistently observed. Most species illustrated by Banier & Sartory (1913) as species of <italic>Citromyces</italic> are accommodated in this section (<italic>fide</italic> <xref ref-type="bibr" rid="R178">Thom 1930</xref>). Members of the section <italic>Ramigena</italic> share the following characters: a slow growth rate on agar media, a monoverticillate branching system with non-vesiculate stipes. Conidia are relatively large (3–4 μm), smooth and ellipsoidal or pyriform (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>). <italic>Penicillium ornatum</italic> is the sole member known in this section with a teleomorph (<xref ref-type="bibr" rid="R182">Udagawa 1968</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>). The ascospores of this species are ornamented with two and sometimes four longitudinal flanges. The ex-type culture of <italic>P. implicatum</italic> in the CBS collection (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=232.38&link_type=cbs">CBS 232.38</ext-link>) is a <italic>Penicillium citrinum</italic>, and therefore this species is not accepted as distinct (<xref ref-type="bibr" rid="R38">Frisvad <italic>et al</italic>. 1990b</xref>, <xref ref-type="bibr" rid="R56">Houbraken <italic>et al</italic>. 2010b</xref>). Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) neotypified <italic>P. implicatum</italic> with <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=184.81&link_type=cbs">CBS 184.81</ext-link> and <xref ref-type="fig" rid="F10">Fig. 10</xref> shows that this strain is closely related to the type of <italic>Penicillium hispanicum</italic> <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=691.77&link_type=cbs">CBS 691.77</ext-link>. This neotypification is not accepted here and <italic>P. implicatum sensu</italic> Pitt is considered as a synonym of <italic>P. hispanicum</italic>. Pitt <italic>et al</italic>. (<xref ref-type="bibr" rid="R130">2000</xref>) accepted <italic>P. dierckxii, P. cyaneum</italic> and <italic>P. sublateritium</italic> as single species in their overview of accepted species in <italic>Penicillium</italic>. This concept is followed here; however, partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F10">Fig. 10</xref>) shows that these three species are very closely related and might represent one species.</p><p id="P219"><list list-type="simple"><list-item><p id="P220"><italic>Penicillium capsulatum</italic> Raper & Fennell, Mycologia 40: 528. 1948.</p></list-item><list-item><p id="P221"><italic>Penicillium cyaneum</italic> (Bainier & Sartory) Biourge, Cellule 33: 102. 1923.</p></list-item><list-item><p id="P222"><italic>Penicillium dierckxii</italic> Biourge, Cellule 33: 313. 1923.</p></list-item><list-item><p id="P223"><italic>Penicillium hispanicum</italic> Ramírez, Martínez & Ferrer, Mycopathol. 66: 77. 1978 (syn. <italic>Penicillium implicatum sensu</italic> Pitt).</p></list-item><list-item><p id="P224"><italic>Penicillium ornatum</italic> Udagawa, Trans. Mycol. Soc. Japan 9: 49.1968.</p></list-item><list-item><p id="P225"><italic>Penicillium ramusculum</italic> Batista & Maia, Anais Soc. Biol. Pernamb. 13: 27. 1955 (syn. <italic>P. brevissimum</italic> Rai & Wadhwani) (this study, <xref ref-type="bibr" rid="R118">Peterson & Horn 2009</xref>).</p></list-item><list-item><p id="P226"><italic>Penicillium sublateritium</italic> Biourge, Cellule 33: 315. 1923.</p></list-item></list></p></sec><sec id="S58"><title>Clade 8: section <italic>Torulomyces</italic> (Delitsch) Stolk & Samson, Adv. Pen. Asp. Syst.: 169. 1985.</title><p id="P227">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P228">Type: <italic>Penicillium lagena</italic> (Delitsch) Stolk & Samson</p><p id="P229">The genus <italic>Torulomyces</italic> is synonymised with <italic>Penicillium</italic> and consequently the majority of the species described in <italic>Torulomyces</italic> are transferred to <italic>Penicillium</italic> (this study). <xref ref-type="fig" rid="F7">Figure 7</xref> shows that <italic>P. lagena</italic> is related to <italic>P. cryptum</italic> and <italic>P. lassenii</italic>. These species have a slow growth rate on the agar media CYA and MEA and form short-stiped monoverticillate or terminal biverticillate conidiophores. Phialides are predominantly singly formed in <italic>P. lagena</italic>, short, 4–7 μm long, with a narrowed base and a swollen middle that tapers abruptly into a narrow neck (<xref ref-type="fig" rid="F6">Fig. 6</xref>).</p><p id="P230"><list list-type="simple"><list-item><p id="P231"><italic>Penicillium cryptum</italic> Gochenaur, Mycotaxon 26: 349. 1986.</p></list-item><list-item><p id="P232"><italic>Penicillium lagena</italic> (Delitsch) Stolk & Samson, Stud. Mycol. 23:100. 1983.</p></list-item><list-item><p id="P233"><italic>Penicillium laeve</italic> (K. Ando & Manoch) Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P234"><italic>Penicillium lassenii</italic> Paden, Mycopathol. Mycol. Appl. 43: 266. 1971.</p></list-item><list-item><p id="P235"><italic>Penicillium ovatum</italic> (K. Ando & Nawawi) Houbraken & Samson, Stud. Mycol. 70: 48. 2011 (this study).</p></list-item><list-item><p id="P236"><italic>Penicillium parviverrucosum</italic> (K. Ando & Pitt) Houbraken & Samson, Stud. Mycol. 70: 48. 2011 (this study).</p></list-item><list-item><p id="P237"><italic>Penicillium porphyreum</italic> Houbraken & Samson, Stud. Mycol. 70: 48. 2011 (this study).</p></list-item></list></p></sec><sec id="S59"><title>Clade 9: section <italic>Fracta</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563129&link_type=mb">MB563129</ext-link>.</title><p id="P238">Sectio in Penicillio subgen. Aspergilloide. Coloniis in agaro tarde crescentibus; ascosporis spinulosis; phialidibus ampullifomibus vel lanceolatis; conidiis ellipsoideis.</p><p id="P239">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P240">Type: <italic>Penicillium ornatum</italic> Udagawa</p><p id="P241"><italic>Penicillium inusitatum</italic> and <italic>P. fractum</italic> belong to section <italic>Fracta</italic> and both are able to form a teleomorph. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) noted that these two species are closely related, differing principally in conidiophore structure. Both species share unusual ascospore morphology for <italic>Penicillium</italic> species: the ascospores are spheroidal without flanges or furrows and ornamented by spines. Furthermore, both species grow slowly on agar media, form ampulliform to lanceolate phialides and ellipsoidal conidia. Phylogenetically, section <italic>Fracta</italic> might be related to section <italic>Torulomyces</italic> (72 % bs, < 0.95 pp). However, ascospores produced by the members of the latter section have two ridges (<italic>P. lagena, P. lassenii, P. cryptum</italic>).</p><p id="P242"><list list-type="simple"><list-item><p id="P243"><italic>Penicillium fractum</italic> Udagawa, Trans. Mycol. Soc. Japan 9: 51. 1968.</p></list-item><list-item><p id="P244"><italic>Penicillium inusitatum</italic> Scott, Mycopathol. Mycol. Appl. 36: 20. 1968.</p></list-item></list></p></sec><sec id="S60"><title>Clade 10: section <italic>Exilicaulis</italic> Pitt, The Genus <italic>Penicillium</italic>: 205. 1980.</title><p id="P245">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P246">Type: <italic>Penicillium restrictum</italic> Gilman & Abbott</p><p id="P247"><list list-type="simple"><list-item><p id="P248">= <italic>Eupenicillium</italic> section <italic>Lapidosa</italic> (Pitt) Stolk & Samson, Stud. Mycol. 23: 55. 1983.</p></list-item></list></p><p id="P249">Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) defined section <italic>Exilicaulis</italic> for monoverticillate species with stipes lacking a terminal vesicular swelling. The phylogenetic delimitation is broader and also several species with an additional branch are included (<italic>e.g. P. raciborski, P. melinii, P. velutinum, P. corylophilum</italic>). This section largely corresponds with group 4 of Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>); the only difference is that Peterson placed <italic>P. turbatum</italic> in this clade, while our data shows that this species belongs to section <italic>Turbata</italic> (group 6 <italic>fide</italic> Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>)). Based on <xref ref-type="fig" rid="F10">Fig. 10</xref> and data of Peterson <italic>et al</italic>. and <xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, the following species are included in section <italic>Exilicaulis</italic>:</p><p id="P250"><list list-type="simple"><list-item><p id="P251"><italic>Penicillium alutaceum</italic> Scott, Mycopathol. Mycol. Appl. 36: 17. 1968.</p></list-item><list-item><p id="P252"><italic>Penicillium atrosanguineum</italic> Dong, Ceská Mycol. 27: 174. 1973.</p></list-item><list-item><p id="P253"><italic>Penicillium burgense</italic> Quintanilla, Avances Nutr. Mejora Anim. Aliment. 30: 176. 1990.</p></list-item><list-item><p id="P254"><italic>Penicillium catenatum</italic> Scott, Mycopathol. Mycol. Appl. 36: 24. 1968.</p></list-item><list-item><p id="P255"><italic>Penicillium chalybeum</italic> Pitt & Hocking, Mycotaxon 22: 204. 1985.</p></list-item><list-item><p id="P256"><italic>Penicillium cinerascens</italic> Biourge, Cellule 33: 308. 1923.</p></list-item><list-item><p id="P257"><italic>Penicillium cinereoatrum</italic> Chalabuda, Bot. Mater. Otd. Sporov. Rast. 6: 167, 1950 (<xref ref-type="bibr" rid="R39">Frisvad <italic>et al</italic>. 1990c</xref>).</p></list-item><list-item><p id="P258"><italic>Penicillium citreonigrum</italic> Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.</p></list-item><list-item><p id="P259"><italic>Penicillium corylophilum</italic> Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.</p></list-item><list-item><p id="P260"><italic>Penicillium decumbens</italic> Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 71. 1910.</p></list-item><list-item><p id="P261"><italic>Penicillium dimorphosporum</italic> Swart, Trans. Br. Mycol. Soc. 55: 310. 1970.</p></list-item><list-item><p id="P262"><italic>Penicillium dravuni</italic> Janso, Mycologia 97: 445. 2005.</p></list-item><list-item><p id="P263"><italic>Penicillium erubescens</italic> Scott, Mycopathol. Mycol. Appl. 36: 14. 1968.</p></list-item><list-item><p id="P264"><italic>Penicillium fagi</italic> Ramírez & Martínez, Mycopathol. 63: 57. 1978.</p></list-item><list-item><p id="P265"><italic>Penicillium flavidostipitatum</italic> Ramírez & González, Mycopathol. 88: 3. 1984 (preliminary sequencing results show that this species is closely related to <italic>P. namyslowskii</italic>).</p></list-item><list-item><p id="P266"><italic>Penicillium guttulosum</italic> Gilman & Abbott, Iowa State Coll. J. Sci. 1: 298. 1927 (<xref ref-type="bibr" rid="R120">Peterson <italic>et al</italic>. 2011</xref>).</p></list-item><list-item><p id="P267"><italic>Penicillium heteromorphum</italic> Kong & Qi, Mycosystema 1: 107. 1988.</p></list-item><list-item><p id="P268"><italic>Penicillium katangense</italic> Stolk, Ant. van Leeuwenhoek 34: 42. 1968.</p></list-item><list-item><p id="P269"><italic>Penicillium lapidosum</italic> Raper & Fennell, Mycologia 40: 524. 1948.</p></list-item><list-item><p id="P270"><italic>Penicillium maclennaniae</italic> Yip, Trans. Br. Mycol. Soc. 77: 202. 1981.</p></list-item><list-item><p id="P271"><italic>Penicillium melinii</italic> Thom, Penicillia: 273. 1930.</p></list-item><list-item><p id="P272"><italic>Penicillium menonorum</italic> Peterson, IMA Fungus 2: 122. 2011.</p></list-item><list-item><p id="P273"><italic>Penicillium meridianum</italic> Scott, Mycopathol. Mycol. Appl. 36: 12. 1968.</p></list-item><list-item><p id="P274"><italic>Penicillium namyslowskii</italic> Zaleski, Bull. Int. Aead. Polonc. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 479. 1927.</p></list-item><list-item><p id="P275"><italic>Penicillium nepalense</italic> Takada & Udagawa, Trans. Mycol. Soc. Japan 24: 146. 1983.</p></list-item><list-item><p id="P276"><italic>Penicillium parvum</italic> Raper & Fennell, Mycologia 40: 508. 1948 (this study).</p></list-item><list-item><p id="P277"><italic>Penicillium philippinense</italic> Udagawa & Y. Horie, J. Jap. Bot. 47: 341. 1972.</p></list-item><list-item><p id="P278"><italic>Penicillium pimiteouiense</italic> Peterson, Mycologia 91: 271. 1999.</p></list-item><list-item><p id="P279"><italic>Penicillium raciborskii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 454. 1927.</p></list-item><list-item><p id="P280"><italic>Penicillium restrictum</italic> Gilman & Abbott, Iowa State Coll. J. Sci. 1: 297. 1927.</p></list-item><list-item><p id="P281"><italic>Penicillium rubefaciens</italic> Quintanilla, Mycopathol. 80: 73. 1982.</p></list-item><list-item><p id="P282"><italic>Penicillium rubidurum</italic> Udagawa & Horie, Trans. Mycol. Soc. Japan 14: 381. 1973.</p></list-item><list-item><p id="P283"><italic>Penicillium smithii</italic> Quintanilla, Avances Nutr. Mejora Anim. Aliment. 23: 340. 1982 (syn. <italic>P. corynephorum, P. sabulosum</italic>).</p></list-item><list-item><p id="P284"><italic>Penicillium striatisporum</italic> Stolk, Ant. van Leeuwenhoek 35: 268. 1969.</p></list-item><list-item><p id="P285"><italic>Penicillium terrenum</italic> Scott, Mycopathol. Mycol. Appl. 36: 1. 1968.</p></list-item><list-item><p id="P286"><italic>Penicillium toxicarium</italic> Miyake, Rep. Res. Inst. Rice Improvement 1: 1. 1940 (<italic>nom. inval</italic>., Art. 36) (Serra <italic>et al</italic>. 2008).</p></list-item><list-item><p id="P287"><italic>Penicillium velutinum</italic> van Beyma, Zentralbl. Bakteriol., 2. Abt., 91: 353. 1935.</p></list-item><list-item><p id="P288"><italic>Penicillium vinaceum</italic> Gilman & Abbott, Iowa State Coll. J. Sci. 1: 299. 1927.</p></list-item></list></p></sec><sec id="S61"><title>Clade 11: Section <italic>Lanata-divaricata</italic> Thom, The Penicillia: 328. 1930.</title><p id="P289"><list list-type="simple"><list-item><p id="P290">= section <italic>Funiculosa</italic> Thom, The Penicillia: 358. 1930.</p></list-item><list-item><p id="P291">= section <italic>Divaricatum</italic> Pitt, The Genus <italic>Penicillium</italic>: 238. 1980.</p></list-item><list-item><p id="P292">= section <italic>Furcatum</italic> Pitt, The Genus <italic>Penicillium</italic>: 272. 1980.</p></list-item><list-item><p id="P293">= <italic>Eupenicillium</italic> section <italic>Javanica</italic> (Pitt) Stolk & Samson, Stud. Mycol. 23: 55. 1983.</p></list-item></list></p><p id="P294">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P295">Type: <italic>Penicillium janthinellum</italic> Biourge</p><p id="P296">Most of the species, but not all, of section <italic>Lanata-divaricata</italic> grow rapidly and form broadly spreading colonies. The majority of the species belonging to this section are strongly divaricate and the metulae are born terminally, subterminally and in intercalary positions, and in the latter case intergrading with monoverticillate conidiophores. Furthermore, the terminal cluster often consists of a prolongation of the main axis. Species belonging to section <italic>Lanata-divaricata</italic> are mainly soil inhabitants, but may also occur on leaf litter and vegetable remains in the later stage of decomposition (<xref ref-type="bibr" rid="R135">Raper & Thom 1949</xref>, <xref ref-type="bibr" rid="R59">Houbraken <italic>et al</italic>. 2011c</xref>). Many species of this section are unusually tolerant for heavy metals and some species have been proposed as efficient biosorbent agents in the bioleaching of zinc oxide, copper, lead and nickel (<xref ref-type="bibr" rid="R20">Burgstaller <italic>et al</italic>. 1992</xref>, <xref ref-type="bibr" rid="R187">Valix <italic>et al</italic>. 2001</xref>, <xref ref-type="bibr" rid="R79">Li <italic>et al</italic>. 2008</xref>).</p><p id="P297">Section <italic>Funiculosa</italic> is placed in synonymy with this section. Thom (<xref ref-type="bibr" rid="R178">1930</xref>) already noted that species belonging section <italic>Funiculosa</italic> have affinity with members of section <italic>Lanata-typica</italic> and that separation is hard to define. This observation is supported by our data: many species mentioned in Thom's section <italic>Funiculosa</italic> belong to section <italic>Lanata-divaricata</italic>. Raper & Thom's (<xref ref-type="bibr" rid="R135">1949</xref>) subsection <italic>Divaricata</italic> largely corresponds with our section <italic>Lanata-divaricata</italic>. They noted that members of their subsection have a definite relationship to <italic>Penicillium javanicum.</italic> Stolk & Samson (<xref ref-type="bibr" rid="R166">1983</xref>) also discussed this relationship and they placed 26 species in synonymy with <italic>Eupenicillium javanicum</italic> and <italic>P. simplicissimum</italic>. Recently, a phylogenetic study showed that many of these synonyms should be treated as separate species (<xref ref-type="bibr" rid="R112">Peterson 2000a</xref>, <xref ref-type="bibr" rid="R59">Houbraken <italic>et al</italic>. 2011c</xref>). This section largely corresponds with Peterson's (<xref ref-type="bibr" rid="R112">2000a</xref>) group 5 and the list provided here for this section is mainly based on this data supplemented with data of Houbraken <italic>et al</italic>. (<xref ref-type="bibr" rid="R59">2011c</xref>). <italic>Penicillium cluniae, P. griseopurpureum</italic> and <italic>P. glaucoroseum</italic> were not included in these studies, though unpublished data shows that these three species also belong to this section.</p><p id="P298">The typification of <italic>P. brefeldianum, P. javanicum, P. levitum</italic> and <italic>P. ehrlichii</italic> warrants further attention. Dodge (<xref ref-type="bibr" rid="R26">1933</xref>) described <italic>P. brefeldianum</italic> as a holomorphic species. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) did not accept teleomorph species in <italic>Penicillium</italic> and a neotype (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=233.81&link_type=cbs">CBS 233.81</ext-link> = FRR 71 = IMI 216895) was selected because the original type culture of <italic>P. brefeldianum</italic> distributed by Dodge no longer produced cleistothecia. Subsequently, Dodge's strain (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.81&link_type=cbs">CBS 235.81</ext-link> = FRR 710 = IMI 216896 = NRRL 710) was used for the description of the anamorph of <italic>Eupenicillium brefeldianum</italic> (as <italic>Penicillium dodgei</italic>). Teleomorphs are allowed in <italic>Penicillium</italic> and therefore Dodge's <italic>P. brefeldianum</italic> is re-instated. Furthermore, <xref ref-type="fig" rid="F11">Fig. 11</xref> shows that Dodge's type strain (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=235.81&link_type=cbs">CBS 235.81</ext-link>) differs from Pitt's neotype (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=233.81&link_type=cbs">CBS 233.81</ext-link>) and this neotype is similar to the type of <italic>P. caperatum</italic> (<ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=443.75&link_type=cbs">CBS 443.75</ext-link><sup>T</sup>). <italic>Penicillium levitum, P. javanicum</italic> and <italic>P. ehrlichii</italic> were described including a teleomorph. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) introduced the new species names <italic>P. rasile, P. indonesiae</italic> and <italic>P. klebahnii</italic> respectively, for the anamorphs of <italic>P. levitum, P. javanicum</italic> and <italic>P. ehrlichii</italic>. These names are not used here for the same the reason as mentioned under <italic>P. brefeldianum</italic>.</p><p id="P299"><list list-type="simple"><list-item><p id="P300"><italic>Penicillium abidjanum</italic> Stolk, Ant. van Leeuwenhoek 34: 49. 1968.</p></list-item><list-item><p id="P301"><italic>Penicillium araracuarense</italic> Houbraken, C. López-Q, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 61: 1469. 2011.</p></list-item><list-item><p id="P302"><italic>Penicillium brasilianum</italic> Batista, Anais Soc. Biol. Pernambuco 15: 162. 1957.</p></list-item><list-item><p id="P303"><italic>Penicillium brefeldianum</italic> Dodge, Mycologia 25: 92. 1933 (syn. <italic>P. dodgei</italic>).</p></list-item><list-item><p id="P304"><italic>Penicillium caperatum</italic> Udagawa & Horie, Trans. Mycol. Soc. Japan 14: 371. 1973 (syn. <italic>E. brefeldianum sensu</italic> Pitt).</p></list-item><list-item><p id="P305"><italic>Penicillium cluniae</italic> Quintanilla, Avances Nutr. Mejora Anim. Aliment. 30: 174. 1990. (unpubl. data)</p></list-item><list-item><p id="P306"><italic>Penicillium coeruleum</italic> Sopp <italic>apud</italic> Biourge, Cellule 33: 102. 1923.</p></list-item><list-item><p id="P307"><italic>Penicillium cremeogriseum</italic> Chalabuda, Bot. Mater. Otd. Sporov. Rast. 6: 168. 1950.</p></list-item><list-item><p id="P308"><italic>Penicillium daleae</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 495. 1927.</p></list-item><list-item><p id="P309"><italic>Penicillium ehrlichii</italic> Klebahn, Ber. Deutsch. Bot. Ges. 48: 374. 1930.</p></list-item><list-item><p id="P310"><italic>Penicillium elleniae</italic> Houbraken, C. López-Q, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 61: 1470. 2011.</p></list-item><list-item><p id="P311"><italic>Penicillium glaucoroseum</italic> Demelius, Verh. Zool.-Bot. Ges. Wien 72: 72. 1923. (unpubl. data)</p></list-item><list-item><p id="P312"><italic>Penicillium griseopurpureum</italic> Smith, Trans. Br. Mycol. Soc. 48: 275. 1965 (unpubl. data).</p></list-item><list-item><p id="P313"><italic>Penicillium janthinellum</italic> Biourge, Cellule 33: 258. 1923.</p></list-item><list-item><p id="P314"><italic>Penicillium javanicum</italic> van Beyma, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect., 26: 17. 1929 (syn. <italic>P. oligosporum, P. indonesiae</italic>).</p></list-item><list-item><p id="P315"><italic>Penicillium levitum</italic> Raper & Fennell, Mycologia 40: 511. 1948 (syn. <italic>P. rasile</italic>).</p></list-item><list-item><p id="P316"><italic>Penicillium limosum</italic> Ueda, Mycoscience 36: 451. 1995.</p></list-item><list-item><p id="P317"><italic>Penicillium lineolatum</italic> Udagawa & Horie, Mycotaxon 5: 493. 1977.</p></list-item><list-item><p id="P318"><italic>Penicillium ludwigii</italic> Udagawa, Trans. Mycol. Soc. Japan 10: 2. 1969.</p></list-item><list-item><p id="P319"><italic>Penicillium mariaecrucis</italic> Quintanilla, Avances Nutr. Mejora Anim. Aliment. 23: 334. 1982.</p></list-item><list-item><p id="P320"><italic>Penicillium meloforme</italic> Udagawa & Horie, Trans. Mycol. Soc. Japan 14: 376. 1973.</p></list-item><list-item><p id="P321"><italic>Penicillium ochrochloron</italic> Biourge, Cellule 33: 269. 1923.</p></list-item><list-item><p id="P322"><italic>Penicillium onobense</italic> Ramírez & Martínez, Mycopathol. 74: 44. 1981.</p></list-item><list-item><p id="P323"><italic>Penicillium oxalicum</italic> Currie & Thom, J. Biol. Chem. 22: 289. 1915.</p></list-item><list-item><p id="P324"><italic>Penicillium paraherquei</italic> Abe ex Smith, Trans. Br. Mycol. Soc. 46: 335. 1963.</p></list-item><list-item><p id="P325"><italic>Penicillium penarojense</italic> Houbraken, C. López-Q, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 61: 1471. 2011.</p></list-item><list-item><p id="P326"><italic>Penicillium piscarium</italic> Westling, Ark. Bot. 11: 86. 1911.</p></list-item><list-item><p id="P327"><italic>Penicillium pulvillorum</italic> Turfitt, Trans. Br. Mycol. Soc. 23: 186. 1939 (Syn. <italic>P. ciegleri</italic>).</p></list-item><list-item><p id="P328"><italic>Penicillium raperi</italic> Smith, Trans. Br. Mycol. Soc. 40: 486. 1957.</p></list-item><list-item><p id="P329"><italic>Penicillium reticulisporum</italic> Udagawa, Trans. Mycol. Soc. Japan 9: 52. 1968. (syn. <italic>P. arvense</italic>).</p></list-item><list-item><p id="P330"><italic>Penicillium rolfsii</italic> Thom, Penicillia: 489. 1930.</p></list-item><list-item><p id="P331"><italic>Penicillium simplicissimum</italic> (Oudemans) Thom, Penicillia: 335. 1930.</p></list-item><list-item><p id="P332"><italic>Penicillium skrjabinii</italic> Schmotina & Golovleva, Mikol. Fitopatol. 8: 530. 1974.</p></list-item><list-item><p id="P333"><italic>Penicillium svalbardense</italic> Frisvad, Sonjak & Gunde-Cimerman, Ant. van Leeuwenhoek 92: 48. 2007.</p></list-item><list-item><p id="P334"><italic>Penicillium vanderhammenii</italic> Houbraken, C. López-Q, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 61: 1473. 2011.</p></list-item><list-item><p id="P335"><italic>Penicillium vasconiae</italic> Ramírez & Martínez, Mycopathol. 72: 189. 1980.</p></list-item><list-item><p id="P336"><italic>Penicillium wotroi</italic> Houbraken, C. López-Q, Frisvad & Samson, Int. J. Syst. Evol. Microbiol. 61: 1474. 2011.</p></list-item><list-item><p id="P337"><italic>Penicillium zonatum</italic> Hodges & Perry, Mycologia 65: 697. 1973.</p></list-item></list></p></sec><sec id="S62"><title>Clade 12: section <italic>Stolkia</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563130&link_type=mb">MB563130</ext-link>.</title><p id="P338">Sectio in Penicillio subgen. Aspergilloide. Conidiophoris pigmentatis, metulis subapicalibus sympodialiter proliferantibus; phialidibus nullis.</p><p id="P339">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P340">Type: <italic>Penicillium stolkiae</italic> Scott</p><p id="P341">Brown conidiophores occur in two phylogenetic unrelated sections of <italic>Penicillium s. str</italic>. One includes species belonging to section <italic>Thysanophora</italic> (previously assigned to the genus <italic>Thysanophora</italic>) (<xref ref-type="bibr" rid="R62">Iwamoto <italic>et al</italic>. 2002</xref>, <xref ref-type="bibr" rid="R122">Peterson & Sigler 2002</xref>) and the second lineage is centered around <italic>P. stolkiae</italic>, another species with conidiophores that also may be hyaline to definitely brown (<xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>). Peterson & Sigler (<xref ref-type="bibr" rid="R122">2002</xref>) described four species with darkly melanised conidiophores, which are all closely related to <italic>P. stolkiae</italic>, namely <italic>P. subarticum, P. canariense, P. pullum</italic> and <italic>P. boreae</italic>. None of these species demonstrate the sympodial proliferation of subapical metulae and phialides present in section <italic>Thysanophora</italic>. The following species are placed in section <italic>Stolkia</italic> based on the data presented in <xref ref-type="fig" rid="F11">Fig. 11</xref> and of Peterson & Sigler (<xref ref-type="bibr" rid="R122">2002</xref>).</p><p id="P342"><list list-type="simple"><list-item><p id="P343"><italic>Penicillium boreae</italic> Peterson & Sigler, Mycol. Res. 106: 1112. 2002.</p></list-item><list-item><p id="P344"><italic>Penicillium canariense</italic> Peterson & Sigler, Mycol. Res. 106: 1113. 2002.</p></list-item><list-item><p id="P345"><italic>Penicillium donkii</italic> Stolk, Persoonia 7: 333. 1973.</p></list-item><list-item><p id="P346"><italic>Penicillium pullum</italic> Peterson & Sigler, Mycol. Res. 106: 1115. 2002.</p></list-item><list-item><p id="P347"><italic>Penicillium stolkiae</italic> Scott, Mycopathol. Mycol. Appl. 36: 8. 1968.</p></list-item><list-item><p id="P348"><italic>Penicillium subarcticum</italic> Peterson & Sigler, Mycol. Res. 106: 1116. 2002.</p></list-item></list></p></sec><sec id="S63"><title>Clade 13: section <italic>Gracilenta</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563131&link_type=mb">MB563131</ext-link>.</title><p id="P349">Sectio in Penicillio subgen. Aspergilloide. Coloniis 37 °C haud crescentibus, reverso olivaceo-brunneo vel brunneo, conidiis saepe late ellipsoideis vel ellipsoideis.</p><p id="P350">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P351">Type: <italic>Penicillium gracilentum</italic> Udagawa & Horie</p><p id="P352">Four species are placed in section <italic>Gracilenta</italic>. Comparison of the phenotypic characters did not reveal many significant similarities among these species. All species did not grow at 37 °C and have an olive-brown to brown reverse on agar media. With exception of <italic>P. macrosclerotiorum</italic>, all species produced broadly ellipsoidal to ellipsoidal conidia (<xref ref-type="bibr" rid="R1">Abe 1956</xref>, <xref ref-type="bibr" rid="R183">Udagawa & Horie 1973</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R175">Takada & Udagawa 1983</xref>, <xref ref-type="bibr" rid="R194">Wang <italic>et al</italic>. 2007</xref>). The taxonomy and phylogeny of these species is not well studied and future research might reveal more shared characters.</p><p id="P353"><list list-type="simple"><list-item><p id="P354"><italic>Penicillium angustiporcatum</italic> Takada & Udagawa, Trans. Mycol. Soc. Japan 24: 143. 1983.</p></list-item><list-item><p id="P355"><italic>Penicillium estinogenum</italic> Komatsu & Abe ex Smith, Trans. Br. Mycol. Soc. 46: 335. 1963.</p></list-item><list-item><p id="P356"><italic>Penicillium macrosclerotiorum</italic> Wang, Zhang & Zhuang, Mycol. Res. 111: 1244. 2007.</p></list-item><list-item><p id="P357"><italic>Penicillium gracilentum</italic> Udagawa & Horie, Trans. Mycol. Soc. Japan 14: 373. 1973.</p></list-item></list></p></sec><sec id="S64"><title>Clade 14: section <italic>Citrina</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563132&link_type=mb">MB563132</ext-link>.</title><p id="P358">Sectio in Penicillio subgen. Aspergilloide. Formatione conidiophorum symmetricorum biverticillatorum.</p><p id="P359">In: <italic>Penicillium</italic> subgenus <italic>Aspergilloides</italic></p><p id="P360">Type: <italic>Penicillium citrinum</italic> Thom</p><p id="P361">Species of section <italic>Citrina</italic> are commonly occurring in soil and the majority of the species form symmetrical biverticillate conidiophores. This section corresponds with group 1 of Peterson (<xref ref-type="bibr" rid="R112">2000a</xref>). The taxonomy of section <italic>Citrina</italic> is recently revised by Houbraken <italic>et al</italic>. (<xref ref-type="bibr" rid="R56">2010b</xref>, <xref ref-type="bibr" rid="R58">2011b</xref>) and based on this data and <xref ref-type="fig" rid="F12">Fig. 12</xref>, the following species are placed in section <italic>Citrina</italic>:</p><p id="P362"><list list-type="simple"><list-item><p id="P363"><italic>Penicillium anatolicum</italic> Stolk, Ant. van Leeuwenhoek 34: 46. 1968.</p></list-item><list-item><p id="P364"><italic>Penicillium argentinense</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 78. 2011.</p></list-item><list-item><p id="P365"><italic>Penicillium atrofulvum</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 80. 2011.</p></list-item><list-item><p id="P366"><italic>Penicillium aurantiacobrunneum</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 80. 2011.</p></list-item><list-item><p id="P367"><italic>Penicillium cairnsense</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 83. 2011.</p></list-item><list-item><p id="P368"><italic>Penicillium christenseniae</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 85. 2011.</p></list-item><list-item><p id="P369"><italic>Penicillium chrzaszczii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 464. 1927.</p></list-item><list-item><p id="P370"><italic>Penicillium citrinum</italic> Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 61. 1910.</p></list-item><list-item><p id="P371"><italic>Penicillium copticola</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 88. 2011.</p></list-item><list-item><p id="P372"><italic>Penicillium cosmopolitanum</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 91. 2011.</p></list-item><list-item><p id="P373"><italic>Penicillium decaturense</italic> Peterson, Bayer & Wicklow, Mycologia 96: 1290. 2004.</p></list-item><list-item><p id="P374"><italic>Penicillium euglaucum</italic> van Beyma, Ant. van Leeuwenhoek 6: 269. 1940.</p></list-item><list-item><p id="P375"><italic>Penicillium galliacum</italic> Ramírez, Martínez & Berenguer, Mycopathol. 72: 30. 1980.</p></list-item><list-item><p id="P376"><italic>Penicillium godlewskii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 466. 1927.</p></list-item><list-item><p id="P377"><italic>Penicillium gorlenkoanum</italic> Baghdadi, Nov. Sist. Niz. Rast. 5: 97. 1968.</p></list-item><list-item><p id="P378"><italic>Penicillium hetheringtonii</italic> Houbraken, Frisvad & Samson, Fung. Div. 44: 125. 2010.</p></list-item><list-item><p id="P379"><italic>Penicillium manginii</italic> Duché & Heim, Trav. Cryptog. Louis L. Mangin: 450. 1931 (syn. <italic>P. pedemontanum</italic>, <xref ref-type="bibr" rid="R58">Houbraken <italic>et al</italic>. 2011b</xref>).</p></list-item><list-item><p id="P380"><italic>Penicillium miczynskii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 482. 1927.</p></list-item><list-item><p id="P381"><italic>Penicillium neomiczynskii</italic> Cole, Houbraken, Frisvad & Samson, Stud. Mycol. 70: 105. 2011.</p></list-item><list-item><p id="P382"><italic>Penicillium nothofagi</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 105. 2011.</p></list-item><list-item><p id="P383"><italic>Penicillium pancosmium</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 108. 2011.</p></list-item><list-item><p id="P384"><italic>Penicillium pasqualense</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 108. 2011.</p></list-item><list-item><p id="P385"><italic>Penicillium paxilli</italic> Bainier, Bull. Soc. Mycol. France 23: 95. 1907.</p></list-item><list-item><p id="P386"><italic>Penicillium quebecense</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 111. 2011.</p></list-item><list-item><p id="P387"><italic>Penicillium raphiae</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 114. 2011.</p></list-item><list-item><p id="P388"><italic>Penicillium roseopurpureum</italic> Dierckx, Ann. Soc. Sci. Bruxelles 25: 86. 1901.</p></list-item><list-item><p id="P389"><italic>Penicillium sanguifluum</italic> (Sopp) Biourge, La Cellule 33: 105. 1923. <italic>Penicillium shearii</italic> Stolk & Scott, Persoonia 4: 396. 1967.</p></list-item><list-item><p id="P390"><italic>Penicillium sizovae</italic> Baghdadi, Novosti Sist. Nizs. Rast. 1968: 103. 1968.</p></list-item><list-item><p id="P391"><italic>Penicillium steckii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 469. 1927.</p></list-item><list-item><p id="P392"><italic>Penicillium sumatrense</italic> Szilvinyi, Archiv. Hydrobiol. 14, Suppl. 6: 535. 1936.</p></list-item><list-item><p id="P393"><italic>Penicillium terrigenum</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 125. 2011.</p></list-item><list-item><p id="P394"><italic>Penicillium tropicoides</italic> Houbraken, Frisvad & Samson, Fung. Div. 44: 127. 2010.</p></list-item><list-item><p id="P395"><italic>Penicillium tropicum</italic> Houbraken, Frisvad & Samson, Fung. Div. 44: 129. 2010.</p></list-item><list-item><p id="P396"><italic>Penicillium ubiquetum</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 127. 2011.</p></list-item><list-item><p id="P397"><italic>Penicillium vancouverense</italic> Houbraken, Frisvad & Samson, Stud. Mycol. 70: 131. 2011.</p></list-item><list-item><p id="P398"><italic>Penicillium waksmanii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 468. 1927.</p></list-item><list-item><p id="P399"><italic>Penicillium wellingtonense</italic> Cole, Houbraken, Frisvad & Samson, Stud. Mycol. 70: 133. 2011.</p></list-item><list-item><p id="P400"><italic>Penicillium westlingii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 473. 1927.</p></list-item></list></p></sec><sec id="S65"><title>Clade 15: Section <italic>Fasciculata</italic> Thom, The Penicillia: 374. 1930.</title><p id="P401"><list list-type="simple"><list-item><p id="P402">= Section <italic>Lanata</italic>-<italic>typica</italic> Thom, The Penicillia: 305. 1930.</p></list-item><list-item><p id="P403">= Section <italic>Viridicata</italic> Frisvad & Samson, Stud. Mycol. 49: 27. 2004.</p></list-item></list></p><p id="P404">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P405">Type: <italic>Penicillium hirsutum</italic> Dierckx</p><p id="P406">Sections <italic>Lanata-typica</italic> and <italic>Viridicata</italic> are placed in synonymy with section <italic>Fasciculata. Lanata-typica</italic> was erected for species with vegetative aerial mycelium consisting of lanose, cottony or floccose colonies and only a small portion of the species currently present this section produce such structures (<italic>P. camemberti, P. commune, P. caseifulvum</italic>). Most species of section <italic>Fasciculata</italic> have a granulose or fasciculate colony texture and therefore the name <italic>Fasciculata</italic> is given priority to <italic>Lanata-typica</italic>. The current definition of <italic>Fasciculata</italic> is similar to that of <italic>Viridicata</italic> (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>). All species grow rather quickly, except species in series <italic>Verrucosa</italic>, which grow slowly. Most species this section have globose conidia and rough-walled conidiophore stipes. All species are psychrotolerant and grow well at low water activities (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>). Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) accommodated 28 species in section <italic>Viridicata</italic> (= <italic>Fasciculata</italic>). We excluded <italic>P. atramentosum</italic> from this section and placed this species in section <italic>Paradoxa</italic>. This species was placed in section <italic>Fasciculata</italic> based on its ability to grow on creatine as sole nitrogen source and its occurrence on cheese. However, Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) also noted that its ability to grow at very high pH values and the formation of smooth-walled stipes sets it apart from section <italic>Fasciculata. Penicillium osmophilum</italic> is tentatively accommodated in section <italic>Viridicata</italic>. <xref ref-type="fig" rid="F13">Figure 13</xref> shows that this species is most closely related to this section, but bootstrap support is lacking.</p><p id="P407"><list list-type="simple"><list-item><p id="P408"><italic>Penicillium albocoremium</italic> (Frisvad) Frisvad, Int. Mod. Tax. Meth. Pen. Asp. Clas.: 275. 2000.</p></list-item><list-item><p id="P409"><italic>Penicillium allii</italic> Vincent & Pitt, Mycologia 81: 300. 1989.</p></list-item><list-item><p id="P410"><italic>Penicillium aurantiogriseum</italic> Dierckx, Ann. Soc. Scient. Brux. 25: 88. 1901.</p></list-item><list-item><p id="P411"><italic>Penicillium camemberti</italic> Thom, Bull. Bur. Anim. Ind. USDA 82: 33. 1906.</p></list-item><list-item><p id="P412"><italic>Penicillium caseifulvum</italic> Lund, Filt. & Frisvad, J. Food Mycol. 1: 97. 1998.</p></list-item><list-item><p id="P413"><italic>Penicillium cavernicola</italic> Frisvad & Samson, Stud. Mycol. 49: 31. 2004.</p></list-item><list-item><p id="P414"><italic>Penicillium commune</italic> Thom, Bull. Bur. Anim. Ind. USDA 118: 56. 1910.</p></list-item><list-item><p id="P415"><italic>Penicillium crustosum</italic> Thom, Penicillia: 399. 1930.</p></list-item><list-item><p id="P416"><italic>Penicillium cyclopium</italic> Westling, Ark. Bot. 11: 90. 1911.</p></list-item><list-item><p id="P417"><italic>Penicillium discolor</italic> Frisvad & Samson, Ant. Van Leeuwenhoek, 72: 120. 1997.</p></list-item><list-item><p id="P418"><italic>Penicillium echinulatum</italic> Fassatiová, Acta Univ. Carol. Biol. 12: 326. 1977.</p></list-item><list-item><p id="P419"><italic>Penicillium freii</italic> Frisvad & Samson, Stud. Mycol. 49: 28. 2004.</p></list-item><list-item><p id="P420"><italic>Penicillium hirsutum</italic> Dierckx, Ann. Soc. Scient. Brux. 25: 89. 1901.</p></list-item><list-item><p id="P421"><italic>Penicillium hordei</italic> Stolk, Ant. van Leeuwenhoek 35: 270. 1969.</p></list-item><list-item><p id="P422"><italic>Penicillium melanoconidium</italic> (Frisvad) Frisvad & Samson, Stud. Mycol. 49: 28. 2004.</p></list-item><list-item><p id="P423"><italic>Penicillium neoechinulatum</italic> (Frisvad, Filt. & Wicklow) Frisvad & Samson, Stud. Mycol. 49: 28. 2004.</p></list-item><list-item><p id="P424"><italic>Penicillium nordicum</italic> Dragoni & Cantoni ex Ramírez, Adv. Pen. Asp. Syst.: 139. 1985.</p></list-item><list-item><p id="P425"><italic>Penicillium osmophilum</italic> Stolk & Veenbaas-Rijks, Ant. van Leeuwenhoek 40: 1. 1974.</p></list-item><list-item><p id="P426"><italic>Penicillium palitans</italic> Westling, Ark. Bot. 11: 83. 1911.</p></list-item><list-item><p id="P427"><italic>Penicillium polonicum</italic> Zaleski, Bull. Int. Acad. Pol. Sci. Lett., Sér. B 1927: 445. 1927.</p></list-item><list-item><p id="P428"><italic>Penicillium radicicola</italic> Overy & Frisvad, Syst. Appl. Microbiol.: 633. 2003.</p></list-item><list-item><p id="P429"><italic>Penicillium solitum</italic> Westling, Ark. Bot. 11: 65. 1911.</p></list-item><list-item><p id="P430"><italic>Penicillium thymicola</italic> Frisvad & Samson, Stud. Mycol. 49: 29. 2004.</p></list-item><list-item><p id="P431"><italic>Penicillium tricolor</italic> Frisvad, Seifert, Samson & Mills, Can. J. Bot. 72: 937. 1994.</p></list-item><list-item><p id="P432"><italic>Penicillium tulipae</italic> Overy & Frisvad, Syst. Appl. Microbiol. 634. 2003.</p></list-item><list-item><p id="P433"><italic>Penicillium venetum</italic> (Frisvad) Frisvad, Int. Mod. Tax. Meth. Pen. Asp. Clas.: 275. 2000.</p></list-item><list-item><p id="P434"><italic>Penicillium verrucosum</italic> Dierckx, Ann. Soc. Scient. Brux. 25: 88. 1901.</p></list-item><list-item><p id="P435"><italic>Penicillium viridicatum</italic> Westling, Ark. Bot. 11: 88. 1911.</p></list-item></list></p></sec><sec id="S66"><title>Clade 16: Section <italic>Digitata</italic> (as “<italic>Digitatum</italic>”) Frisvad & Samson, Stud. Mycol. 49: 26. 2004.</title><p id="P436">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P437">Type: <italic>Penicillium digitatum</italic> (Pers.:Fr.) Sacc.</p><p id="P438">Section <italic>Digitata</italic> is represented by one species, <italic>P. digitatum</italic>. This species is unique in its combination of features. Conidiophore and conidial structures are irregular and exceptionally large for <italic>Penicillium</italic>, usually biverticillate rather than terverticillate and the conidia are olive-green. The conidia are large and ellipsoidal to cylindrical (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>). Partial β-tubulin (<xref ref-type="bibr" rid="R147">Samson <italic>et al</italic>. 2004</xref>) and <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) shows that this section is situated in subgenus <italic>Penicillium</italic>. Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) is followed here and this section is retained for <italic>P. digitatum</italic>.</p><p id="P439"><italic>Penicillium digitatum</italic> (Pers.:Fr.) Sacc., Fung. Ital.: 894. 1881.</p></sec><sec id="S67"><title>Clade 17: Section <italic>Penicillium</italic></title><p id="P440"><list list-type="simple"><list-item><p id="P441">= <italic>Bulliardium</italic> Biourge, La Cellule 33: 107. 1923 (= <italic>Asymetrica</italic>).</p></list-item></list></p><p id="P442">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P443">Type: <italic>Penicillium expansum</italic> Link</p><p id="P444">Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) are followed here in their delimitation of section <italic>Penicillium</italic>. The recently described species <italic>P. brevistipitatum</italic> is added to this list, because it is closely related to <italic>P. coprophilum</italic> (<xref ref-type="fig" rid="F13">Fig. 13</xref>). The analysis of our partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) indicate sthat this section is polyphyletic. In contrast, partial β-tubulin data (<xref ref-type="bibr" rid="R147">Samson <italic>et al</italic>. 2004</xref>) showed that members of this section are on a single branch with 100 % bootstrap support. Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) are followed and the following species are accommodated in section <italic>Penicillium</italic>:</p><p id="P445"><list list-type="simple"><list-item><p id="P446"><italic>Penicillium brevistipitatum</italic> Wang & Zhuang, Mycotaxon 93: 234. 2005.</p></list-item><list-item><p id="P447"><italic>Penicillium clavigerum</italic> Demelius, Verh. Zool.-Bot. Ges. Wien 72: 74. 1922.</p></list-item><list-item><p id="P448"><italic>Penicillium concentricum</italic> Samson, Stolk & Hadlok, Stud. Mycol. 11: 17. 1976.</p></list-item><list-item><p id="P449"><italic>Penicillium coprobium</italic> Frisvad, Mycologia 81: 853. 1989.</p></list-item><list-item><p id="P450"><italic>Penicillium coprophilum</italic> (Berk. & Curt.) Seifert & Samson, Adv.Pen. Asp. Syst.: 145. 1985.</p></list-item><list-item><p id="P451"><italic>Penicillium dipodomyicola</italic> (Frisvad, Filt. & Wicklow) Frisvad, Int. Mod. Meth. Pen. Asp. Clas.: 275. 2000.</p></list-item><list-item><p id="P452"><italic>Penicillium expansum</italic> Link, Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 16. 1809.</p></list-item><list-item><p id="P453"><italic>Penicillium formosanum</italic> Hsieh, Su & Tzean, Trans. Mycol. Soc. R.O.C. 2: 159. 1987.</p></list-item><list-item><p id="P454"><italic>Penicillium gladioli</italic> McCulloch & Thom, Science, N.Y. 67: 217. 1928.</p></list-item><list-item><p id="P455"><italic>Penicillium glandicola</italic> (Oud.) Seifert & Samson, Adv. Pen. Asp. Syst.: 147. 1985.</p></list-item><list-item><p id="P456"><italic>Penicillium griseofulvum</italic> Dierckx, Ann. Soc. Scient. Brux. 25: 88. 1901.</p></list-item><list-item><p id="P457"><italic>Penicillium italicum</italic> Wehmer, Hedwigia 33: 211. 1894.</p></list-item><list-item><p id="P458"><italic>Penicillium marinum</italic> Frisvad & Samson, Stud. Mycol. 49: 20. 2004.</p></list-item><list-item><p id="P459"><italic>Penicillium sclerotigenum</italic> Yamamoto, Scient. Rep. Hyogo Univ. Agric., Agric. Biol. Ser. 2, 1: 69. 1955.</p></list-item><list-item><p id="P460"><italic>Penicillium ulaiense</italic> Hsieh, Su & Tzean, Trans. Mycol. Soc. R.O.C. 2: 161. 1987.</p></list-item><list-item><p id="P461"><italic>Penicillium vulpinum</italic> (Cooke & Massee) Seifert & Samson, Adv. Pen. Asp. Syst.: 144. 1985.</p></list-item></list></p></sec><sec id="S68"><title>Clade 18: section <italic>Roquefortorum</italic> (as “<italic>Roqueforti</italic>”) Frisvad & Samson, Stud. Mycol. 49: 16. 2004.</title><p id="P462">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P463">Type: <italic>Penicillium roqueforti</italic> Thom</p><p id="P464">Frisvad & Samson (<xref ref-type="bibr" rid="R36">2004</xref>) erected section <italic>Roqueforti</italic> for rapidly growing species forming strictly velutinous colonies. All species form terverticillate rough walled conidiophores and are able to grow at low pH values (<italic>e.g.</italic> on media containing 0.5 % acetic acid), at high alcohol concentrations and at elevated CO<sub>2</sub> levels. Members of this section appear to have a symbiotic relationship with lactic acid bacteria and certain acid-tolerant yeasts. Currently, four species are described in this section (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>, <xref ref-type="bibr" rid="R55">Houbraken <italic>et al</italic>. 2010a</xref>):</p><p id="P465"><list list-type="simple"><list-item><p id="P466"><italic>Penicillium carneum</italic> (Frisvad) Frisvad, Microbiology, UK, 142: 546. 1996.</p></list-item><list-item><p id="P467"><italic>Penicillium paneum</italic> Frisvad, Microbiology (UK) 142: 546. 1996.</p></list-item><list-item><p id="P468"><italic>Penicillium psychrosexualis</italic> Houbraken & Samson, IMA Fungus 1:174. 2010.</p></list-item><list-item><p id="P469"><italic>Penicillium roqueforti</italic> Thom, Bull. Bur. Anim. Ind. US Dept. Agric. 82: 35, 1906.</p></list-item></list></p></sec><sec id="S69"><title>Clade 19: section <italic>Chrysogena</italic> Frisvad & Samson, Stud. Mycol. 49: 17. 2004.</title><p id="P470">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P471">Type: <italic>Penicillium chrysogenum</italic> Thom</p><p id="P472">Members of the section <italic>Chrysogena</italic> are characterised by the formation of ter- and/or quarterverticillate, smooth walled conidiophores with relatively small phialides. Colonies have a velvety texture and species are tolerant to salt and the majority is capable to produce penicillin (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>). Four teleomorph species belong to section<italic> Chrysogena</italic>: <italic>P. sinaicum, P. egyptiacum, P. molle</italic> and <italic>P. kewense</italic> (<xref ref-type="fig" rid="F13">Fig. 13</xref>)<italic>. Penicillium egyptiacum</italic> was described as a holomorphic species (<xref ref-type="bibr" rid="R17">van Beyma 1933</xref>). Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) transferred the teleomorphic state to <italic>Eupenicillium</italic> (<italic>E. egyptiacum</italic>) and introduced a new name for the <italic>Penicillium</italic> morph (<italic>P. nilense</italic>). This name is not used here and <italic>P. egyptiacum</italic> is re-instated. There are several taxonomic problems concerning <italic>P. kewense</italic>. Brefeld (<xref ref-type="bibr" rid="R19">1874</xref>) was the first who described the formation of a teleomorph in <italic>Penicillium</italic>. He identified the studied species as “<italic>Penicillium crustaceum</italic> Fries, <italic>Penicillium glaucum</italic> Link”. It is, however, very questionable whether the strains studied by Brefeld truly represented the species described by Link and Fries (<xref ref-type="bibr" rid="R168">Stolk & Scott 1967</xref>). Stolk & Scott (<xref ref-type="bibr" rid="R168">1967</xref>) are followed here; they agreed that the fungus described by Smith (<xref ref-type="bibr" rid="R157">1961b</xref>) as <italic>Penicillium kewense</italic> resembles Brefeld's fungus. Based on the data of Samson <italic>et al</italic>. (<xref ref-type="bibr" rid="R147">2004</xref>), Houbraken <italic>et al.</italic> (<xref ref-type="bibr" rid="R57">2011a</xref>) and <xref ref-type="fig" rid="F13">Fig. 13</xref>, the following species are accommodated in section <italic>Chrysogena</italic>.</p><p id="P473"><list list-type="simple"><list-item><p id="P474"><italic>Penicillium aethiopicum</italic> Frisvad, Mycologia 81: 848. 1990.</p></list-item><list-item><p id="P475"><italic>Penicillium chrysogenum</italic> Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 58. 1910.</p></list-item><list-item><p id="P476"><italic>Penicillium confertum</italic> (Frisvad <italic>et al</italic>.) Frisvad, Mycologia 81: 852. 1990.</p></list-item><list-item><p id="P477"><italic>Penicillium dipodomyis</italic> (Frisvad, Filtenborg & Wicklow) Banke, Frisvad & Rosendahl, Int. Mod. Meth. Pen. Asp. Clas., 270. 2000.</p></list-item><list-item><p id="P478"><italic>Penicillium egyptiacum</italic> van Beyma, Zentralbl. Bakteriol., 2. Abt., 88: 137. 1933. (syn. <italic>P. nilense</italic>).</p></list-item><list-item><p id="P479"><italic>Penicillium flavigenum</italic> Frisvad & Samson, Mycol. Res. 101: 620. 1997.</p></list-item><list-item><p id="P480"><italic>Penicillium kewense</italic> Smith, Trans. Br. Mycol. Soc. 44: 42. 1961 (syn. <italic>E. crustaceum</italic>).</p></list-item><list-item><p id="P481"><italic>Penicillium molle</italic> Pitt, The Genus <italic>Penicillium</italic>: 148, 1980 [“1979”].</p></list-item><list-item><p id="P482"><italic>Penicillium mononematosum</italic> (Frisvad <italic>et al</italic>.) Frisvad, Mycologia 81:857. 1990.</p></list-item><list-item><p id="P483"><italic>Penicillium nalgiovense</italic> Laxa, Zentralbl. Bakteriol., 2. Abt., 86: 160. 1932.</p></list-item><list-item><p id="P484"><italic>Penicillium persicinum</italic> Wang, Zhou, Frisvad & Samson, Ant. van Leeuwenhoek 86: 177. 2004.</p></list-item><list-item><p id="P485"><italic>Penicillium rubens</italic> Biourge, Cellule 33: 265. 1923.</p></list-item><list-item><p id="P486"><italic>Penicillium sinaicum</italic> Udagawa & Ueda, Mycotaxon 14: 266. 1982.</p></list-item></list></p></sec><sec id="S70"><title>Clade 20: section <italic>Turbata</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563133&link_type=mb">MB563133</ext-link>.</title><p id="P487">Sectio in Penicillio subgen. Penicillo. Conidiophoris delicatis et symmetricis, biverticillatis; formatione acoris extroliti penicillici.</p><p id="P488">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P489">Type: <italic>Penicillium turbatum</italic> Westling</p><p id="P490">Section <italic>Turbata</italic> is phylogenetically closely related to section <italic>Paradoxa</italic>, and <italic>P. matriti, P. bovifimosum</italic> and <italic>P. turbatum</italic> are accommodated in this section. These species form rather delicate and symmetric biverticillate Penicillium conidiophores. Furthermore, penicillic acid is produced by all these species, and <italic>P. bovifimosum, P. turbatum</italic> and selected strains of <italic>P. matriti</italic> produce a fumagillin-like compound (<xref ref-type="bibr" rid="R180">Tuthill & Frisvad 2002</xref>).</p><p id="P491"><list list-type="simple"><list-item><p id="P492"><italic>Penicillium bovifimosum</italic> (Tuthill & Frisvad) Houbraken & Samson, Stud. Mycol. 70: 47. 2011 (this study).</p></list-item><list-item><p id="P493"><italic>Penicillium matriti</italic> Smith, Trans. Br. Mycol. Soc. 44: 44. 1961.</p></list-item><list-item><p id="P494"><italic>Penicillium turbatum</italic> Westling, Ark. Bot. 11: 128. 1911 (syn. <italic>E. baarnense, P. baarnense</italic>, this study).</p></list-item></list></p></sec><sec id="S71"><title>Clade 21: section <italic>Paradoxa</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563134&link_type=mb">MB563134</ext-link>.</title><p id="P495">Sectio in Penicillio subgen. Penicillo. Speciebus saepe cum conidiophoris typi Aspergillus et odore molesti efferenti.</p><p id="P496">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P497">Type: <italic>Aspergillus paradoxus</italic> Fennell & Raper</p><p id="P498"><italic>Aspergillus paradoxus, A. malodoratus, A. crystallinus</italic> and <italic>P. atramentosum</italic> form a well-supported clade (85 % bs, 1.00 pp). Phylogenetic and extrolite analysis shows that the first three species belong in <italic>Penicillium</italic> and will be transferred to this genus (R.A. Samson, unpubl. data). Besides a similar type of <italic>Aspergillus</italic> anamorph, these three species also produce a strong, unpleasant smell. <italic>Penicillium atramentosum</italic> is phylogenetically basal to these three species. This species is alkaliphilic and unpublished results show that this character is shared with <italic>A. paradoxus</italic>. More research is needed to determine whether <italic>A. malodoratus</italic> and <italic>A. crystallinus</italic> also share this feature.</p><p id="P499"><list list-type="simple"><list-item><p id="P500"><italic>Penicillium atramentosum</italic> Thom, Bull. Bur. Anim. Ind. US Dept. Agric. 118: 65. 1910.</p></list-item><list-item><p id="P501"><italic>Aspergillus crystallinus</italic> Kwon-Chung & Fennell, The Genus <italic>Aspergillus</italic>: 471. 1965.</p></list-item><list-item><p id="P502"><italic>Aspergillus malodoratus</italic> (Kwon-Chung & Fennell), The Genus <italic>Aspergillus</italic>: 468. 1965.</p></list-item><list-item><p id="P503"><italic>Aspergillus paradoxus</italic> Fennell & Raper, Mycologia 47: 69.</p></list-item></list></p></sec><sec id="S72"><title>Clade 22: section <italic>Brevicompacta</italic> Thom, The Penicillia: 289. 1930.</title><p id="P504"><list list-type="simple"><list-item><p id="P505">= section <italic>Coronata</italic> Pitt, The Genus <italic>Penicillium</italic>: 392, 1980.</p></list-item></list></p><p id="P506">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P507">Type: <italic>Penicillium brevicompactum</italic> Dierckx</p><p id="P508">Members of the section <italic>Brevicompacta</italic> are characterised by conidiophores with long and broad stipes. The conidial heads look superficially like <italic>Aspergillus</italic> heads in the stereomicroscope. Section <italic>Coronata</italic>, typified with <italic>P. olsonii</italic>, is placed here in synonymy. Recently, <italic>P. neocrassum</italic> and <italic>P. astrolobatum</italic> were described in this section (<xref ref-type="bibr" rid="R153">Serra & Peterson 2007</xref>) and partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) show that also <italic>P. tularense</italic> and <italic>P. fennelliae</italic> belong here. The production of the extrolites asperphenamate and the unknown metabolite O (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>) is shared by <italic>P. olsonii, P. brevicompactum</italic> and <italic>P. bialowiezense</italic>. More research in needed to determine whether these metabolites are also produced by the other members of section <italic>Brevicompacta</italic>. Based on literature (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>, <xref ref-type="bibr" rid="R114">Peterson 2004</xref>, <xref ref-type="bibr" rid="R153">Serra & Peterson 2007</xref>) and partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) the following species are accommodated in section <italic>Brevicompacta</italic>:</p><p id="P509"><list list-type="simple"><list-item><p id="P510"><italic>Penicillium astrolobatum</italic> Serra & Peterson, Mycologia 99: 80. 2007.</p></list-item><list-item><p id="P511"><italic>Penicillium bialowiezense</italic> Zaleski, Bull. Int. Acad. Pol. Sci. Lett., Sér. B, 1927: 462. 1927 (syn. <italic>P. biourgeianum</italic>).</p></list-item><list-item><p id="P512"><italic>Penicillium brevicompactum</italic> Dierckx, Ann. Soc. Scient. Brux. 25: 88. 1901.</p></list-item><list-item><p id="P513"><italic>Penicillium fennelliae</italic> Stolk, Ant. van Leeuwenhoek 35: 261. 1969.</p></list-item><list-item><p id="P514"><italic>Penicillium neocrassum</italic> Serra & Peterson, Mycologia 99: 81. 2007.</p></list-item><list-item><p id="P515"><italic>Penicillium olsonii</italic> Bainier & Sartory, Ann. Mycol. 10: 398. 1912.</p></list-item><list-item><p id="P516"><italic>Penicillium tularense</italic> Paden, Mycopathol. Mycol. Appl. 43: 264. 1971.</p></list-item></list></p></sec><sec id="S73"><title>Clade 23: section <italic>Ramosa</italic> (as “<italic>Ramosum</italic>”) Stolk & Samson, Adv. Pen. Asp. Syst.: 179. 1985.</title><p id="P517">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P518">Type: <italic>Penicillium lanosum</italic> Westling</p><p id="P519"><xref ref-type="fig" rid="F13">Figure 13</xref> shows that section <italic>Ramosa</italic> is not well resolved and members of this section are on a well-supported branch with section <italic>Brevicompacta</italic> members (100 % bs, 1.00 pp). We split this clade in two sections based on phenotypic characters and extrolite patterns. Members of the section <italic>Lanosa</italic> form biverticillate or terverticillate conidiophores with divergent rami (twice biverticillate), while members of sect. <italic>Brevicompacta</italic> have appressed branches. <italic>Penicillium jamesonlandense, P. lanosum, P. ribeum, P. raistrickii, P. soppii</italic> and <italic>P. swiecickii</italic> produce different combinations of cycloaspeptide, kojic acid and griseofulvin (<xref ref-type="bibr" rid="R34">Frisvad & Filtenborg 1990</xref>, <xref ref-type="bibr" rid="R35">Frisvad <italic>et al</italic>. 2006</xref>) and these extrolites are not been found in section <italic>Brevicompacta</italic> (<xref ref-type="bibr" rid="R36">Frisvad & Samson 2004</xref>). More research is needed to determine if the other members of this section also produce cycloaspeptide, kojic acid and/or griseofulvin.</p><p id="P520"><italic>Penicillium scabrosum</italic> is basal to the members of sections <italic>Brevicompacta</italic> and <italic>Ramosa</italic>. This species is tentatively accommodated in sect. <italic>Ramosa</italic> based on the formation of divaricate branches (<xref ref-type="bibr" rid="R37">Frisvad <italic>et al</italic>. 1990a</xref>). In contrast, cyclopenin, cyclopenol, viridicatin, penigequinolone A and B and fumagillin are produced by <italic>P. scabrosum</italic> and these extrolites are not detected in species belonging to sect. <italic>Ramosa</italic> (<xref ref-type="bibr" rid="R37">Frisvad <italic>et al</italic>. 1990a</xref>, <xref ref-type="bibr" rid="R77">Larsen <italic>et al</italic>. 1999</xref>). In the original description of <italic>P. virgatum</italic>, a relationship with <italic>P. daleae</italic> was suggested (<xref ref-type="bibr" rid="R74">Kwasna & Nirenberg 2005</xref>). However, these two species are unrelated and our partial <italic>RPB2</italic> data suggest <italic>P. virgatum</italic> is related to members of section <italic>Ramosa</italic> (<xref ref-type="fig" rid="F13">Fig. 13</xref>). Based on data presented in <xref ref-type="fig" rid="F13">Fig. 13</xref> and in Frisvad <italic>et al</italic>. (<xref ref-type="bibr" rid="R35">2006</xref>), the following species are placed in section <italic>Ramosa</italic>:</p><p id="P521"><list list-type="simple"><list-item><p id="P522"><italic>Penicillium jamesonlandense</italic> Frisvad & Overy, Int. J. Syst. Evol. Microbiol. 56: 1435. 2006.</p></list-item><list-item><p id="P523"><italic>Penicillium kojigenum</italic> Smith, Trans. Br. Mycol. Soc. 44: 43. 1961.</p></list-item><list-item><p id="P524"><italic>Penicillium lanosum</italic> Westling, Ark. Bot. 11: 97. 1911.</p></list-item><list-item><p id="P525"><italic>Penicillium raistrickii</italic> Smith, Trans. Br. Mycol. Soc.18: 90. 1933.</p></list-item><list-item><p id="P526"><italic>Penicillium ribeum</italic> Frisvad & Overy, Int. J. Syst. Evol. Microbiol. 56: 1436. 2006.</p></list-item><list-item><p id="P527"><italic>Penicillium sajarovii</italic> Quintanilla, Avances Nutr. Mejora Anim. Aliment. 22: 539. 1981.</p></list-item><list-item><p id="P528"><italic>Penicillium scabrosum</italic> Frisvad, Samson & Stolk, Persoonia 14: 177. 1990.</p></list-item><list-item><p id="P529"><italic>Penicillium simile</italic> Davolos, Pietrangeli, Persiani & Maggi, J. Syst. Evol. Microbiol., <italic>in press</italic>.</p></list-item><list-item><p id="P530"><italic>Penicillium soppii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 476. 1927.</p></list-item><list-item><p id="P531"><italic>Penicillium swiecickii</italic> Zaleski, Bull. Int. Acad. Pol. Sci. Lett., Sér. B 1927: 474. 1927.</p></list-item><list-item><p id="P532"><italic>Penicillium virgatum</italic> Nirenberg & Kwasna, Mycol. Res. 109: 977. 2005.</p></list-item></list></p></sec><sec id="S74"><title>Clade 24: section <italic>Canescentia</italic> Houbraken & Samson, sect. nov. MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB563135&link_type=mb">MB563135</ext-link>.</title><p id="P533">Sectio in Penicillio subgen. Penicillo. Structuris symmetricis biverticillatis, raro cum ramulis pluribus. Phialidibus simplicibus, brevibus (7–9 μm), cum collo brevi, interdum distincte attenuato.</p><p id="P534">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P535">Type: <italic>Penicillium canescens</italic> Sopp</p><p id="P536">Members of section <italic>Canescentia</italic> are soil-borne and are characterised by the formation of symmetrical biverticillate structures with infrequently an additional branch. Phialides are simple and short (7–9 μm) with a broadly cylindrical to slightly or more definitely swollen base and a short, occasionally more pronounced narrowed neck. This section has not been a subjected to a thorough phylogenetic study and unpublished sequence results show that several synonyms should be raised to species level. Partial <italic>RPB2</italic> data (<xref ref-type="fig" rid="F13">Fig. 13</xref>) shows that following species are placed in section <italic>Canescentia</italic>.</p><p id="P537"><list list-type="simple"><list-item><p id="P538"><italic>Penicillium canescens</italic> Sopp, Skr. Vidensk.-Selsk. Christiana, Math.-Naturvidensk. Kl. 11: 181. 1912.</p></list-item><list-item><p id="P539"><italic>Penicillium jensenii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 494. 1927.</p></list-item><list-item><p id="P540"><italic>Penicillium yarmokense</italic> Baghdadi, Nov. Sist. Niz. Rast. 5: 99. 1968.</p></list-item><list-item><p id="P541"><italic>Penicillium janczewskii</italic> Zaleski, Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat., 1927: 488. 1927.</p></list-item><list-item><p id="P542"><italic>Penicillium antarcticum</italic> Hocking & McRae, Polar Biology 21: 103. 1999.</p></list-item><list-item><p id="P543"><italic>Penicillium atrovenetum</italic> Smith, Trans. Br. Mycol. Soc. 39: 112. 1956.</p></list-item><list-item><p id="P544"><italic>Penicillium novae-zeelandiae</italic> van Beyma, Ant. van Leeuwenhoek 6: 275. 1940.</p></list-item><list-item><p id="P545"><italic>Penicillium coralligerum</italic> Nicot & Pionnat, Bull. Soc. Mycol. France 78: 245. 1963 [“1962”].</p></list-item></list></p></sec><sec id="S75"><title>Clade 25: section <italic>Eladia</italic> (Smith) Stolk & Samson, Adv. Pen. Asp. Syst.: 169. 1985.</title><p id="P546">In: <italic>Penicillium</italic> subgenus <italic>Penicillium</italic></p><p id="P547">Type: <italic>Penicillium sacculum</italic> Dale</p><p id="P548">The genus <italic>Eladia</italic> is synonymised with <italic>Penicillium</italic> and two species are placed here in section <italic>Eladia</italic>: <italic>P. sacculum</italic> and <italic>P. senticosum</italic> (<xref ref-type="fig" rid="F7">Fig. 7</xref>, clade 25 and <xref ref-type="fig" rid="F13">Fig. 13</xref>). <italic>Penicillium sacculum</italic> and <italic>P. senticosum</italic> grow rather well on MEA (and poorly on Czapek agar) and their colonies on MEA are velvety and dull-green, brownish-green or olive-brown coloured. Phialides are born irregularly on the stipes, subterminally as well as terminally, short, 4–7 μm, with a swollen base, and at the apex tapering abruptly into a short narrow neck. Conidia are distinctly ornamented (<xref ref-type="bibr" rid="R157">Smith 1961b</xref>, <xref ref-type="bibr" rid="R126">Pitt 1980</xref>, <xref ref-type="bibr" rid="R166">Stolk & Samson 1983</xref>, <xref ref-type="bibr" rid="R167">Stolk & Samson 1985</xref>). No type material could be obtained from <italic>Eladia pachyphialis</italic> and <italic>Eladia tibetensis</italic> and their taxonomic position remains uncertain. Based on their protologues, it is likely that these species belong to <italic>Penicillium</italic>.</p><p id="P549"><list list-type="simple"><list-item><p id="P550"><italic>Penicillium sacculum</italic> Dale <italic>apud</italic> Biourge, Cellule 33: 323. 1923.</p></list-item><list-item><p id="P551"><italic>Penicillium senticosum</italic> Scott, Mycopathol. Mycol. Appl. 36: 5. 1968.</p></list-item></list></p></sec></sec><sec id="S76"><title>Excluded and unclassified Penicillia</title><p id="P552">Over 250 <italic>Penicillium</italic> and <italic>Eupenicillium</italic> species are mentioned in the list of accepted <italic>Penicillium</italic> species (<xref ref-type="bibr" rid="R130">Pitt <italic>et al</italic>. 2000</xref>) and a fair amount of these do not belong to <italic>Penicillium s. str</italic>. The majority of these excluded species are currently classified in <italic>Talaromyces</italic> and an overview of species is given by Samson <italic>et al</italic>. (<xref ref-type="bibr" rid="R149">2011</xref>). Only a small number of species do not belong to either genus. These include <italic>P. arenicola, P. inflatum, P. kabunicum, P. lineatum, P. megasporum</italic> and <italic>P. moldavicum</italic>. <xref ref-type="fig" rid="F1">Figure 1</xref> shows that <italic>P. arenicola</italic> is closely related to <italic>Phialomyces</italic> (clade 6) and <italic>P. megasporum</italic> belongs to the clade 3 (<italic>Hamigera</italic>/<italic>Warcupiella</italic>). Both species should be transferred to other genera. Unpublished data (R.A. Samson) shows that <italic>P. inflatum</italic> belongs to <italic>Aspergillus</italic> and this species will be combined in that genus. <italic>Penicillium kabunicum</italic> and <italic>P. moldavicum</italic> are phylogenetically related and were included in the initial analyses of <italic>Trichocomaceae</italic>. Both species were together on a single branch and did not fit with any members of this family (J. Houbraken, unpubl. data). These two species belong to another (related) family and might represent a new genus. <italic>Penicillium lineatum</italic> was described as the anamorph of <italic>Hamigera striata</italic> (<xref ref-type="bibr" rid="R126">Pitt 1980</xref>). <italic>Hamigera striata</italic> is accommodated in clade 3 (<xref ref-type="fig" rid="F1">Fig. 1</xref>) and does therefore not belong to <italic>Penicillium s. str. Penicillium syriacum</italic> was included in the list of accepted names (<xref ref-type="bibr" rid="R130">Pitt <italic>et al</italic>. 2000</xref>), but the illustration and description of <italic>P. syriacum</italic> by Baghdadi (<xref ref-type="bibr" rid="R9">1968</xref>) and examination of ex-type material from ATCC, CBS and IMI indicated a mixed culture. This species is considered a <italic>nomen ambiguum</italic> (<xref ref-type="bibr" rid="R21">Christensen <italic>et al</italic>. 1999</xref>).</p><p id="P553">The phylogenetic position of <italic>P. resedanum</italic> needs further attention. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) and Ramírez (<xref ref-type="bibr" rid="R132">1982</xref>) placed <italic>P. resedanum</italic> in section <italic>Aspergilloides</italic> based on the formation of monoverticillate conidiophores. Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) already noted that this species form acerose phialides with weak growth on G25N, suggesting a relationship with <italic>Talaromyces</italic> (and subgenus <italic>Biverticillium</italic>). A BLAST search on GenBank with ITS sequences of NRRL 578<sup>T</sup> (AF033398) indicates a relationship with <italic>Talaromyces</italic>.</p><p id="P554"><italic>Penicillium griseolum</italic> is listed as a synonym of <italic>P. restrictum</italic> (<xref ref-type="bibr" rid="R130">Pitt <italic>et al.</italic> 2000</xref>). However, <xref ref-type="fig" rid="F7">Fig. 7</xref> shows that these species are phylogenetically unrelated. In our study, we did not find any species closely related to <italic>P. griseolum</italic> and this species might represent a separate section. We have chosen not to proceed with the description of this new section for this species until additional related species are described.</p><p id="P555"><list list-type="simple"><list-item><p id="P556"><italic>Penicillium arenicola</italic> Chalabuda, Bot. Mater. Otd. Sporov. Rast. 6: 162. 1950 (= clade 6, related to <italic>Phialomyces</italic>).</p></list-item><list-item><p id="P557"><italic>Penicillium inflatum</italic> Stolk & Malla, Persoonia 6: 197. 1971. (= <italic>Aspergillus inflatus</italic>, R.A. Samson, unpubl. data).</p></list-item><list-item><p id="P558"><italic>Penicillium kabunicum</italic> Baghdadi, Novosti Sist. Nizs. Rast.: 98. 1968 (unrelated to <italic>Penicillium</italic>, J. Houbraken, unpubl. data).</p></list-item><list-item><p id="P559"><italic>Penicillium lineatum</italic> Pitt, The Genus <italic>Penicillium</italic>: 485. 1980 [“1979”] (= <italic>Hamigera striata</italic>).</p></list-item><list-item><p id="P560"><italic>Penicillium megasporum</italic> Orpurt & Fennell, Mycologia 47: 233. 1955 (= clade 3, related to <italic>Hamigera</italic> and <italic>Warcupiella</italic>).</p></list-item><list-item><p id="P561"><italic>Penicillium moldavicum</italic> Milko & Beliakova, Novosti Sist. Nizs. Rast. 1967: 255. 1967 (unrelated to <italic>Penicillium</italic>, J. Houbraken, unpubl. data).</p></list-item><list-item><p id="P562"><italic>Penicillium syriacum</italic> Baghdadi, Novosti Sist. Nizs. Rast. 1968: 111. 1968 (<italic>nomen ambiguum,</italic> <xref ref-type="bibr" rid="R21">Christensen <italic>et al.</italic> 1999</xref>).</p></list-item></list></p></sec><sec id="S77"><title>Character analysis</title><p id="P563">The classification proposed in the monographs of Raper & Thom (<xref ref-type="bibr" rid="R135">1949</xref>), Pitt (<xref ref-type="bibr" rid="R126">1980</xref>) and Ramírez (<xref ref-type="bibr" rid="R132">1982</xref>) is not concordant with the new classification system proposed here. One of the most important characters in these monographs is the branching pattern of the <italic>Penicillium</italic> conidiophore. Our study shows that monoverticillate (Aspergilloid) conidiophores occur in various sections (<italic>e.g.</italic> clades 1, 2, 6, 8, 10, 12, 25). Sections <italic>Aspergilloides</italic> (clade 1) and <italic>Eladia</italic> (clade 25) comprise only strictly monoverticillate species, while mono- and biverticillate species are intermingled in the other clades. The occurrence of both structures in multiple phylogenetic clades (sections) indicates that reduction of the <italic>Penicillium</italic> conidiophore might have occurred various times. Most of the species belonging to section <italic>Citrina</italic> (clade 14) are symmetrically biverticillate and occasionally additional branches with the same branching pattern as the main axis (“double symmetrically biverticillate”) occurs. Species belonging to section <italic>Lanata-divaricata</italic> are mainly divaricate and the metulae are borne terminally, subterminally and in intercalary positions. Terverticillate conidiophores mainly occur in clades 15–18 and section <italic>Chrysogena</italic> (clade 19) comprises species with quarterverticillate condiophores. The monoverticillate species <italic>Penicillium sacculum</italic> and <italic>P. senticosum</italic> belong to clade 25. This clade is positioned in subgenus <italic>Penicillium</italic> and has therefore a unique branching pattern for this subgenus. Growth rates on agar media are also frequently used for classification. Some sections mainly comprise fast growing species (<italic>e.g.</italic> clades 1, 2, 11, 16, 18, 19, 25) while in other clades slow growing species predominate (<italic>e.g.</italic> clades 3, 6, 8, 9). The new proposed sectional classification will serve as a starting point to investigate phenotypic characters used for classification.</p></sec></sec></sec><sec id="S78"><title>TAXONOMIC IMPLICATIONS</title><p id="P564"><bold><italic>Penicillium asymmetricum</italic></bold> (Subramanian & Sudha) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561963&link_type=mb">MB561963</ext-link>.</p><p id="P565"><italic>Basionym</italic>: <italic>Thysanophora asymmetrica</italic> Subramanian & Sudha, Kavaka 12: 88. 1985.</p><p id="P566"><bold><italic>Penicillium bovifimosum</italic></bold> (Tuthill & Frisvad) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561957&link_type=mb">MB561957</ext-link>.</p><p id="P567"><italic>Basionym</italic>: <italic>Eupenicillium bovifimosum</italic> Tuthill & Frisvad, Mycologia 94: 241. 2002.</p><p id="P568"><bold><italic>Penicillium coniferophilum</italic></bold> Houbraken & Samson, <bold>nom. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561968&link_type=mb">MB561968</ext-link>.</p><p id="P569"><italic>Basionym</italic>: <italic>Thysanophora striatispora</italic> Barron & Cooke, Mycopathologia et Mycologia Applicata 40: 353. 1970, non <italic>Penicillium striatisporum</italic> Stolk, Ant. van Leeuwenhoek 35: 268. 1969.</p><p id="P570"><italic>Note</italic>: The name <italic>P. striatisporum</italic> is already occupied and therefore a new name is proposed.</p><p id="P571"><bold><italic>Penicillium glaucoalbidum</italic></bold> (Desmazières) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561965&link_type=mb">MB561965</ext-link>.</p><p id="P572"><italic>Basionym</italic>: <italic>Sclerotium glaucoalbidum</italic> Desmazières, Annales des Sciences Naturelles, Botanique 16: 329. 1851.</p><p id="P573"><list list-type="simple"><list-item><p id="P574">= <italic>Thysanophora glaucoalbida</italic> (Desm.) Morelet, Annales de la Société des Sciences Naturelles et Archéologie de Toulon et Var 20: 104. 1968.</p></list-item><list-item><p id="P575">= <italic>Thysanophora penicillioides</italic> (Roumeguère) Kendrick, Can. J. Bot. 39: 820. 1961.</p></list-item></list></p><p id="P576"><italic>Note</italic>: Virtually all of the published information relating to <italic>P. glaucoalbidum</italic> has used the binomial <italic>Thys. penicillioides.</italic> Iwamoto <italic>et al</italic>. (2005) aggregated sequence data of seven European and North American <italic>P. glaucoalbidum</italic> (as <italic>Thys. penicillioides</italic>) strains with Japanese strains. The strains formed nine lineages and according to phylogenetic species recognition by the concordance of genealogies, respective lineages correspond to phylogenetic species.</p><p id="P577"><bold><italic>Penicillium hennebertii</italic></bold> Houbraken & Samson, <bold>nom. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561964&link_type=mb">MB561964</ext-link>.</p><p id="P578"><italic>Basionym</italic>: <italic>Thysanophora canadensis</italic> Stolk & Hennebert, Persoonia 5: 189. 1968, non <italic>Penicillium canadense</italic> Smith, Trans. Br. mycol. Soc. 39: 113. 1956.</p><p id="P579"><italic>Note</italic>: A new name was sought for this species, as the species name “canadensis” is already occupied.</p><p id="P580"><bold><italic>Penicillium laeve</italic></bold> (K. Ando & Manoch) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561960&link_type=mb">MB561960</ext-link>.</p><p id="P581"><italic>Basionym</italic>: <italic>Torulomyces laevis</italic> K. Ando & Manoch, Mycoscience 39: 317. 1998.</p><p id="P582"><bold><italic>Penicillium longisporum</italic></bold> (Kendrick) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561966&link_type=mb">MB561966</ext-link>.</p><p id="P583"><italic>Basionym</italic>: <italic>Thysanophora longispora</italic> Kendrick, Can. J. Bot. 39: 826. 1961.</p><p id="P584"><bold><italic>Penicillium malachiteum</italic></bold> (Yaguchi & Udagawa) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561971&link_type=mb">MB561971</ext-link>.</p><p id="P585"><italic>Basionym</italic>: <italic>Chromocleista malachitea</italic> Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 102. 1993.</p><p id="P586"><list list-type="simple"><list-item><p id="P587">= <italic>Geosmithia malachitea</italic> Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 102. 1993.</p></list-item></list></p><p id="P588"><bold><italic>Penicillium melanostipe</italic></bold> Houbraken & Samson, <bold>nom. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561970&link_type=mb">MB561970</ext-link>.</p><p id="P589"><italic>Basionym</italic>: <italic>Thysanophora verrucosa</italic> Mercado, Gené & Guarro, Mycotaxon 67: 419. 1998, non <italic>Penicillium verrucosum</italic> Dierckx, Annales de la Société Scientifique de Bruxelles 25: 88. 1901.</p><p id="P590"><italic>Note</italic>: The name <italic>Penicillium verrucosus</italic> is already occupied and therefore the name melanostipe, which is referring to the pigmented stipe of this species, is proposed.</p><p id="P591"><bold><italic>Penicillium ovatum</italic></bold> (K. Ando & Nawawi) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561961&link_type=mb">MB561961</ext-link>.</p><p id="P592"><italic>Basionym</italic>: <italic>Torulomyces ovatus</italic> K. Ando & Nawawi, Mycoscience 39: 317. 1998.</p><p id="P593"><bold><italic>Penicillium parviverrucosum</italic></bold> (K. Ando & Pitt) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561962&link_type=mb">MB561962</ext-link>.</p><p id="P594"><italic>Basionym</italic>: <italic>Torulomyces parviverrucosus</italic> K. Ando & Pitt, Mycoscience 39: 317. 1998.</p><p id="P595"><bold><italic>Penicillium porphyreum</italic></bold> Houbraken & Samson, <bold>nom. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561959&link_type=mb">MB561959</ext-link>.</p><p id="P596"><italic>Basionym</italic>: <italic>Monocillium humicola</italic> Barron var. <italic>brunneum</italic> M. Christensen & Backus, Mycologia 56: 498. 1964, non <italic>Penicillium brunneum</italic> Udagawa, J. agric. Sci. Tokyo Nogyo Daigaku 5: 16. 1959.</p><p id="P597"><list list-type="simple"><list-item><p id="P598">= <italic>Torulomyces brunneus</italic> (M. Christensen & Backus) K. Ando, Mycoscience 39: 314. 1998.</p></list-item></list></p><p id="P599"><italic>Note</italic>: The name <italic>Penicillium brunneum</italic> is already occupied (<xref ref-type="bibr" rid="R181">Udagawa <italic>et al.</italic> 1959</xref>) and therefore the name <italic>P. porphyreum</italic> is proposed. The epithet porphyreum refers to the red-brown reverse of this species.</p><p id="P600"><bold><italic>Penicillium saturniforme</italic></bold> (Wang & Zhuang) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561958&link_type=mb">MB561958</ext-link>.</p><p id="P601"><italic>Basionym</italic>: <italic>Eupenicillium saturniforme</italic> Wang & Zhuang, Mycopathologia 167: 300. 2009.</p><p id="P602"><bold><italic>Penicillium taiwanense</italic></bold> (Matsushima) Houbraken & Samson, <bold>comb. nov.</bold> MycoBank <ext-link ext-link-type="uri" xlink:href="http://www.studiesinmycology.org/cgi/external_ref?access_num=MB561969&link_type=mb">MB561969</ext-link>.</p><p id="P603"><italic>Basionym</italic>: <italic>Phialomyces taiwanensis</italic> Matsushima, Matsushima Mycological Memoirs 4: 12. 1985.</p><p id="P604"><list list-type="simple"><list-item><p id="P605">= <italic>Thysanophora taiwanensis</italic> (Matsush.) Mercado, Gené & Guarro, Mycotaxon 67: 421. 1998.</p></list-item></list></p><p id="P606"><italic>Note</italic>: This species was originally described as <italic>Phialomyces taiwanensis</italic>. Based on micro-morphological features, Mercado-Sierra <italic>et al</italic>. (<xref ref-type="bibr" rid="R97">1998</xref>) transferred this species to <italic>Thysanophora taiwanensis</italic>.</p></sec><sec sec-type="supplementary-material"><title>Supplemental Information</title><supplementary-material content-type="local-data" id="SI1"><label>Table S1.</label><caption><p>Penicillium strains used in the study of the infrageneric classification (addition to those mentioned in Table 1).</p></caption><media xlink:href="51_supp_info.pdf" xlink:type="simple" id="d32e16540" position="anchor" mimetype="application" mime-subtype="pdf"></media></supplementary-material></sec></body><back><ack><p id="P607">Neriman Yilmaz and Barbara Favie are thanked for testing various primer pairs and generating sequences. Uwe Braun is thanked for providing the Latin diagnosis and advice on nomenclature issues. Jens Frisvad is acknowledged for providing various isolates and the reviewers for their useful suggestions. 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